recovered material

1.2.3.2. Taxonomy-Dependent Principles 1.2.2.2. Phenetics Unlike cladistics, phenetics and traditional systematics (or, in its refined form, phyletics) proceed from the integrity of living systems and, accordingly, from the discreteness of biodiversity, and are engaged in searching for hiata. Therefore, the fundamental concept must include a process that generates such discreteness. Initially, creationism readily explained discreteness; today the epigenetic theory of evolution appears to be the only realistic contender for this role (see above). Of the three competing approaches, phenetics is the most straightforward in its aims and methods. Since the discreteness of biodiversity is described in terms of similarity rather than kinship, pheneticists maintain that the systematist's task should be formulated in the same terms — namely, that the system should be explicitly constructed in terms of similarity. Subjectivity can be avoided here through the calculus of similarity. The methods of calculation are collectively termed taxometry, or more frequently, though less felicitously, numerical taxonomy. Unfortunately, these methods were not developed to the proclaimed level — in my view not because that level is unattainable, but because phenetics was displaced by cladistics before it had completed its useful work. Indeed, the methods of taxometry are numerous and diverse; choosing a method adequate to the task is not straightforward and at the current level is scarcely possible without a considerable degree of subjectivity. That said, this is a problem for every approach, and cladistics is no exception (see, for example, Mickevich, 1978): identifying the optimal domain of application of a specific method, its strengths and weaknesses, can only be achieved at considerable cost. Before yielding the field to cladistics, phenetics had advanced substantially — though not sufficiently — in this direction. However, the fundamental problem of phenetics, in my view, is different. The similarity with which it works is not quite the similarity that the system of organisms requires. Pheneticists work with studied characters, the number of which should be reasonably large, but no more than that. The system, in order to optimally fulfil its functions, must rest on similarities and differences across all characters — studied and unstudied alike, including those that may never be studied. Of course, such similarity is not accessible to direct investigation, but it is important enough to warrant an indirect estimate. This is precisely the goal that phyletics pursues. 1.2.2.3. Phyletics Phyletics, like cladistics, proceeds from the assumption that the organisation of living beings is determined not only, and not even primarily, by the conditions of their existence, but by their history. Similarity of history (consanguinity) largely predetermines similarity of organisation as well. The difference lies in the fact that the methodology of cladistics ignores everything except kinship, phenetics ignores kinship, while phyletics attempts to account for both kinship and similarity. For a phyletic taxon to simultaneously reflect both similarity and genealogical relationships, it must, first, be phenetically (by similarity) maximally homogeneous and maximally distinct from other taxa and, second, be monophyletic. Monophyly is understood in a broader sense than in cladistics — that is, encompassing holophyly (monophyly in the cladistic sense; see Ashlock, 1971) and paraphyly. The phylogenetic qualification of taxa is determined by lines of descent crossing the taxon's boundaries. If a single line (one root) crosses its lower boundary, the taxon is termed monophyletic (in the broad sense); if more lines cross it, the taxon is polyphyletic. A more precise formulation, however, is the following: a taxon should be considered monophyletic as long as and insofar as there is no reliable evidence of its polyphyly (the presumption of taxon monophyly). Cladists redefined the concept of monophyly with reference not only to the origin (roots) of a taxon but also to its subsequent fate. If the upper boundary of a monophyletic (in our sense) taxon is not crossed by phyletic lines — that is, the taxon has no obvious descendants — we call it holophyletic, and cladists call it monophyletic. If, however, its upper boundary is crossed (descendants exist), both phyleticists and cladists call it a paraphyletic taxon. Cladists call a holophyletic taxon monophyletic because, from their point of view, a paraphyletic taxon has no more right to exist than a polyphyletic one. They have no need for a term combining holophyletic and paraphyletic taxa: such a combination is meaningless to them. In phyletics, however, such a term is absolutely necessary, since both forms of taxon (holo- and paraphyletic) are recognised and treated as equally valid. This phyletic understanding was the original sense of the term monophyly, and it is adopted here as well. There is an even broader understanding of monophyly — as the origin of a natural taxon by multiple roots from a common ancestor within the ancestral taxon (pluriphyly, pachyphyly, broad monophyly). This is how the origin of mammals, reptiles, and many other large groups is currently envisaged, hence the terms mammalisation, arthropodisation, angiospermisation, and the like. Broad monophyly not only falls under the definition of polyphyly in the cladistic formulation, but in general exhausts the real content of that term. By definition, what lies beyond broad monophyly is only the multiple origin of a taxon whose naturalness is denied (we will encounter these in Section 1.2.3.3). But for an artificial taxon, the multiplicity of its origin is trivial, and it is unlikely that anyone would think to discuss this separately. Maintaining a distinct term for such a case seems superfluous to me; accordingly, I will use polyphyly for all cases of multiple origin of a taxon, as formulated above. Thus it turns out that many (if not all) well-studied taxa are polyphyletic in the sense indicated. At the same time it is difficult to doubt the naturalness of reptiles, mammals, arthropods, angiosperms, and many other taxa for which origin from more than one root from an ancestral taxon has been demonstrated more or less convincingly. Thus polyphyly may prove to be the norm in taxonomy. And yet I maintain that it is methodologically more correct to proceed from the presumption of monophyly. I see two reasons for this: first, at the current very low level of knowledge of most taxa, the discovery of multiple origins for a particular group is more likely to falsify the hypothesis of its naturalness than the presumption of monophyly. If the group is genuinely natural, additional investigation provoked by the discovery of polyphyly (see below) will only confirm its naturalness and in the end will compel recognition that the presumption of monophyly is inapplicable in that particular case. There is another reason why abandoning the presumption of monophyly (in the sense adopted here, excluding broad monophyly) seems methodologically hazardous. This would mean renouncing the use of phylogenesis to control the naturalness of a taxon (see below) and, consequently, shifting to the position of phenetics. Such a step seems at the very least premature. Let us return to the phyletic procedure. A phyletic taxon was defined as a monophyletic continuum (Ponomarenko, Rasnitsyn, 1971). The definition of monophyly has been given above; the continuum is understood as an unbroken chain — branching or otherwise — constructed from subordinate taxa (monophyletic continua of smaller scope) such that each is phenetically closer to any of its immediate neighbours in the chain than to members of any other taxon (Fig. 6). Thus the continuum is separated by a hiatus (a break in continuity of similarity) from all other continua. [IMG_1] Fig. 6. Phenetic structure of lower Hymenoptera at the family level, demonstrating continua at several hierarchical levels (from Rasnitsyn, 1972b). Lines represent levels of similarity: a single line indicates the lowest level depicted, a triple line the highest. Abbreviations: Ar — Argidae, Ax — Anaxyelidae, Bl — Blasticotomidae, Cb — Cimbicidae, Cp — Cephidae, Dp — Diprionidae, Gs — Gidasiricidae, Md — Megalodontesidae, Mm — Myrmiciisae, Or — Orussidae, Pg — Pergidae, Pl — Pamphiliidae, Po — Paroryssidae, Pp — Parapamphiliidae (now in Sepulcidae), Sc — Sepulcidae, Sr — Siricidae, Tt — Tenthredinidae, Xd — Xyelydidae, Xl — Xyelidae, Xt — Xyelotomidae, Xy — Xiphydriidae. Building a system based on two different criteria (in this case, kinship and similarity) carries the risk of arbitrariness in their application — unless the domains of competence of the two criteria are clearly delimited. Cladistics and phenetics solve the problem in the simplest possible way — by renouncing the use of one or the other criterion: cladistics ignores similarity, phenetics ignores kinship. Phyletics acts more subtly: it uses both criteria, but differently. Similarity is used as the criterion that defines the system; kinship is used as the criterion that controls the quality of the system. A taxon is delineated by tracing hiata (breaks in similarity), and then tested for unity of origin. Thus phyletics requires that the members of its taxon be sufficiently similar to satisfy the criterion of the continuum, and that they show no signs of polyphyly (origin from more than one line of ancestors). If the continuum proves polyphyletic, the criteria come into conflict. In this case the systematist will hardly discard all the work done and start over. His natural reaction will be to attempt falsification of this result through additional study of the relevant group. The attempt usually succeeds in one sense or another. For example, a broader analysis may reveal that the similarity of polyphyletic groups is superficial. The order Neuroptera long included groups whose close kinship was subsequently not confirmed. Thus Sharp (1910) still grouped under this name, in addition to neuropteroids in the modern sense, also the embiids (Embioptera), termites (Isoptera), booklice (Psocoptera), biting lice (Mallophaga), dragonflies (Odonata), stoneflies (Plecoptera), mayflies (Ephemeroptera), scorpionflies (Mecoptera), and caddisflies (Trichoptera). A similar fate befell the mammalian order Pachydermata: the taxon was used in the eighteenth and nineteenth centuries to unite elephants, tapirs, rhinoceroses, hippopotamuses, and pigs, but was subsequently rejected in favour of more natural (monophyletic) orders. In other cases it proved possible to falsify the hypothesis of polyphyly, as with the lagomorphs. They were at one time separated into a distinct order Lagomorpha, supposedly only convergently similar to the rodents (Gidley, 1912). Although the order status of lagomorphs was retained, by modern understanding they form a monophyletic group Glires together with rodents (Wilson, 1989). It is quite possible that both hypotheses (of similarity and of polyphyly) will prove equally plausible. Most likely this will mean that our attempt was insufficient and we need to try again to analyse the situation and resolve the conflict. But it cannot be excluded that we have indeed arrived at the real multiple-rootedness of a natural taxon, which we have already mentioned repeatedly. I have no recipe for distinguishing one situation from the other, but in any case it is better to once again attempt to analyse the system and phylogeny of the group on broader material. There are also entirely obvious cases where traditional systematics retains unquestionably polyphyletic taxa that cannot be replaced without significant loss by monophyletic ones. I have in mind the various parataxon concepts in fields such as palaeontology, parasitology, mycology, and also many prokaryote taxa (for more detail see Section 1.2.3). The existence of parataxons once again confirms that traditional systematics is ultimately oriented toward similarity, and employs kinship merely as a means of indirectly estimating total similarity (including similarity by characters not yet studied) that is inaccessible to direct assessment. Consequently, kinship is used here in approximately the same way as, in the Hennigian version of cladistics, apomorphy is used to establish divergences, and in the Bremer version divergences serve to determine the sequence of apomorphies. Thus all the approaches considered are equally consistent (monistic, non-eclectic): Hennigian cladistics is ultimately oriented toward the sequence of divergence events, Bremerian cladistics toward the sequence of apomorphies, phenetics toward existing (directly accessible) similarity, and phyletics toward total similarity (inaccessible to direct observation, but indirectly estimated through kinship). We may now return to the question of the comparative merits and shortcomings of cladistics and phyletics. In my view, the most important merit of phyletics is its ultimate reliance on similarity — that is, on characters as such. It can be content with similarity in available characters when any reasonably reliable evidence of kinship is absent, and will then be just as empirical as phenetics — in fact, it will simply be phenetics. But it will also employ all (and only!) sufficiently reliable phylogenetic data to enhance the reliability of the system it constructs, without making it excessively hypothetical. In contrast, the cladistic system, both directly and ultimately, rests on kinship — that is, on the interpretation of characters (on assumptions regarding the evolutionary sequence of character states). As a result, a cladistic system is considerably more hypothetical than a phyletic one. Another aspect of what has been said is that phyletics is "omnivorous" whereas cladistics is "fastidious." Cladistics places far more stringent demands on the characters it employs than does phyletics (or phenetics), because characters that cannot be reliably polarised are useless. This, admittedly, applies only to Hennigian cladistics (manual, or "pen-and-paper cladistics"). In computer cladistics any characters may be used, but if the program cannot confidently reconstruct the sequence of changes for a given character (and it can hardly do this much better than a good systematist), that character will be superfluous in the calculations. Moreover, characters that work well in one part of the study group but show numerous parallel changes (homoplasies) in other parts are, as we have seen, devalued and given very little weight — even where they are in fact phylogenetically informative. Less well-studied taxa — for example, extinct ones — are characterised by fewer characters and therefore frequently fail to find an exact position in computer cladograms, even when they retain phylogenetically important characters. As a result, cladists are compelled to declare as parataxons ("plesiomorphs") many groups that are treated as normal by phyletics (orthataxa) but are insufficiently known for their phylogenetic relationships with better-studied taxa to be precisely characterised. Naturally, this reduces the taxonomic efficiency of cladistics. A phyletic system, accounting for both similarity and kinship, is more informative than others. J.S. Farris (Farris, 1979) objects that a cladistic system in its most complete form indicates the origin of all characters and is therefore more informative than a phenetic (and, by extension, phyletic) system. This conclusion is not unequivocal, since exhaustive completeness of a system is unattainable. Indeed, the more information we accumulate about the polarisation of known characters, the more new characters are discovered whose evolutionary history is as yet unknown and which therefore cannot be used in "manual" cladistic analysis. As a result one is forced to work with an incomplete set of characters compared with phyletics, and in this sense the system conveys less information. Computer cladistics can use all available characters, but it performs the polarisation of transformation series automatically, based simply on the distribution of character states (by the outgroup method) — that is, from very limited information. All other knowledge about the history of characters, widely used by phyletics and manual cladistics, is ignored here, so the overall information loss is large. In practice, each of the taxonomic methods under comparison uses only a portion of the available information, and these portions do not overlap very strongly. Computer cladistics covers a vast field of data but in a highly selective and superficial manner. Phenetics acts similarly but without such selectivity. Manual cladistics uses a limited "elite" body of material selected from an unlimited corpus. Phyletics also "sieves" an unlimited volume of raw data, but is freer than manual cladistics in sorting it and freer than phenetics and computer cladistics in its interpretation. It is therefore capable of extracting more extensive and varied information than other approaches. There is also another aspect to the problem. It is evident from the above that information resides not in the system itself, but in its description or, more precisely, in what underlies it. The system as such contains only very limited information about the topology of the corresponding dendrogram and the length of its internodes interpreted in taxonomic ranks. Thus a symmetric dichotomous dendrogram of four terminal taxa, even without additional information about their ranks, contains three bits of information and no more. Detailing the ranking of taxa enriches the informational content of the system, and in this respect the most information-rich are Hennig's original systems with their incredible number of numbered ranks exhaustively reflecting the supposed sequence of divergence events. However, this approach was found impractical and was effectively abandoned in favour of the traditional Linnaean rank hierarchy (Wiley, 1979). This form of ranking contains limited cladistic information and uses it arbitrarily (the assignment of a rank to one of many successive divergences is arbitrary), thereby making post-Hennigian cladistics eclectic. Artificial rules such as Nelson's phyletic sequencing (Nelson, 1973b; Cracraft, 1974) are of little help. We can agree that a pectinate cladogram should be transformed into a group of taxa of one rank, listed in order of their divergence beginning with the first to branch off — but in the last pair the order will inevitably be arbitrary. Nor will we be able to distinguish such a list from another constructed arbitrarily or based on a more complex cladogram that is not strictly pectinate and/or includes a polytomy. It is asserted that phenetics and phyletics, unlike cladistics, employ taxa characterised by the absence of a character (Platnick, 1979). This refers to paraphyletic taxa characterised by the absence of apomorphies. Above, however, it was shown that an apomorphy is not a character or a character state. An apomorphy is a hypothesis about the history of a character, so the absence of an apomorphy and the absence of a character are different things. Lizards lack the apomorphies that separate them from snakes, but they possess a character — legs. Also deserving discussion is the problem of symbiotic and hybrid taxa, important if only because they are too common to be dismissed as exotic. The first category includes no less a taxon than the eukaryotes themselves, and hybrids are very characteristic, for example, of many plant taxa. According to N.N. Tzvelev's review (1993), in the grass family (Poaceae) the tribe Triticeae includes 500 species, of which 300 are karyotypically clearly intergeneric hybrids, which in fact constitute the largest genera. The same is claimed to be true of many other higher plant taxa. Symbiotic and hybrid taxa are polyphyletic (or constitute part of a polyphyletic taxon) by definition, since their boundary (or the boundary of the senior taxon) is crossed by more than one ancestral line, which categorically violates the principles of cladistics. Nevertheless, such taxa cannot in principle be partitioned into holophyletic components and must therefore be excluded from the competence of cladistics — a conclusion with which few will agree. There are other considerations bearing on the comparison of competing taxonomic concepts. However, what has been said is sufficient to conclude that each of them has its own merits and shortcomings, which, moreover, do not always balance one another. Cladistics reduces the system of a group to its genealogical history, phenetics to the characters of similarity available for analysis, while traditional systematics (reflected in phyletics) attempts to use both, and not arbitrarily but in an ordered way. If any reasonably reliable evidence of kinship is absent, traditional systematics will be satisfied with mere similarity in available characters — and then it will simply be phenetics. But when possible, it will also employ all available phylogenetic data to enhance the reliability of the system it constructs. Phenetics is strictly empirical, while cladistics, by contrast, is highly hypothetical, since both directly and ultimately it relies on kinship — that is, on the interpretation of similarity. Phyletics, on the other hand, is free to occupy any position in this spectrum depending on the properties of the material available in each particular case. Phyletics is "omnivorous" and can work with any available material, employing when necessary any phenetic and cladistic methods. Cladistics, by contrast, is "fastidious" and places far more stringent demands on the characters it employs. That is why a cladistic system so often proves to be either unstable — when minimal changes in computational parameters substantially alter the form of the cladogram — or uninteresting ("unresolved," with the relationships of many taxa left undetermined). For the same reason, cladists are compelled to declare as parataxons ("plesiomorphs") many groups that are perfectly normal for phyletics.

1.2.1.3. Computer Cladistics 1.2.2. TAXONOMY 1.2.2.1. Cladistics A.P. Rasnitsyn. Theoretical Foundations of Evolutionary Biology // V.V. Zherikhin, A.G. Ponomarenko, A.P. Rasnitsyn. Introduction to Palaeoentomology. Moscow: KMK. 2008. 371 pp. 1.2.2.2. Phenetics 1.2.2.3. Phyletics 1.2.2.3. Phyletics (continued) 1.2.3. TAXONOMIC NOMENCLATURE 1.2.3.1. Taxonomy-Independent Principles 1.2.3.2. Taxonomy-Dependent Principles 1.2.2.2. Phenetics 1.2.2.3. Phyletics 1.2.2.2. Phenetics 1.2.2.3. Phyletics A.P. Rasnitsyn. Theoretical Foundations of Evolutionary Biology // V.V. Zherikhin, A.G. Ponomarenko, A.P. Rasnitsyn. Introduction to Palaeoentomology. Moscow: KMK. 2008. 371 pp. 1.2.2.3. Phyletics (continued) 1.2.3. TAXONOMIC NOMENCLATURE 1.2.3.1. Taxonomy-Independent Principles 1.2.3.2. Taxonomy-Dependent Principles 1.2.3.2. Taxonomy-Dependent Principles (continued) 1.2.3.3. Taxa Violating Principles of Nomenclature Nevertheless, cladistics is the dominant school of thought in the West, and the same tendency is evident in our country. It is simply easier. The methodology is clear, there is less room for intuitive evaluative functions — such as assessing the level of similarities and differences — and there are no separate tasks of constructing the system and the phylogeny. The advantages are particularly great in computer (parsimony) cladistics. Here only a single stage remains at which one must actually think: the delimitation of the group for analysis and the search for and selection of characters for constructing the cladogram. Cladists, however, do not trouble themselves with an analysis of the methodology of this most crucial part of the work. Beyond that, everything is simple: compile a matrix, enter it into a complex but thoroughly described and therefore fully user-accessible computer program, launch a powerful computer, and wait for the calculations to finish. If the cladogram is unsatisfactory, one can change the computational parameters and let the computer calculate again. Eventually it will produce an acceptable result, at least minimally consistent with common sense (i.e., with a priori expectations). In fact, cladistics is a striking example of deductive rather than hypothetico-deductive reasoning. Indeed, cladistics employs as a postulate the highly plausible assertion that a common phylogenetic past determines the similarity of related groups. From this, a conclusion is drawn without further verification: that the system of organisms must be strictly isomorphic to their genealogy as explicated in the cladogram. In doing so, no serious attention is paid to evidence that at least some of the most important postulates of cladistics lack biological justification, and that the results of their consistent application frequently contradict common sense and the results obtained by other methods, including the palaeontological and molecular ones (see above). There is no biological reason for divergence to proceed only in the form of strict dichotomy with the obligatory disappearance of the ancestral species at the moment of divergence — on the contrary, very reliable observations on fossil organisms (e.g., Nevesskaya et al., 1986; Nevesskaya et al., 2001) indicate that an ancestral species may give rise to new daughter species over millions of years. Equally, genealogical closeness can effectively reflect relationships along other organismal properties only if the evolutionary process is substantially uniform, but — as has already been shown — palaeontology abundantly furnishes compelling evidence of extreme irregularity in the evolutionary process. It follows that, all things considered, it is better to engage in traditional systematics, but to clearly understand its foundations, criteria, and goals as formulated by phyletics, and to employ the methods of cladistics and phenetics only where they can offer genuine assistance. 1.2.3. TAXONOMIC NOMENCLATURE Nomenclature is the body of conventions governing how names of taxa are created and used. It is usually perceived as an area of elaborate detail and apparently self-evident basic principles. This is incorrect, first, because not all of the operative principles of biological nomenclature have yet been consciously formulated in explicit terms. Some are mentioned in the codes (see ICZN, 1988, 2000; ICBN, 2000) but not as basic principles, and many principles are simply absent from them. Second, some of the basic principles are not pure conventions but reflect specific taxonomic concepts or represent corollaries of specific evolutionary hypotheses. Linnaeus's principle of binominal nomenclature is perfectly conventional and therefore easily compatible with any taxonomic concept that recognises a hierarchy of genera and species. The principle of the type is another matter, as discussed in detail below. Distinguishing between these two types of principles seems useful; hereafter they will be termed taxonomy-independent and taxonomy-dependent principles respectively (Table 4). Table 4. Principles of nomenclature Taxonomy-independent 1 non-interference in taxonomic decisions beyond the constraints imposed by taxonomy-dependent principles 2 limited scope of application 3 homonymy 4 priority 5 standardised, rank-specific names 6 supreme authority of the International Commission on Nomenclature Taxonomy-dependent 7 type (no reasonable grounds for rejection) 8 hierarchy (no reasonable grounds for rejection) 9 synonymy (application to parataxons limited to the framework of the respective partial system) 1.2.3.1. Taxonomy-Independent Principles Taxonomy-independent principles are generally simple, require little comment, and are considered first here. 1. The current International Code of Zoological Nomenclature states that it "refrains from interfering with taxonomic judgements that are not subject to regulation" (ICZN, 2000: 24). This assertion is not entirely accurate because of the existence of taxonomy-dependent principles that preclude certain taxonomic decisions. The formulation therefore requires the addition of the words "beyond the constraints imposed by the Code." 2. In zoology, the domain of application of the principles of nomenclature is restricted to suprapopulation groups from subspecies to superfamily (for details see ICZN, 2000, Art. 1.2). The exclusion of higher taxa from the scope of the code appears to be an error (Rasnitsyn, 1982, 1989, 1991, 2002b). My reasoning can be summarised briefly as follows. The standardisation of name formation and use ensured by the codes has demonstrated through long practice its beneficial effect on all of biology. The refusal to extend it to higher taxa is unlikely to be connected to any deep-seated peculiarity of those taxa. No serious indication of such peculiarity is known, and the numerous attempts to standardise the untypified names of higher taxa (the ending -ptera in insect order names, -formes in bird order names, -ida in many names of extinct orders of various invertebrate groups) remove all doubt as to the necessity of such unification. The point, of course, is not unification but the typification of names of higher taxa. Typification (more precisely, through-typification) is the procedure ensuring a firm link between each name of each taxon and a specific subordinate taxon within it, top-down through the entire hierarchy down to the type specimen of the species (or subspecies). What are typified are names, not the taxa themselves: when we speak of the holotype of a species or the type species of a genus, we mean the type of the name (nomen). The principle of the type, which corresponds to the very nature of the taxon (see below), is the principal achievement of biological nomenclature: it ensures stability of use and orderly change of the name of a taxon under the most varied changes in its composition (boundaries) and interpretation (change of rank, diagnostic characters, etc.). The first attempts to typify higher taxa date back more than two centuries (Laicharting, 1781), and understanding of its necessity is also growing. This view prevailed in botany (ICBN, 1985) and finds some support in zoology (Rodendorf, 1977; Starobogatov, 1991; Klyuge, 2000). The point is that an untypified name is referred to a taxon as a whole and is practical only as long as the interpretation of the taxon itself remains stable. Otherwise, problems arise with the application of the name, resulting in confusion and renaming. However, the introduction of typified names for higher taxa and their consequent renaming affects the interests of numerous users who have little interest in the problems of high-level systematics but are alarmed by the prospect of learning new words. For those working with higher taxa the situation proves difficult: taxonomic and phylogenetic errors and imprecisions become entrenched in untypified names and accumulate. The longer this continues, the more difficult it will be to transition to typification of higher taxa. But such a transition will inevitably occur, and posterity will not thank us for our selfish conservatism. 3. Identical names for different taxa should not be used as valid names (ICZN, 2000, Art. 52). The domain of application of the principle of homonymy is arbitrarily limited so as not to affect cases of homonymy: first, between species names in different genera; second, between plant and animal taxa; and third, with and between taxa of rank above superfamily. 4. The choice between competing names (synonyms or homonyms) must be made in favour of the name proposed earlier (principle of priority; ICZN, 2000, Art. 23). 5. The principle of standardised and rank-specific form of taxon names. This principle provides the basis for numerous rules concerning the language, grammar, and syntax of names, and includes the principle of binominal nomenclature (ICZN, 2000, Arts. 4-6, 11, 25-34). 6. The last of the taxonomy-independent principles is the principle of the supremacy of the International Commission on Nomenclature, which may make decisions in violation of any provision of the Code other than those concerning itself (ICZN, 2000, Art. 78). 1.2.3.2. Taxonomy-Dependent Principles There are three taxonomy-dependent principles: 7. The principle of the type. This is the most important of the taxonomy-dependent principles, and it well illustrates the constraints that a taxonomic concept can impose on nomenclature. In the 1988 Code this principle was formulated as follows: "The nomenclatural type is the objective standard by means of which the application of a name is determined independently of possible changes in the limits of the taxon" (ICZN, 1988, Art. 61a; the formulation of ICZN 2000, Art. 61.1, seems less felicitous: "The fixation of the nomenclatural type of a nominal taxon provides the objective standard for the application of the name of that taxon"). In other words, a taxon cannot be introduced into the system except by ultimate reference to the type that bears its name. To appreciate the significance of this assertion, let us consider the alternatives. The first that comes to mind is the possibility of introducing a taxon into the system by reference to its characters. This means that the characters of the taxon perform a defining rather than merely a diagnostic function, and the taxon itself corresponds to the concept of a "class" (Ghiselin, 1974, 1987). There are various ways of introducing a taxon into the system in this manner, the simplest being a combinatorial system — that is, a multidimensional matrix in which each cell corresponds to a specific combination of characters and constitutes the place of the taxon possessing that combination. A version of such a matrix is the polytomous key sometimes used by systematists — a rectangular matrix with rows representing taxa and columns corresponding to characters, so that each cell reflects the state of one specific character in a given taxon. This form of system is indeed simple and sometimes very useful, but does not correspond to the requirements of a general system of organisms. It lacks flexibility: we cannot correct it locally — for example, to better align it with the characters of a specific taxon — since the introduction of a new character or the redefinition or removal of an old one requires revision of the characteristics of all taxa. Therefore, even as a means of identification, the combinatorial system is rarely used. More flexible is a system in which taxa are defined by ranked characters. In an ordinary dichotomous key, the first character used has the highest rank, since it divides the group under determination into the largest subgroups. Characters introduced subsequently provide further subdivision and have lower rank. Such a system is indeed convenient, though not as good as a general system of organisms, owing to the obviously arbitrary ranking of characters. Nevertheless, in a somewhat modified form (with a small number of high-rank characters sufficient for constructing the system) this approach was very popular among systematists. In Linnaeus (Linnaeus, 1751), for example, the characters of plant fructification played a closely analogous role. This approach was formulated in its most explicit form by A.A. Lyubishchev (1923, 1966). Lyubishchev called for the search for a few highest-rank characters (parameters) determining the distribution of all other characters — just as nuclear charge determines the properties of atoms and, accordingly, the position of elements in the periodic table. The result was expected to be the construction of a parametric system of organisms that would allow prediction of all important properties of the corresponding taxa from these key characters (parameters). The task Lyubishchev set seemed hopeless, and he himself was unable to solve it. Nevertheless, a solution of some kind was found — but in quite a different place from where he sought it. By an irony of fate, this happened in the field that Lyubishchev himself, as a convinced anti-selectionist, regarded as false. The solution proved to be the cladistic system, with its central assertion that the characters of an organism and, accordingly, its position in the system are best determined by a single character — kinship. Kinship turned out to be Lyubishchev's parameter by definition. The significance and possibilities of the cladistic system were discussed above; here I wish only to draw attention to the fact that this system requires no typification of its taxa, since reference to the value of the parameter (the nature of genealogical relationships) is sufficient to introduce a taxon into the system. Indeed, "names are synonymous if they refer to clades originating from the same ancestor" (de Queiroz, Gauthier, 1990: 307). The logical corollary of this assertion — namely, the rejection of the principle of the type (to say nothing of the Linnaean system of standard taxonomic ranks and other such matters) — was drawn by F. Pleijel and co-authors (Sundberg, Pleijel, 1994; Pleijel, Rouse, 2002), and the corresponding code of phylogenetic nomenclature (PhyloCode) is under development. Under this code, the role of the type fixing the application of a taxon name will be played by a cladogram node, an internode, or an apomorphy. The authors claim great advantages for phylogenetic nomenclature in terms of both ease of application and its stability (Pleijel, Rouse, 2002). This is difficult for me to accept, given the high degree of hypotheticality of the cladogram, which represents a system of hypotheses about the unobservable history of observable characters (see above). At this point, however, it is more sensible to pause and allow time to do its work. It will show how viable a taxonomic nomenclature built on cladistic principles can be. The logical alternative to the class as a group defined by its characters is the individual, and the proposal to treat a taxon as an individual (Ghiselin, 1974, 1987, and the bibliography cited therein) is perfectly natural. Possessing the properties of an individual, a taxon can be introduced into the system ostensively — that is, by direct reference (including reference to its name). For this purpose its integrity in space and time must be sufficiently high to allow the taxon to be born and to die, but not to disintegrate into parts, each of which could claim to inherit the taxon's name. Although a person on the way from infancy to old age may not retain a single common atom of matter or a single common character, through all his metamorphoses it is always possible to trace that he is the same individual. An individual is so completely integrated that the use of its name causes no problems. One can simply point a finger — there it is — without specifying precisely what the name refers to — the head, the heart, the ear, or something else. To everything at once and to each detail separately. This is exactly where the boundary between the ostensive method and the principle of the type lies: if any part of an individual equally bears its name, the special rules for selecting and using the nomenclatural type that serves to introduce a taxon into the system become superfluous. If the method of the type is inapplicable to an individual, then the taxon is not an individual. At least it is not a typical individual (see, for example, the commentary on the article by Ghiselin, 1981), so the problem requires further discussion. The paradigmatic example of an individual, the organism, is integrated because of the continuous interaction of its parts. The same has been asserted of the species, whose integrity is ensured by gene exchange (Mayr's biological species concept). However, this model has limited application, if only because "the biological species concept applies only to what I have called the 'non-dimensional situation,' where populations of the species are in actual [reproductive] contact" (Mayr, 1988: 301-302). In reality the problems here are even more numerous, as we have already seen (see the section on the ontology of evolution). However, the class-versus-individual dichotomy does not exhaust all possibilities. A taxon possesses features of both extremes. As a class it possesses characters — as witnessed by the existence of a diagnosis — and members (contrary to Ghiselin, I am not only a part of the species Homo sapiens but also an instance of it — that is, an example, a representative). As an individual, a taxon possesses parts (populations in relation to the species) and, more importantly, the capacity to evolve without losing its individuality. By evolving, the taxon preserves its integrity through time, in the multidimensional space of characters, and, to a greater or lesser extent, in geographical space. Such integrity allows the taxon to be identified as an individual — that is, by its name. Consequently, the taxon, possessing features of both a class and an individual, fills the logical space between these concepts and transforms class and individual into two poles of a single spectrum. Thus a taxon is something intermediate between a class and an individual. It is sufficiently integrated to preserve its name as a means of identification through change, but insufficiently integrated for a simple "there it is" to suffice for its identification. Insufficiently integrated, because a taxon is a continuum — a condensation of points, a cloud in the multidimensional space of characters. A cloud that can evolve — that is, change its size and position in character space — can disappear (become extinct) or, conversely, split in two or bud off a new cloud, and so on. The continuum is characterised both by its integrity — revealed by the presence of a hiatus (a discontinuity) between it and other cloud-continua — and by the characters of its diagnosis. However, both of these criteria lack rigidity and change readily both in the course of evolution and as our knowledge of the taxon accumulates. At any moment their inability to identify and delimit taxa clearly may become apparent. The possibility of identifying a continuum, unlike an individual, is determined by similarity — but unlike a class, it is a matter of a relation (overall similarity) rather than similarity with respect to specific characters. Therefore, pointing a finger cannot be addressed to the cloud as a whole, nor to its specific characters. It must be addressed to a specific point of the cloud — its nomenclatural type. Whatever happens to the taxon-cloud — in the course of its evolution, or as our understanding of it changes — the name fixed to the nomenclatural type, like a label on a museum specimen, will always indicate what is being discussed. Provided, of course, that the type is preserved — or at least described with adequate completeness. 1.2.2.2. Phenetics 1.2.2.3. Phyletics A.P. Rasnitsyn. Theoretical Foundations of Evolutionary Biology // V.V. Zherikhin, A.G. Ponomarenko, A.P. Rasnitsyn. Introduction to Palaeoentomology. Moscow: KMK. 2008. 371 pp. 1.2.2.3. Phyletics (continued) 1.2.3. TAXONOMIC NOMENCLATURE 1.2.3.1. Taxonomy-Independent Principles 1.2.3.2. Taxonomy-Dependent Principles 1.2.3.2. Taxonomy-Dependent Principles (continued) 1.2.3.3. Taxa Violating Principles of Nomenclature 1.2.2.3. Phyletics (continued) 1.2.3. TAXONOMIC NOMENCLATURE 1.2.3.1. Taxonomy-Independent Principles 1.2.3.2. Taxonomy-Dependent Principles 1.2.2.3. Phyletics (continued) 1.2.3. TAXONOMIC NOMENCLATURE 1.2.3.1. Taxonomy-Independent Principles 1.2.3.2. Taxonomy-Dependent Principles A.P. Rasnitsyn. Theoretical Foundations of Evolutionary Biology // V.V. Zherikhin, A.G. Ponomarenko, A.P. Rasnitsyn. Introduction to Palaeoentomology. Moscow: KMK. 2008. 371 pp. 1.2.3.2. Taxonomy-Dependent Principles (continued) 1.2.3.3. Taxa Violating Principles of Nomenclature 1.3. Conclusion References What has been said makes it clear that the nomenclatural type is a natural attribute of the traditional — that is, phyletic — as well as the phenetic system, and accordingly follows from the epigenetic theory of evolution. As for cladistics, the cladistic taxon is defined, as we have seen, by reference to a divergence event or to the synapomorphy acquired by that taxon (its first member) at the moment of its origin. It is precisely the divergence and/or acquisition of a synapomorphy that is conceived as the event creating the taxon. This event is the character that uniquely and exclusively defines the cladistic taxon. Thus a cladistic taxon is a class by definition, and no nomenclatural type is needed to identify it: all that is necessary and sufficient is, as already noted, the act of divergence and/or the synapomorphy. The same conclusion can be reached by a different and shorter line of reasoning. The aim of any classification is to cover all the diversity to be classified with taxa without remainder and, as far as possible, without overlapping taxon boundaries. If we accept that both the taxa themselves as natural groupings and the system they form exist in nature rather than having been invented by systematists (a view that also exists; see, e.g., S. Rasnitsyn, 2000), then the principal aim of classification is twofold. First, tracing natural discontinuities in the space being classified (using Plato's metaphor as quoted by Hull, 1983: 186, we carve nature at its joints). Second, we fix and rank the identified hiata as taxonomic boundaries (in artificial classification we impose these boundaries rather than tracing them). Classes do not lend themselves to the procedure described, because they are defined by characters that naturally overlap with other characters (unless one defines a class as elaborately as I did above in analysing the nature of the cladistic taxon). As for individuals, no natural forces or laws are known that could pack individuals tightly without leaving empty gaps (again, unless one sees a cladistic taxon as an individual on the grounds that it is born with the divergence of its ancestor, only to disappear in consequence of extinction or divergence). A continuum-taxon, by contrast, is ideally suited to the procedure of classification described, since the only legitimate way to define a continuum is by tracing and ranking its boundaries. One should not suppose that this is a unique property of the general system of organisms. It follows clearly from the reasoning of S.V. Meyen (1989) that the same is true, for example, of the meron — that is, the taxon of meronomy (the morphological system of parts of the organism) — and the straton, the taxon of stratigraphic classification. Beyond the principle of the type, there are two further taxonomy-dependent principles of nomenclature that, like the principle of the type, follow from the continual nature of the taxon. 8. The principle of hierarchy. Being continua, taxa can be united only into broader continua (senior taxa), which must encompass the included continua in their entirety. Equally, a continuum-taxon cannot simultaneously belong to two or more senior taxa, either in whole or in part, since otherwise these senior taxa would overlap, thereby violating the definition of a continuum. All of this means that the system has a strictly hierarchical form. 9. The principle of synonymy. If a taxon can belong to only one senior taxon, it has a unique position in the system and consequently requires only one name. All other names must be declared invalid — that is, treated as synonyms. 1.2.3.3. Taxa Violating Principles of Nomenclature Life is complex, and science too. Sometimes we are compelled to violate the principles of nomenclature — not out of ignorance or negligence, but of necessity, when serious reasons arise for doing so. Let us examine these possible reasons. Taxonomy-independent principles represent agreements among systematists and are in this sense subjective. The reasons for declining to follow them are always subjective as well, in the sense that they are not justified by any taxonomic considerations. They are therefore not discussed here. The problem of taxonomically dependent principles is more important and deserves analysis. A more detailed analysis was conducted earlier (Rasnitsyn, 1986a) and is presented here in abbreviated form. We are compelled to violate taxonomically dependent principles of nomenclature when the lack of information necessary for the classification of certain taxonomic material conflicts with the need to classify it — owing to its scientific or applied significance. The incompleteness of information in such cases is usually associated with the specific defectiveness of the material, as is well known in palaeontology and in the systematics of groups with complicated ontogeny (for example, parasitic worms and fungi). The International Codes of Nomenclature recognize certain forms of taxa with special status, but do not resolve the problem in its entirety, and the regulations contained in the codes are in many respects contradictory. The unsatisfactory state of affairs with taxa of special status can hardly be regarded as surprising, at least in zoology, where such taxa are traditionally considered rare and highly undesirable. In general (both in zoology and in botany) they are indeed not very numerous, but in certain specialized fields (palaeontology, parasitology, mycology) the refusal to employ them threatens a significant loss of information about objects of considerable stratigraphic, medical, agro- or silvicultural importance. The use of "deficient" taxa is unavoidable, but their incomplete regulation by the codes leads to nomenclatural inconsistency and improvisation. In palaeontology, for instance, essentially similar groupings are in some cases designated as special taxonomic categories (ichnotaxa in ichnology, including "induzirodae" and "induzipodrody" in the systematics of fossil cases built by larval insects — caddisflies (Vyalov, Sukacheva, 1976), turmae, anteturmae, and so on in palaeopalynology, form genera and recently abolished organ genera in the systematics of plant macrofossils, etc.). In other cases the name of the taxon is placed in quotation marks (Belokrys, 1984) or the taxonomic category is quoted (Ponomarenko, 1985); in yet other cases the status of the taxon is simply stated in the description or preamble. Occasionally this leads to the development of special nomenclatural codes (Sarjeant, Kennedy, 1973). This inconsistency is inconvenient, though not dangerous. Moreover, it is even useful as a means of searching for optimal methods of working with "deficient" taxa. What is dangerous is the desire — engendered by a negative attitude toward such taxa — to present them as normal and fully valid. It is remarkable, for example, that many specialists in conodonts (isolated microscopic skeletal elements of ancient animals) strive to demonstrate the validity of their taxa (Melville, 1981a,b). Even more illustrative is the situation with the extant genus Laberius Kieffer (Hymenoptera: Dryinidae). This genus was described (under the name Labeo Hal., nom. praeocc.) as an ordinary taxon, but was subsequently used as an aggregate group for males of the tribes Dryinini and Gonatopodini (Kieffer, 1914). These tribes are characterized by pronounced sexual dimorphism; their system is constructed on the basis of female characters, and the males, unless experimentally associated with females, could until recently not be identified even to tribe. However, in the revision of the family (Olmi, 1984) the author rejected such an interpretation of the genus Laberius and synonymized it with the genus Dicondylus Haliday (without discussing the characters of the type species of Laberius, that is, without substantiating his decision). At the same time, an extinct species described within the genus Laberius as understood in the sense of an aggregate group (N. Ponomarenko, 1981) was transferred to the genus Dryinus Latreille, with the note that the sole specimen of this species is preserved "so poorly that this does not allow its assignment to any genus of the family Dryinidae" (Olmi, 1984). Thereby Dryinus (the type genus of the family) was effectively converted into an aggregate group — that is, the most deficient form of a taxon (see below) — although the author's aim was clearly the opposite: to rid himself of the deficient taxon. Thus, the problem of taxa that fail to satisfy one or another rule of nomenclature remains topical. Let us analyze in greater detail the nature and forms of such taxa (Table 5). Table 5. Forms of taxa Taxon Belongs to system Type and diagnosis Upon synonymy with an orthotaxon orthotaxon general own subject to synonymization taxon of uncertain position general own subject to synonymization morphotaxon (parataxon) special own not subject to synonymization collective taxon (parataxon) special of superior taxon not subject to synonymization The simplest case is that of a taxon of uncertain position (taxon incertae sedis). It differs from a normal taxon (orthotaxon) in that the incompleteness of its characterization does not permit clarification of its position only at a certain level of the hierarchy. For example, a genus of uncertain position (genus incertae sedis) may at a given moment be assigned with sufficient confidence to a particular order, but not to a family. The extinct genus Baissobius Rasn., described among Hymenoptera of the suborder Apocrita, "possesses a peculiar type of reduced venation not known, so far as is known, in any other Hymenoptera" (Rasnitsyn, 1975, p. 128). At the same time, its venation was sufficiently peculiar to make determination of the systematic position of the insect difficult (other important characters are not visible on the fossil), but not peculiar enough to justify establishing a special family. Indeed, new finds made it possible to part with this taxon of uncertain position, converting it into a normal genus belonging to the family Embolemidae (Rasnitsyn, 1996b). The use of taxa of uncertain position may be understood as a local abandonment of the principle of synonymy. A genus subordinated to an order, bypassing the family, effectively functions as an independent but unnamed family — most probably synonymous with one of the existing families, only it is not yet known with which one. As is clear from the example of Baissobius, this synonymy may in time indeed become apparent. More complex are the cases united under the concept of parataxonomy. The definition treating a parataxon as a taxon in partial systems for isolated organs and body parts of animals (Melville, 1979) already narrows the practical application of this concept. Indeed, the specific features and problems associated with the introduction and use of a taxonomic name differ little in partial systems created for isolated organs and body parts of organisms, for a particular sex or life-cycle stage, or for traces of vital activity. Therefore a different definition is adopted here: a parataxon, in contrast to an orthotaxon and a taxon of uncertain position, belongs to an independent system connected with the main system by the principle of homonymy but not by the principle of synonymy, and therefore is not subject to synonymization even in cases where its synonymy with a particular orthotaxon has been demonstrated. Thus a parataxon is a taxon that is not subject to the requirement of uniqueness of taxonomic position — that is, the requirement of inclusion in the single hierarchy common to all organisms. It is to this requirement, in particular, that all five distinctive features of fossil material formulated by S. V. Meyen reduce (Meyen, Traverse, 1979; Meyen, 1983). Important considerations on the nature and functions of the parataxon are presented in another work (Meyen, 1990). Several kinds of parataxon are distinguished. The term formal taxon was proposed (Rasnitsyn, 1986a) as a broader interpretation of the botanical concept of form genus. Recently (ICBN, 2000) botanists replaced this term with the more apt concept of morphotaxon, which we shall employ here. A morphotaxon is the equivalent of a normal taxon (orthotaxon), but is employed only within the framework of a special system. This system runs parallel both to the main system and to other analogous partial systems, but is independent of them with respect to the principle of synonymy. Such are the systems of isolated fossil leaves, seeds, or imprints of beetle elytra, of fossil tracks of organism movement and other forms of their vital activity (for example, cases of caddisfly larvae), and the system of larval stages of modern parasitic worms or of males not associated with females in certain groups of insects where the system is oriented toward female characters. A morphotaxon differs from an orthotaxon only in that it may be — and sometimes certainly is — synonymous with a particular orthotaxon, yet it is not subject to synonymization. It is well known, for example, that in a number of cases for dispersed parts of ancient plants (leaves, trunks, roots, male and female fructifications, seeds and pollen), each described in its own special system, finds are also known from organic connection, making it possible to "reconstruct" the whole plant. Nevertheless, each such part retains its name received as a morphotaxon. Thus, the well-preserved Carboniferous trunks Lepidodendron, trunks Aspidiaria, Bergeria, Aspidiopsis, Knorria bearing bark shed to varying depth, roots Stigmaria, female strobili Lepidocarpon, male strobili Lepidostrobus, and microspores Lycospora (Fig. 7) could have belonged to the same plant, since the organic association of various pairs from this list has been established with greater or lesser reliability (Meyen, 1987b). And no one thereby contests the validity of all these generic names! [IMG_2] Fig. 7. Formal taxa proposed for the classification of isolated parts of lycopods (after Meyen, 1987b, 1988): a — Lepidodendron for trunks with intact bark (Aspidiaria, Bergeria, Aspidiopsis, Knorria for trunks that have lost their bark to varying degrees), Stigmaria for roots, Lepidocarpon for female strobili (showing general view and section of the megasporangium), and the microspore Lycospora extracted from the microsporangium of Lepidostrobus; b — all these names in certain cases designate the same plant, whose parts in various combinations have repeatedly been found in organic connection, as shown by lines (dotted lines indicate less reliable evidence of organic connection). With regard to certain cases of this kind it is maintained that they also bear on the principle of the type. Sometimes the reasoning behind such claims is difficult to understand (as in the case of the refusal to typify ichnotaxa — that is, taxa of the fossil trace system — in the version of the ICZN, 1988, Art. 66); in other cases the reasons are discussed in detail. A. G. Ponomarenko (1985) established non-typified formal taxa, defined solely by the characters of their diagnosis ("The genus [Flichea Handlirsch — A.R.] is understood as a formal assemblage of isolated beetle elytra without distinct grooves on the upper surface and with a noticeable notch in the middle of the outer margin", p. 76). As the author explained to me, the reason was the persistent tendency of certain systematists to synonymize and thereby eliminate parataxa, however arbitrary such synonymization might be, and in particular the example cited above of the genus Laberius Kieffer. The non-typified taxa of Ponomarenko are in essence the same morphotaxa, but artificially detypified and thereby placed outside the jurisdiction of the nomenclatural code (since the code regulates the application of a name by reference to a type, see above). Detypification indeed prevents unsubstantiated synonymization, but the problem is that it excludes all synonymization and therefore proves to be a cure worse than the disease itself. It seems to me that an understanding of the goals and functions of parataxonomy will be a better safeguard against arbitrary synonymization. The last and most impoverished kind of parataxon is the collective taxon (collective taxon). This is a parataxon, usually at the rank of genus, which is referred to a particular superior taxon, but in contrast to a morphotaxon cannot be placed within a special system parallel to the main one. In essence, species in collective groups are simply species of uncertain position (species incertae sedis). For example, the name "Cercaria O. F. Muller, 1773 was established for a genus of worms, and many authors in the 19th century treated it as the name of a nominal genus, as though its type species were C. lemna O. F. Muller, 1773. At present Cercaria is used as the name of a collective group for trematode larvae that cannot with confidence be referred to normal genera" (ICZN, 2000, Art. 67.14). The difference between taxa of uncertain position and species of a collective genus consists only in that a specific epithet may not appear alone, without a generic name. Special names of collective groups must therefore be introduced for such species; these names have the form of generic names but serve as names of the superior taxon to which the author refers the species included in the given collective genus. Thus Cercaria in the modern sense is the equivalent of a trematode order, Laberius in the interpretation of J. Kieffer (Kieffer, 1914) is the equivalent of Dryinini + Gonatopodini, and Dryinus in the interpretation of M. Olmi (Olmi, 1984) is the equivalent of Dryinidae. Collective taxa are considered detypified, but this assertion is erroneous. The definition given above means that a collective taxon corresponds in characterization and scope to the respective superior taxon, and its members, like orthotaxa, are ultimately introduced into the system by reference to the nomenclatural type of the latter. Species of Cercaria are definable as such simply by virtue of being sufficiently similar to the larvae of the type species of the Trematoda. And the fact that the type species of the Trematoda has not yet been formally fixed is immaterial: I have no doubt that superior taxa will be typified in zoology as well. Another example: the collective genus Carabilarva Ponomarenko, proposed for Mesozoic beetle larvae of the superfamily Caraboidea that cannot be identified to family (A. G. Ponomarenko, 1985). The type species of the genus is therefore the type of the superfamily, Carabus granulatus Linnaeus, 1758. Consequently, Carabilarva is a junior objective synonym of the genus Carabus, but is not subject to synonymization with it so long as it remains a parataxon. The last example implies the existence of yet another form of multiple subordination in nomenclature, in addition to the already discussed case where different preservational forms of one natural taxon are described as distinct morphotaxa. Here the same species proves to be the type of numerous collective groups. Indeed, the genus Carabus is simultaneously the type of the family Carabidae (ground beetles), the subfamily Carabinae, the tribe Carabini, and so on, and for the indeterminate members of each of these taxa special collective groups may theoretically be required, with separate ones for larvae and for various isolated body parts of the adult beetle. It is easy to imagine how many collective genera might as a result be based on the same nomenclatural type. For all its unfamiliarity, the situation can hardly give rise to serious nomenclatural or taxonomic problems if we understand what collective taxa are for. After all, genera and superior taxa of uncertain position trouble no one — why then should the treatment of species of uncertain position be any different? There is yet another important aspect of the problem of the deficient taxon (taxon of special status). A taxon belongs to this category simply because it is distinguished on the basis of "poor" characters — poor in comparison with other, normal taxa. If, however, there are no other taxa, the same deficient taxon proves to be fully adequate. If modern caddisflies did not exist, fossil cases would probably be studied without any connection to fossil caddisflies, and their system, based on the very same "poor" characters, would be considered normal. In many groups of insects, extant species are now distinguished almost exclusively by the characters of the male genitalia, while their extinct representatives are distinguished mainly by wing venation. So long as the fossils are referred to special, extinct genera, no nomenclatural problems arise, and it is possible to work with the fossils as orthotaxa. In cases where an extinct species falls into the same genus (subgenus) as extant ones, it frequently ends up in the category of the deficient. Faint consolation is afforded by the circumstance that long-described extant species whose genitalic structure has yet to be described deserve the same status. In addition to the conditional nature of the qualification of a taxon as an ortho- or parataxon, there are difficulties of another kind. Whether a character is good or poor is, first, a matter of degree (how great is the difference?), and second, a matter of taste. It is a common occurrence that a dispute between systematists reduces, in the final analysis, to the question of which set of characters should form the basis for elaborating the system of a particular group. One can only be surprised that in such disputes there has not yet arisen the mutual accusation that the system of one's opponent, based on such-and-such characters, is useful only as a partial system of parataxa, but not as the general (main) system of the group. All these considerations permit the conclusion that orthotaxon and parataxon are formally clearly distinct, but there is no substantive separateness here. Depending on one's point of view, and still more, as we have seen, depending on the level of our knowledge (the example of Baissobius), a taxon may shift from one category to the other. Apparently, parataxonomy should be treated pragmatically: if the use of parataxa genuinely facilitates the cognition and inventory of biodiversity, they should be introduced and employed. If, however, the gain is small, one should not complicate the situation by introducing parallel, overlapping systems of organisms. 1.2.2.3. Phyletics (continued) 1.2.3. TAXONOMIC NOMENCLATURE 1.2.3.1. Taxonomically independent principles 1.2.3.2. Taxonomically dependent principles A.P. Rasnitsyn. Theoretical foundations of evolutionary biology // V.V. Zherikhin, A.G. Ponomarenko, A.P. Rasnitsyn. Introduction to Palaeoentomology. Moscow: KMK. 2008. 371 p. 1.2.3.2. Taxonomically dependent principles (continued) 1.2.3.3. Taxa violating the principles of nomenclature 1.3. Conclusion References 1.2.3.2. Taxonomically dependent principles (continued) 1.2.3.3. Taxa violating the principles of nomenclature 1.2.3.2. Taxonomically dependent principles (continued) 1.2.3.3. Taxa violating the principles of nomenclature A.P. Rasnitsyn. Theoretical foundations of evolutionary biology // V.V. Zherikhin, A.G. Ponomarenko, A.P. Rasnitsyn. Introduction to Palaeoentomology. Moscow: KMK. 2008. 371 p. 1.3. Conclusion References 1.3. CONCLUSION The problems of the evolutionary wing of modern biology — which may be taken to encompass those of its branches that investigate the history and present state of biological diversity, including their theory and methodology — are connected above all with the search for a paradigm. The debate between two approaches — the "elementarist" ("reductionist") and the "holist" paradigms — has now continued for more than a century. Remarkably, no "demarcation" between these so divergent approaches has occurred, and the products of each of them coexist in the minds and publications of the great majority of scholars, who are usually entirely unaware of this. The elementarist paradigm, which reduces the system under study to its elements, corresponds to the population-genetic (synthetic) theory of evolution, representing the evolutionary process as the dynamics of the frequencies of discrete hereditary determinants (genes) in populations under conditions of selection. The observed evolutionary process is a reflection of this dynamics, and its specific character is determined, first, by the specific character of the genetic level of organization, and second, by the specific character of selection. Accordingly, the evolutionary process appears as essentially uniform, and the only regular source of discreteness — making possible the subdivision of the products of evolution into taxa — proves to be divergence and extinction. Adequate to this paradigm is the taxonomic concept of cladism, in which a taxon arises in the course of evolution as a product of divergence and is introduced into the system precisely on this basis, while the sequence of divergences is determined by means of synapomorphies. The alternative holist paradigm reduces the evolving system to the whole and in particular to the epigenotype — that is, to the system of construction of the living organism in ontogeny. Viewed through the lens of the epigenetic theory of evolution, the complex living system permeated by interwoven mutual connections and interactions appears as a tense compromise among the conflicting demands of optimizing different adaptive functions. Change in such a system proves to be substantially constrained, poorly predictable, and more or less saltational, while the specific character of the evolutionary process is determined above all by the specific character of the epigenotype and only to a substantially lesser degree by the specific character of selection. Because of the saltational nature of evolutionary changes, the biological diversity arising in the course of such evolution proves to be discrete, divided by hiatuses into natural entities, which are those that deserve to be used as taxa of the general system. Accordingly, the meaning of taxonomy adequate to the epigenetic concept and the holist paradigm consists in revealing objectively existing discreteness and tracing the hiatuses. Analysis of the available material testifies in favour of its correspondence with the epigenetic rather than the synthetic concept. More important, however, is the very attempt to distinguish the concepts and methodologies employed in the analysis of the evolutionary process and its results as deriving from one or the other of the two competing paradigms. It appears that a rigorous distinction between the two paradigms and their consequences will permit not so much, or not only, a deliberate choice between them as relief from the simultaneous use of mutually incompatible principles and approaches — that is, from a kind of pluralism within a single mind. The conclusion reached permits some advance in identifying the roots and more clearly formulating the methodological concepts employed in the field of the evolutionary wing of biology, and in particular in phylogenetics, taxonomy, and taxonomic nomenclature. However, we first had to examine some of the most general problems of scientific (and not only scientific) analysis. Phylogenetic inferences, like any scientific investigation, rest upon (1) planning of the forthcoming study, (2) observation (including experiment), (3) the search for analogies, (4) the creation of hypotheses about the regularities and mechanisms underlying the visible picture, (5) attempts at falsification of these hypotheses (primarily through analysis of their consequences), and (6) evaluation of the results of these attempts with the aid of presumptions, with the aim of selecting the most promising hypothesis for further work. The set of phylogenetic presumptions includes the presumption of the knowability of phylogeny and a number of more particular presumptions, which may be divided into those employed in the analysis of groups and those employed in the analysis of characters. Analysis of groups investigates ancestor-descendant relationships and rests above all on the palaeontological presumption for groups. The presumptions used in the analysis of characters are further divided into two groups according to whether they are included in the analysis of differences or of similarities. Analysis of differences consists in the polarization of transformation series — that is, the determination of plesiomorphic (ancestral) and apomorphic (derived) character states. Here the most important presumptions include the palaeontological presumption for characters, the biogenetic presumption, and the presumptions of analogy, irreversibility of evolution, functional perfection, complexity, rudiments, and conserved character distribution. The presumptions of the analysis of similarities are employed in resolving the question of whether a given similarity was inherited from a common ancestor or arose independently. These include above all the presumptions of parsimony and of weighted similarity. The aim of taxonomy is the creation of a system whose taxa are meaningful to the widest possible circle of users and, accordingly, are maximally homogeneous internally and divergent among themselves. Of the three principal competing approaches, cladism rests on the synthetic theory of evolution, which ignores the discreteness of biodiversity. The cladistic system therefore accounts only for divergences and recognizes only taxa characterized by synapomorphies. A synapomorphy is not a character but its interpretation; therefore the cladistic system not only ignores the discreteness of biodiversity but is also excessively hypothetical and fastidious in the selection of characters. Phyletics and phenetics rest on the epigenetic theory of evolution and are concerned with tracing the hiatuses between taxa. Phenetics employs available characters as such and is therefore too myopic — this is essentially an empirical approach. 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