Rasnytsyn, 2008. Theoretical Foundations of Evolutionary Biology - 09

1.2.2.2. Phenetics 1.2.2.3. Phylogenetics

A.P. Rasnitsyn. Theoretical Foundations of Evolutionary Biology // V.V. Zherikhin, A.G. Ponomarenko, A.P. Rasnitsyn. Introduction to Paleoentomology. M.: KMK. 2008. 371 p.

1.2.2.3. Phylogenetics (continued)

1.2.3. TAXONOMIC NOMENCLATURE 1.2.3.1. Taxonomically Independent Principles 1.2.3.2. Taxonomically Dependent Principles

1.2.3.2. Taxonomically Dependent Principles (continued) 1.2.3.3. Taxa Violating Nomenclature Principles

1.2.3. TAXONOMIC NOMENCLATURE

Nomenclature is the set of agreements on how to create and use taxon names. It is usually perceived as a field of intricate details and simple, obvious basic principles. This is incorrect, first, because not all operative principles of biological nomenclature are yet recognized and formulated explicitly. Some are mentioned in codes (see ICZN, 1988, 2000; ICNB, 2000), but not as basic principles, and many principles are simply absent. Second, some basic principles are not pure conventions but reflect certain taxonomic concepts or are consequences of specific evolutionary hypotheses. Linnaean principle of binomial nomenclature is fully conventional and therefore easily compatible with any taxonomic concept that recognizes a hierarchy of genera and species. The principle of type, however, is a different matter, as discussed in detail later. Distinguishing these two types of principles is useful, and henceforth they will be called taxonomically independent and taxonomically dependent principles, respectively (Table 4). Table 4. Nomenclatural principles Taxonomically independent 1 non‑interference in taxonomic decisions beyond the limits imposed by taxonomically dependent principles

Taxonomically Independent

non-interference in taxonomic decisions beyond the limitations imposed by taxonomically dependent principles

limited scope of application

homonymy

priority

standardized, rank-specific names

supreme priority of the International Commission on Nomenclature

Taxonomically Dependent

type (no reasonable grounds for rejection)

hierarchy (no reasonable grounds for rejection)

synonymy (application to parataxa limited to the scope of the respective private system)

1.2.3.2. Taxonomically dependent principles There are three taxonomically dependent principles: 7. Principle of type. This is the most important of the taxonomically dependent principles, and it clearly illustrates the constraints that a taxonomic concept can impose on nomenclature. In the 1988 Code this principle is formulated as follows: “The nomenclatural type serves as an objective reference by which the application of a name is determined independently of possible changes in the taxon’s boundaries” (ICZN, 1988, Art. 61a; the wording in ICZN 2000, Art. 61.1, seems less successful: “Fixation of the nomenclatural type of a nominal taxon provides an objective reference for the application of that taxon’s name”). In other words, a taxon cannot be introduced into the system except, ultimately, by reference to a type bearing its name. To assess the meaning of this statement, consider alternative possibilities. The first that comes to mind is introducing a taxon into the system by reference to its characters. This would mean that the taxon’s characters perform a defining, not merely diagnostic, function, and the taxon corresponds to the concept of a “class” (Ghiselin, 1974, 1987). Various possibilities for such introduction exist; the simplest is a combinatorial system, i.e., a multidimensional matrix where each cell corresponds to a specific combination of characters and is the place of a taxon possessing that combination. A version of such a matrix is the polytomic table sometimes used by systematists, i.e., a rectangular matrix with rows representing taxa and columns representing characters. Consequently each cell reflects the state of one particular character in a given taxon. This form of system is indeed simple and sometimes very useful, but it does not meet the goals of a general organismic system. It lacks flexibility: we cannot locally adjust it to better fit the characters of a particular taxon, because introducing a new, redefining, or deleting an old character forces a revision of all taxa’s characterizations. Therefore the combinatorial system is rarely used even as an identification tool. A more flexible system defines taxa by ranked characters. In a usual diagnostic table the first character used has the highest rank because it divides the defined set into the largest groups. Subsequent characters further subdivide and have lower rank. This system is indeed convenient, though not as good as a general organismic system, due to the obviously arbitrary ranking of characters. Nevertheless, in a somewhat modified form (with a small number of high‑rank characters sufficient to form the system) this approach was popular among systematists. In K. Linnaeus (Linnaeus, 1751), for example, plant fruit‑type characters played a similar role. In its clearest form the approach was formulated by A.A. Lyubyshev (1923, 1966). Lyubyshev called for the search for a few highest‑rank characters (parameters) that determine the distribution of all other characters—just as nuclear charge determines atomic properties and, consequently, the position of elements in the periodic table. The result was expected to be a parametric system of organisms that would allow prediction of all important properties of corresponding taxa from these key characters (parameters). Lyubyshev’s task seemed hopeless, and he himself could not solve it. Nevertheless, a solution of a sort was found, but not where he looked. Ironically, this occurred in a field that Lyubyshev, as a convinced anti‑selectionist, considered false. The solution turned out to be the cladistic system with its central claim that an organism’s characters and, consequently, its position in the system are best determined by a single character—relationship. Relationship proved to be Lyubyshev’s parameter by definition. The meaning and possibilities of the cladistic system were discussed above; here I only wish to draw attention to the fact that this system does not require typification of its taxa, since reference to the value of the parameter (the nature of the relationship) is sufficient for introducing a taxon into the system. Indeed, “names are synonymous if they refer to clades [branches] descending from the same ancestor” (de Queiroz, Gauthier, 1990: 307). The logical conclusion from this statement, namely the rejection of the type principle (let alone the Linnaean system of standard taxonomic ranks and other minutiae), was made by F. Pleijel and co‑authors (Sundberg, Pleijel, 1994; Pleijel, Rouse, 2002), and the corresponding Phylogenetic Code of Nomenclature (PhyloCode) is under development. It is assumed that under this code the role of the type fixing name application will be performed either by a node of the cladogram, its internode, or an apomorphy. The authors claim great advantages of phylogenetic nomenclature in terms of both ease of use and stability (Pleijel, Rouse, 2002). I find it hard to believe due to the high degree of hypothetical nature of the cladogram, which represents a system of hypotheses about the unobservable history of observable characters (see above). Nevertheless, it is wiser to stop here and let time work. Time will show how viable a taxonomic nomenclature based on cladistic principles is. A logical alternative to a class as a group defined by its characters is an individual, and the proposal to consider a taxon as an individual (Ghiselin, 1974, 1987 and the bibliography therein) is quite natural. Possessing the properties of an individual, a taxon can be introduced into the system ostensively, i.e., by direct indication (including reference to its name). For this its integrity in space and time must be sufficiently high to allow the taxon to be born and die, but not to fragment into parts each of which could claim inheritance of the taxon’s name. Let a person, from infancy to old age, retain no single common atom of matter and no single common character, yet through all his metamorphoses one can trace that he is the same individual. An individual is so integral that using its name causes no problems. One can simply point—there it is—and need not specify whether the name refers to the head, heart, ear, or something else. To the whole at once and to each detail separately. Here the boundary between the ostensive method and the type principle is crossed: if any part of the individual equally bears its name, special rules for choosing and using a nomenclatural type for introducing the taxon become superfluous. If the type method is inapplicable to an individual, the taxon is not an individual. At least it is not a typical individual (see, for example, comments to Ghiselin, 1981), so the problem requires further discussion. A paradigmatic example of an individual, an organism, is whole because of the continuous interaction of its parts. The same was asserted for a species, whose integrity is ensured by gene flow (Mayr’s biological species concept). However, this model has limited application, partly because “the biological species concept applies only to what I have called ‘non‑extensive situations’, when populations of a species are in real [reproductive] contact” (Mayr, 1988: 301‑302). In fact, the problems are even greater (see the section on the ontology of evolution). Nevertheless, the class‑individual dichotomy does not exhaust all possibilities. A taxon possesses features of both extremes. As a class it has characters, as evidenced by the existence of a diagnosis, and members (contrary to Ghiselin, I am not only part of the species Homo sapiens but also its specimen, i.e., an example, a representative). As an individual the taxon has parts (populations relative to the species) and, more importantly, the capacity to develop (evolve) without losing its individuality. While evolving, the taxon retains its integrity over time, in multidimensional character space and, to some degree, in geographic space. This integrity allows the taxon to be identified as an individual, i.e., by its name. Consequently, a taxon possessing both class and individual traits fills the logical space between these concepts and turns 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 integral to retain its name as an identifier while changing, but not so integral that a simple “there it is” suffices for its identification. It is not sufficiently integral because a taxon is a continuum, a condensation of points, a cloud in multidimensional character space. A cloud that can evolve, i.e., change its size and position in character space, may disappear (go extinct) or, conversely, split in two or give rise to a new cloud, etc. The continuum is characterized both by integrity revealed by the presence of a hiatus (discontinuity) between it and other cloud‑continua, and by diagnostic characters. Yet both criteria lack rigidity and change easily both during evolution and as our knowledge of the taxon accumulates. At any moment their inability to clearly identify and delimit taxa may become apparent. The possibility of identifying a continuum, unlike an individual, is determined by similarity, but unlike a class it concerns relationship (overall similarity), not similarity of specific characters. Therefore pointing does not succeed in addressing either the whole cloud or its specific characters. It must be directed to a particular point of that cloud—its nomenclatural type. Whatever happens to the taxon‑cloud—whether through its evolution or through changes in our concepts of it—the name fixed to the nomenclatural type, like a label on a museum specimen, will always indicate what is meant, provided the type is preserved—or at least described with acceptable completeness. The foregoing makes clear that the nomenclatural type is a natural attribute of the traditional, i.e., phyletic, as well as phenetic system, and consequently derives from the epigenetic theory of evolution. As for cladism, a cladistic taxon is defined, as we have seen, by reference to a divergence event or to a synapomorphy acquired by that taxon (its first member) at its origin. The divergence or/and acquisition of a synapomorphy is conceived as the event creating the taxon. This event is the character that uniquely and unambiguously defines the cladistic taxon. Thus, a cladistic taxon is a class by definition, and its identification requires no nomenclatural type: all that is necessary and sufficient, as already noted, is the act of divergence and/or synapomorphy. The same result can be reached by a different and shorter line of reasoning.{ "translated_text": "The purpose of any classification is to cover, with taxa, the entire diversity subject to classification without remainder and, if possible, without overlapping taxon boundaries. If we assume that both the taxa themselves as natural groupings and the system they form exist in nature, rather than being invented by systematists (there is such a viewpoint, see, for example, S. Rasnytsyn, 2000), then the main goal of classification is dual. First, tracing natural breaks in continuity in the classified space (using the Platonic metaphor cited by Hull, 1983: 186, we dissect nature at its joints). Second, we fix and rank the identified giatuses as taxonomic boundaries (in artificial classification we impose these boundaries instead of tracing them). Classes do not lend themselves to the described procedure, as they are defined by characters that naturally overlap with other characters (unless one defines a class as cleverly as I did above, analyzing the nature of a cladistic taxon). As for individuals, we do not know the natural forces and regularities that could pack individuals tightly, leaving no gaps (again, unless one does not see an individual in a cladistic taxon on the basis that it is born with divergence from its ancestor, only to later disappear due to extinction or divergence). A taxon‑continuum, on the contrary, perfectly fits the stated classification procedure, since the only legitimate way to define a continuum is to trace and rank its boundaries. One should not think that this is a unique property of the overall system of organisms. From the reasoning of S.V. Meyer (1989) it follows with obviousness that the same applies, for example, to meron, i.e., a taxon of meronomy (the morphological system of organism parts), and straton — a taxon of stratigraphic classification. In addition to the type principle, there are two more taxonomy‑dependent nomenclatural principles that, like the type principle, follow from the continuum nature of a taxon. 8. Principle of hierarchy. Being continua, taxa cannot be combined except into the following broader continua (higher taxa), which must completely cover the included continua. Likewise, a taxon‑continuum cannot simultaneously belong to two or more higher taxa, either wholly or in part, because otherwise those higher taxa would overlap, thereby violating the definition of a continuum. All this means that the system has a strictly hierarchical form. 9. Principle of synonymy. If a taxon can be a member of a single higher taxon, it has a single place in the system and, therefore, needs only one name. All other names must be declared invalid, i.e., be designated as synonyms. 1.2.3.3. Taxa that violate nomenclatural principles Life is complex, and science is too. Sometimes we have to violate nomenclatural principles—not out of ignorance or negligence, but out of necessity, when serious reasons arise. Let us consider these possible reasons. Taxonomy‑independent principles are agreements among systematists and are thus subjective. The reasons for refusing to follow them are also always subjective in the sense that they are not justified by any taxonomic considerations. Therefore they are not discussed here. The problem of taxonomy‑dependent principles is more important and deserves analysis. A more detailed analysis was previously conducted (Rasnytsyn, 1986a) and is presented here in abbreviated form. We have to violate taxonomy‑dependent nomenclatural principles when a lack of information necessary for classifying certain taxonomic material conflicts with the need to classify it—due to its scientific or applied significance. In such cases, incompleteness of information is usually linked to specific material defects, as is well known in paleontology and in the systematics of groups with complex ontogeny (for example, parasitic worms and fungi). International nomenclatural codes recognize certain forms of taxa with special status, but they do not solve the problem as a whole, and the regulations contained in the codes are often contradictory. The unsatisfactory situation with taxa of special status is hard to consider unexpected, at least in zoology, where such taxa have traditionally been regarded as rare and highly undesirable. In general (both in zoology and botany) they are indeed not very numerous, but in some specific fields (paleontology, parasitology, mycology) abandoning their use threatens a great loss of information about objects of important stratigraphic, medical, agro‑ or forestry significance. The use of “incomplete” taxa is inevitable, but their incomplete regulation by the codes leads to nomenclatural chaos and ad‑hoc practices. In paleontology, for instance, essentially similar groupings are sometimes designated as special taxonomic categories (ichnotaxa in ichnology, in particular “induzirids” and “induzipodids” in the systematics of fossil dwellings built by insect larvae—streamlets (Vyalo, Sukacheva, 1976), turmas, anteturmas, etc., in paleopalynology, formal genera and recently abolished organ‑genera in the systematics of macroremains of plants, etc.). In other cases the taxon name is placed in quotation marks (Belokrys, 1984) or the taxonomic category (Ponomarenko, 1985), and in a third case the status of the taxon is simply mentioned in the description or preamble. Sometimes this leads to the development of special nomenclatural codes (Sarjeant, Kennedy, 1973). This chaos is inconvenient, though not dangerous. Moreover, it can even be useful as a way to search for optimal ways of working with “incomplete” taxa. What is dangerous is the desire, generated by a negative attitude toward such taxa, to present them as normal, complete entities. It is surprising, for example, the tendency of many specialists in conodonts (isolated microscopic elements of the skeleton of ancient animals) to portray their taxa as complete (Melville, 1981a,b). An even more demonstrative case involves the modern genus Laberius Kieffer (membrane‑winged insects of the family Dryinidae). This genus was described (under the name Labeo Hal., nom. praeocc.) as an ordinary taxon, but was later used as a collective group for the males of the tribes Dryinini and Gonatopodini (Kiefer, 1914). These tribes are characterized by strong sexual dimorphism, with their system built on female characters, while males, unless experimentally associated with females, could not be identified even to the tribe level until recently. However, in the revision of the family (Olmi, 1984) its author rejected this interpretation of the genus Laberius and reduced it to synonyms of the genus Dicondylus Haliday (without discussing the characters of the type species Laberius, i.e., without justifying his decision). At the same time, an extinct species described within the genus Laberius, understood as a collective group (N. Ponomarenko, 1981), was transferred to the genus Dryinus Latreille with the note that the sole specimen of this species was preserved “so poorly that it cannot be assigned to any genus of the family Dryinidae” (Olmi, 1984). Thus Dryinus (the type genus of the family) was effectively turned into a collective group, i.e., the most incomplete form of a taxon (see below), although the author’s aim was clearly the opposite—to eliminate an incomplete taxon. Consequently, the problem of taxa that do not satisfy certain nomenclatural rules remains relevant. Let us analyze in more detail the nature and forms of such taxa (Table 5). Table 5. Forms of taxa Taxon Belongs to system Type and diagnosis With synonymy with orthotaxon orthotaxon general own subject to synonymization taxon of unclear position general own subject to synonymization morphotaxon (parataxon) special own not subject to synonymization collective taxon (parataxon) special senior taxon not subject to synonymization" }The easiest case is that of a taxon incertae sedis. It differs from a normal taxon (optotaxon) in that the incompleteness of its characterization does not allow its position to be specified at a particular hierarchical level. For example, a genus incertae sedis at present may be assigned with reasonable confidence to a certain order, but not to a family. The extinct genus Baissobius Rasn., described among the membracoid insects of the suborder Apocrita, “possesses a distinctive type of reduced venation not known, as far as is known, from other Hymenoptera” (Rasnitsyn, 1975, p. 128). Its venation was sufficiently unusual to hinder the determination of the insect’s systematic position (other important characters are not visible in the fossil), yet not sufficiently unusual to justify the erection of a separate family. Indeed, new finds have allowed this taxon of uncertain position to be resolved, turning it into a normal genus belonging to the family Embolemidae (Rasnitsyn, 1996b).

Classes are not subject to the described procedure, as they are defined by characteristics that naturally overlap with other characteristics (unless the class is defined as subtly as I did above, analyzing the nature of a cladistic taxon). As for individuals, we are unaware of natural forces and laws that could pack individuals tightly without leaving gaps (again, unless an individual is considered in a cladistic taxon based on the fact that it is born with a divergence from its ancestor, only to disappear due to extinction or divergence). A taxon-continuum, on the contrary, perfectly matches the specified classification procedure, as the only legitimate way to define a continuum is to trace and rank its boundaries.

One should not think that this is a unique property of the general system of organisms. From the reflections of S. V. Meyen (1989), it clearly follows that, for example, a meron, i.e., a taxon of meronomy (a morphological system of organism parts), and a straton – a taxon of stratigraphic classification, are the same.

In addition to the type principle, there are two more taxonomically dependent principles of nomenclature which, like the type principle, stem from the continuous nature of a taxon.

8. The Principle of Hierarchy. Being continua, taxa cannot be combined in any other way than into the following broader continua (higher taxa), which must encompass the included continua entirely. Likewise, a taxon-continuum cannot simultaneously belong to two or more higher taxa, either entirely or in part, because otherwise these higher taxa would overlap, thereby violating the definition of a continuum. All of this means that the system has a strictly hierarchical form.

9. Principle of synonymy. If a taxon can be a member of a single higher taxon, it has a single place in the system and, accordingly, needs only one name. All other names must be recognized as invalid, i.e., declared synonyms.

1.2.3.3. Taxa Violating Nomenclature Principles Life is complex, and so is science. Sometimes we have to violate nomenclature principles – not out of ignorance or negligence, but out of necessity, when serious reasons arise. Let's consider these possible reasons.

Taxonomically independent principles are agreements between systematists and are subjective in this sense. The reasons for refusing to follow them are also always subjective in the sense that they are not based on any taxonomic considerations. Therefore, they are not considered here. The problem of taxonomically dependent principles is more important and deserves analysis. A more detailed analysis was carried out earlier (Rasnitsyn, 1986a) and is presented here in a shortened form.

We are forced to violate taxonomically dependent principles of nomenclature when a lack of information necessary for the classification of a particular taxonomic material conflicts with the need to classify it due to its scientific or applied significance. Incompleteness of information in these cases is usually associated with specific defects in the material, as is well known in paleontology and in the systematics of groups with complex ontogeny (e.g., parasitic worms and fungi).

International codes of nomenclature recognize certain forms of taxa of special status, but do not solve the problem as a whole, and the regulations in the codes are often contradictory. The unsatisfactory situation with taxa of special status can hardly be called unexpected, at least in zoology, where such taxa are traditionally considered rare and highly undesirable. In general (both in zoology and botany), they are indeed not very numerous, however, in some specific fields (paleontology, parasitology, mycology), refusing to use them threatens a great loss of information about objects of important stratigraphic, medical, agricultural, or forestry significance. The use of "incomplete" taxa is inevitable, but with their incomplete regulation by the codes, it leads to nomenclatural discord and arbitrariness. In paleontology, for example, fundamentally homogeneous groupings are in some cases designated as special taxonomic categories (ichnotaxa in ichnology, in particular "indusoids" and "indusopodites" in the systematics of fossil cases built by the larvae of caddisflies (Vyalov, Sukacheva, 1976), turmae, antiturmae, etc. in paleopalynology, formal genera and recently abolished organ-genera in the systematics of plant macrostructures, etc.). In other cases, the name of the taxon (Belokrys, 1984) or taxonomic category (Ponomarenko, 1985) is put in quotation marks, in others – the status of the taxon is simply clarified in the description or preamble. Sometimes it comes to the development of special codes of nomenclature (Sarjeant, Kennedy, 1973).

This discord is inconvenient, though not dangerous. Moreover, it is even useful as a way to find optimal forms of working with "incomplete" taxa. It is dangerous to present them as normal, complete taxa, born out of a negative attitude towards such taxa. It is strange, for example, that many specialists in conodonts (isolated microscopic elements of the skeletons of ancient animals) strive to present their taxa as complete (Melville, 1981a,b). The situation is even more demonstrative with the modern genus Laberius Kieffer (hymenopteran insects of the family Dryinidae). This genus was described (under the name Labeo Hal., nom. praeocc.) as a normal taxon, but then used as a catch-all group for males of the tribes Dryinini and Gonatopodini (Kiefer, 1914). These tribes are characterized by strong sexual dimorphism, with their system built on the characteristics of females, and males, if not experimentally associated with females, could not be identified even to the tribe until recently. However, in a revision of the family (Olmi, 1984), its author abandoned this interpretation of the genus Laberius and synonymized it with the genus Dicondylus Haliday (without discussing the characteristics of the type species of Laberius, i.e., without justifying his decision). At the same time, an extinct species described within the genus Laberius, understood as a catch-all group (N. Ponomarenko, 1981), was transferred to the genus Dryinus Latreille, with the indication that the only specimen of this species is preserved "so poorly that it does not allow it to be assigned to any genus of the family Dryinidae" (Olmi, 1984). Thus, Dryinus (the type genus of the family) was effectively turned into a catch-all group, i.e., the most incomplete form of a taxon (see below), although the author's goal was clearly the opposite – to get rid of an incomplete taxon.

Thus, the problem of taxa that do not comply with certain nomenclature rules remains relevant. Let's analyze in more detail the nature and forms of such taxa (Table 5).

Table 5. Forms of taxa

| Taxon | Belongs to system | Type and diagnosis | Under synonymy with orthotaxon | |--------|------------------|---------------|------------------------------| | orthotaxon | general | own | subject to synonymization | | taxon of uncertain position | general | own | subject to synonymization | | morphotaxon (parataxon) | special | own | not subject to synonymization | | aggregate taxon (parataxon) | special | senior taxon | not subject to synonymization |The easiest case is a taxon of uncertain position (taxon incertae sedis). It differs from a regular taxon (optotaxon) in that the incompleteness of its characterization does not allow its placement to be specified at a particular hierarchical level. For example, a genus of uncertain position (genus incertae sedis) may currently be assigned with sufficient confidence to a particular order, but not to a family. The extinct genus Baissobius Rasn., described among the Hymenoptera of the suborder Apocrita, "possesses a peculiar type of reduced venation, which, as far as is known, is not observed in other Hymenoptera" (Rasnitsyn, 1975, p. 128). At the same time, its venation was peculiar enough to complicate the determination of the insect's systematic position (other important features are not visible on the fossil), but not peculiar enough to justify the establishment of a separate family. Indeed, new findings have allowed us to bid farewell to this taxon of uncertain position, transforming it into a regular genus belonging to the family Embolemidae (Rasnitsyn, 1996b).

The use of taxa of uncertain placement can be understood as a local deviation from the principle of synonymy. A genus subordinate to an order, bypassing the family, effectively acts as an independent but unnamed family, likely synonymous with one of the existing families, only it is not yet known which one. As seen in the example of Baissobius, this synonymy may indeed become known over time.

More complex cases, united by the concept of parataxon. The definition that treats a parataxon as a taxon in partial systems for isolated organs and body parts of animals (Melville, 1979) has already lost practical application of this concept. Indeed, the specifics 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 specific sex or life cycle stage, or for trace fossils. Therefore, a different definition is adopted here: a parataxon, unlike an orthotaxon and a taxon of uncertain position, belongs to an independent system connected to the main system by the principle of homonymy, not the principle of synonymy, and therefore is not subject to synonymization even in cases where its synonymy with some orthotaxon is indicated. Thus, a parataxon is a taxon to which the requirement of uniqueness of taxonomic position, i.e., the requirement of belonging to a single hierarchy for all organisms, does not apply. This, in particular, is the basis for all five characteristics of the uniqueness of fossil material formulated by S.V. Meyen (Meyen, Traverse, 1979; Meyen, 1983). Important considerations about the nature and functions of a parataxon are given in another work (Meyen, 1990).

Several types of parataxa are distinguished. The term formal taxon was proposed by Rantsitsyn (1986a) as a broader interpretation of the botanical concept of form-genus. Recently (ICN, 2000), botanists replaced this term with the more appropriate concept of morphotaxon, which we will use. A morphotaxon is an equivalent of a regular taxon (orthotaxon) but is used only within a special system. This system is parallel to the main system and other similar partial systems but is independent of them regarding the principle of synonymy. Such systems exist for isolated fossil leaves, seeds or beetle wing impressions, fossil traces of organism movement, and other forms of their life activities (e.g., caddisfly larval cases), systems of larval stages of modern parasitic worms, or males not associated with females in some insect groups where the system is focused on female characteristics.

[IMG_1] Fig. 7. Formal taxa proposed for the classification of isolated parts of lepidophytes (after Meyen, 1987b, 1988): a — Lepidodendron for stems with preserved bark (Aspidiaria, Bergeria, Aspidiopsis, Krorria for stems that have lost bark to varying degrees), Stigmaria for roots, Lepidocarpon for female strobili (showing a general view and a cross‑section of a megasporangium), and the microspore Lycospora extracted from the megasporangium Lepidostrobus; b — in some cases all these names denote the same plant, whose parts have been repeatedly found together in life‑time associations, as indicated by the lines (dashed — less reliable evidence of life‑time connection). Regarding some cases of this kind it is claimed that they also affect the type principle. Sometimes the reasons for such claims are hard to grasp (as in the refusal to typify “ethnotaxa”, i.e., taxa of the fossil‑trace system, in the ICZN 1988 version, Art. 66), while in other cases the reasons are discussed in detail. A.G. Ponomarenko (1985) identified non‑typified formal taxa defined solely by diagnostic characters (“The genus [Flichea Handlirsch.—A.R.] is understood as a formal aggregation of isolated beetle elytra lacking distinct grooves on the dorsal surface and with a conspicuous notch in the middle of the outer margin”, p. 76). As the author explained to me, the cause was a persistent effort by some systematists to synonymize and thus eliminate parataxa, however arbitrary such synonymization might be, exemplified by the earlier case of the genus Laberius Kieffer. The non‑typified taxa of Ponomarenko are essentially the same morphotaxa, but artificially detypified and thereby removed from the jurisdiction of the nomenclatural code (since the code regulates the use of a name by reference to a type, see above). Detypification indeed prevents unwarranted synonymization, but the problem is that it excludes any synonymization and thus becomes a remedy worse than the disease. It seems to me that understanding the goals and functions of parataxonomy will provide the best protection against arbitrary synonymization.

Regarding some cases of this genus, it is asserted that they also touch upon the principle of type. Sometimes the reasonableness of such statements is difficult to understand (as in the case of refusing to typify their taxa, i.e., taxa of trace fossil systems, in the ICZN version, 1988, Art. 66), in other cases the reasons are discussed in detail. A.G. Ponomarenko (1985) established untypified formal taxa, defined only by the features of their diagnosis ("Genus [Flichea Handlirsch.—A.R.] is understood as a formal association of isolated beetle elytra without distinct grooves on the upper side and with a noticeable notch in the middle of the outer edge", p. 76]). As the author explained to me, the reason was the persistent desire of some systematists to synonymize and thus eliminate parataxa, however arbitrary such synonymization might be, and in particular the aforementioned example with the genus Laberius Kieffer. Ponomarenko's untypified taxa are essentially the same as morphotaxa, but artificially detypified and thus removed from the jurisdiction of the nomenclature code (since the code regulates the application of a name by reference to a type, see above). Detypification indeed prevents unjustified synonymization, but the problem is that it excludes any synonymization and therefore turns out to be a cure worse than the disease. It seems to me that understanding the goals and functions of parataxonomy will be a better protection against arbitrary synonymization.

The most impoverished form of parataxon is the collective taxon. This is a parataxon, usually at the rank of genus, that is assigned to a higher taxon but, unlike a morphotaxon, cannot be placed in a special system parallel to the main one. In effect, species in collective groups are simply species incertae sedis. For example, the name Cercaria O.F. Muller, 1773 was established for a worm genus, and many 19th‑century authors treated it as a nominal‑genus name, as if its type species were C. lemna O.F. Muller, 1773. Today Cercaria is used as the name of a collective group for trematode larvae that cannot be confidently assigned to normal genera (ICZN, 2000, Art. 67.14). The difference between taxa of uncertain position and species of a collective genus lies only in the fact that a specific epithet may not appear alone, without a generic name. Therefore, for such species a collective‑group name must be introduced, which has the form of a generic name but functions as the name of the higher taxon to which the author assigns the included species. Thus, Cercaria in modern usage is equivalent to the order Trematoda, Laberius in the interpretation of J. Kieffer (Kieffer, 1914) is equivalent to Dryinini + Gonatopodini, and Dryinus in the interpretation of M. Olmi (Olmi, 1984) is equivalent to Dryinidae.

Collective taxa are considered detypified, but this statement is erroneous. The definition above means that a collective taxon coincides in its characterization and scope with the corresponding higher taxon, and its members, like orthtaxa, are ultimately introduced into the system by reference to the nomenclatural type of the latter. Species of Cercaria are defined as such simply because they closely resemble the larvae of a typical trematode species. The fact that a type species of trematodes has not yet been formally fixed is irrelevant: I have no doubt that higher taxa will be typified in zoology as well. Another example is the collective genus Carabilarva Ponomarenko, proposed for Mesozoic beetle larvae of the superfamily Caraboidea that cannot be placed to family level (A.G. Ponomarenko, 1985). Consequently, the type species of the genus is the type of the superfamily Carabus granulatus Linnaeus, 1758. Hence, Carabilarva is a junior objective synonym of the genus Carabus, but it cannot be synonymized with it while it remains a parataxon.

The last example signifies the existence of yet another form of multiple subordination in nomenclature, besides the already discussed case where different forms of preservation of one natural taxon are described as different morphotaxa. Here, the same species turns out to be the type for a multitude of collective groups. Indeed, the genus Carabus is simultaneously the type for the family Carabidae (ground beetles), the subfamily Carabinae, the tribe Carabini, etc., and for further undefined members of each of these taxa, their own collective groups may theoretically be needed, which are different for larvae and different isolated parts of the adult beetle's body. It is easy to imagine how many collective genera may ultimately be based on the same nomenclatural type. Despite the unusualness of the situation, it is unlikely to cause serious nomenclatural or taxonomic problems if we understand why collective taxa are needed. After all, no one is bothered by genera and higher taxa of uncertain position, so why should the attitude towards species of uncertain position be different?

There is another important aspect of the problem of an incomplete taxon (a taxon of special status). A taxon belongs to this category simply because it is distinguished by "poor" characteristics – poor in comparison with other, ordinary taxa. If there are no other taxa, the same incomplete taxon turns out to be quite complete. If modern caddisflies did not exist, fossil caddisflies would probably be studied outside any connection with fossil caddisflies, and their system, based on the same "poor" characteristics, would be considered ordinary. In many insect groups, modern species are currently distinguished almost solely by the characteristics of male genitalia, while their extinct representatives are mainly distinguished by wing venation. As long as fossil species are assigned to special, extinct genera, no nomenclatural problems arise, and fossil species can be treated as orthotaxa. Where an extinct species falls into the same genus (subgenus) as modern ones, it often ends up in the category of incomplete. A weak consolation here is the fact that long-described modern species, for which the structure of the genitalia has not yet been described, deserve the same.

Besides the convention of qualifying a taxon as an ortho- or parataxon, there are difficulties of another kind. A good character or a bad one is, firstly, a measure (how large the difference is), and secondly, a matter of taste. It is common for disputes between systematists to ultimately boil down to which set of characters should be relied upon when developing a system for a specific group. One can only be surprised that mutual accusations have not yet arisen in the dispute, that, so to speak, the opponent's system, based on certain characters, is useful only as a partial system of parataxa, and not as a general (main) system of the group.

All these considerations allow us to conclude that an orthotaxon and a parataxon are formally distinct, but there is no conceptual separation here. Depending on the point of view, and even more so, as we have seen, on the level of our knowledge (the example of Baissobius), a taxon can move from one category to another. It seems that parataxonomy should be approached pragmatically: if the use of parataxa genuinely facilitates the understanding and inventory of biodiversity, they should be introduced and applied. If the benefit is small, there is no need to complicate the situation by introducing parallel, overlapping systems of organisms.

The alternative holistic paradigm reduces the evolving system to a whole, in particular to the epigenotype—that is, to the system of constructing a living organism in ontogeny. In the mirror of epigenetic evolutionary theory, a complex, interwoven living system appears as a tense compromise of conflicting demands to optimize various adaptive functions. Change in such a system proves to be substantially hindered, poorly predictable, and more or less punctuated, and the specificity of the evolutionary process is determined primarily by the specificity of the epigenotype and only to a much lesser extent by the specificity of selection. Because of the punctuated nature of evolutionary changes, the biological diversity that arises during such evolution is discrete, broken into giatuses as natural units, which deserve use as taxa of the overall system. Accordingly, the meaning of taxonomy adequate to the epigenetic concept and the holistic paradigm lies in revealing objectively existing discreteness and tracing giatuses. Analysis of the available material supports its correspondence more with the epigenetic than with the synthetic concept. Yet an even more important issue appears to be the very attempt to distinguish the concepts and methodologies used in analyzing the evolutionary process and its results as stemming from one or the other of the two competing paradigms. It seems that a strict differentiation of the two paradigms and their consequences will allow not only a meaningful choice between them but also the elimination of the simultaneous use of mutually incompatible principles and approaches—that is, a kind of pluralism in a single mind. The conclusion drawn enables a modest advance in identifying the roots and more clearly formulating the methodological concepts employed in the field of the evolutionary wing of biology, particularly in phylogenetics, taxonomy, and taxonomic nomenclature. However, at the outset we had to consider some of the most general problems of scientific (and not only) analysis. Phylogenetic conclusions, like any scientific investigation, rely on (1) planning the forthcoming study, (2) observation (including experiment), (3) searching for analogies, (4) formulating hypotheses about the underlying regularities and mechanisms of the visible picture, (5) attempts at falsifying these hypotheses (mainly by analyzing their consequences), and (6) evaluating the results of these attempts using 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 specific presumptions, which can be divided into those used in group analysis and those used in character analysis. Group analysis investigates ancestor‑descendant relationships and relies primarily on the paleontological presumption for groups. Presumptions used in character analysis are further split into two groups according to whether they are included in analyses of differences or similarities. Difference analysis is the polarization of transformation series, i.e., the determination of plesiomorphic (ancestral) and apomorphic (derived) character states. Here, among the more important, operate the paleontological presumption for characters, the biogenetic presumption, presumptions of analogy, irreversibility of evolution, functional perfection, complexity, rudiments, and conserved distribution of characters. Presumptions of similarity analysis are used when deciding whether a given similarity is inherited from a common ancestor or arose independently. These primarily include the presumptions of parsimony and weighted similarity. The purpose of taxonomy is to create a system whose taxa are meaningful for the widest possible range of users and, accordingly, are most homogeneous within themselves and distinct from each other. Of the three main competing approaches, cladism relies on the synthetic theory of evolution, which ignores the discreteness of biodiversity. Therefore, a cladistic system accounts only for divergences and recognizes only taxa characterized by synapomorphies. A synapomorphy is not a character but its interpretation, so the cladistic system not only ignores biodiversity discreteness but is also overly hypothetical and picky in character selection. Phylotaxy and phenetaxy are based on the epigenetic theory of evolution and are engaged in tracing giatuses between taxa. Phenetaxy uses available characters as such and is thus overly myopic—a substantially empirical approach. Phylotaxy occupies an intermediate position between phenetaxy and cladism. It attempts to reflect a full balance of similarities and differences, including those not yet studied. For this, phylotaxy employs the predictive capacities of phylogeny, which serves as a heuristic method for controlling a system built by phenetaxic methods. Phylotaxy defines a taxon as a monophyletic continuum and uses giatuses to delimit taxa, while monophyly serves as a way to assess whether delimitation was successful or the system requires refinement. Taxonomic nomenclature rests on nine principles, six of which are pure conventions for convenience and uniformity in the use of taxon names and are independent of the taxonomic approach employed. The remaining three principles depend on taxonomy. 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History of Insects. Dodrecht etc., Kluwer Academic Publishers. P. 303-324.\n\n1.2.2.2. Phylogenetics\n1.2.2.3. Phyletics\n\nA.P. Rasnitsyn. Theoretical foundations of evolutionary biology // V.V. Zherikhin, A.G. Ponomarenko, A.P. Rasnitsyn. Introduction to paleoentomology. Moscow: KMK. 2008. 371 p.\n\n1.2.2.3. Phyletics (continuation)\n\n1.2.3. TAXONOMIC NOMENCLATURE\n1.2.3.1. Taxonomically independent principles\n1.2.3.2. Taxonomically dependent principles\n\n\n1.2.3.2. Taxonomically dependent principles (continuation)\n1.2.3.3. Taxa violating nomenclatural principles" }

1.2.2.2. Phenetics 1.2.2.3. Phylogenetics

A. P. Rasnitsky. Theoretical Foundations of Evolutionary Biology // V. V. Zherikhin, A. G. Ponomarenko, A. P. Rasnitsky. Introduction to Paleoentomology. M.: KMK. 2008. 371 pp.

1.2.2.3. Phylogenetics (continued)

1.2.3. TAXONOMIC NOMENCLATURE 1.2.3.1. Taxonomically Independent Principles 1.2.3.2. Taxonomically Dependent Principles

1.2.3.2. Taxonomically Dependent Principles (continued) 1.2.3.3. Taxa Violating Nomenclature Principles