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Volume 313 (Supplement 1), 2009. A.F. Alimov and S.D. Stepanjants (Eds.) Species and speciation. Analysis of new views and trends.

Saint Petersburg, Zoological Institute RAS, Company of Scientific Papers KMK, 298 p.

Date of publication — July 25, 2009

Preface. «Species and speciation». Analysis of new views and trends

The volume proposed is Supplement No. 1 to the periodical “Trudy Zoologicheskogo instituta Ross. Akad. Nauk” (Proceedings of the Zoological Institute of the Russian Academy of Sciences). The volume is conceived to present results of modern discussions on the topic “Species and Speciation”. Apart from historical articles dealing with the taxonomic concepts of plant and animal classification proposed by the founders Karl Linnaeus, Peter Artedi, Peter Simon Pallas and others (see articles by N. Bogutskaya, L. Borkin, A. Sytin, A. Ostrovsky), the volume includes estimates and interpretations of botanists, mycologists, and zoologists of orderliness of biological diversity and recognition or non-recognition of species and genera as natural categories. We deliberately present opposite views, so that the readers could form their own opinions of the problems.

Carl Linnaeus (1707–1778) as zoologist

The activity of Carl Linnaeus as zoologist is described. His life is briefly outlined in the context of his zoological interests. All twelve editions of “Systema Naturae” (from 1735 to 1766) are analysed, and the most important changes in the taxonomy of animals are given. So called 13th editions are mentioned as well. The importance of “Fauna Svecica” is considered, with comparison of its both editions (1746 и 1760). Other publications of Linnaeus, as well as numerous dissertations of his students, associated with zoology, are listed. Linnaeus’ contribution to the systematics, including taxonomic ranks and binary nomenclature, is evaluated. His species concept, his views on Homo sapiens and fossil animals are presented as well.

On a category of genus in the 18th century botany: Linnaeus and his opponents

The period of 1737 through 1739 was exceedingly important in the life of Linnaeus in Holland where he published his well-known «Genera Plantarum» (1737). This work is of importance in modern taxonomy as a source of descriptions of 935 genera (in all a total of 1336 Linnaeus’ genera of plants were diagnosed). Basically the Linnaean concept of the genus was in accord with the French botanist J.P. Tournefort that the fundamental category of classification was the genus, and that plants having in common two or three characters of reproductive structures were usually to be treated as members of the same genus. Linnaeus so– called sexual system «of plant classification» comprised 24 classes, 23 of which contain the flowering plants, with stamens and pistils was highly artificial was expanded and served as the basis for «Genera plantarum». At the same time Linnaeus preparing the manuscript for a sumptuous work «Hortus Cliffortianus» (1737), in which were named and described many temperate and tropical plan grown in the botanical garden by George Clifford, Director of the Dutch East India Company. The genus Сliffortia was selected to honour of the owner of plant collection. The Genera of Linnaeus were based largely on the belief that a genus is a category whose components (species) have the essential character, it discriminated between species to morphological distinctions of leaf forms: С. ilicifolia, C. polygonifolia, С. ruscifolia, С. trifoliata; discriminated through stem habit types: Phascum acaulon, P. caulescens, P. repens; geography and its relationship to plant distribution: Parietaria cretica, P. lusitanica, P. zeylanica – in a literal sense is illustrated Linnaeus’ metaphora: «plantae omnes utrinque affinitatem monstrant, uti Territorium in Mappa geograpica» («Philosophia botanica » (PhB) Systemata. II. 77). It is evident that according to Linnaeus’ concept of the genus the naming of plants must be reflected a significant selected characters(«Nomen specificum legitimum plantam ab omnibus congeneribus distinguat”(PhB) Differentia. VIII. 257). It is likely that the Linnaeus’ metod integrated an elements of system, was elucidated the structure of a genus as a natural and phyletic unit. Therefore this concept provides the basic for modern plant classification and flowering plant phylogeny. However, a few years later Peter Simon Pallas (1741–1811), a professor of St. Petersburg Academy of Sciences in his taxonomical works recognized several species in the genus Polycnemum (now partly Petrosimonia Bunge; Chenopodiaceae). Perhaps engage in controversy against Linnaeus’s study and pursue polemical goals he named plants with according to number of stamens: P. monandrum Pall. – with only one stamen in androecium; P. triandrum Pall. – with three stamens. It was a corrective action on the advancement of the method of natural classification and, on opposite sides, a parody on the Linnaeus’s artificial classification. A delimitation of taxonomical content of a genus as a natural units – the second most important problem by most contemporary scientist , but on this point the Linnaeus’s authority was incontestable for Pallas. He even renounced one’s point of view on the name of Rindera Pall., in consequence of Linnaeus’s critical opinion and this plant name unwillingly accepted as Cynoglossum L. in his «Flora Rossica» (1788, 1, 2 : 97). Only in last works as a first monograph of genus Astragalus «Species astragalorum descriptae et iconibus coloratis illustratae» (1800–1803), Pallas used as the basic personal method and recognized into the genus six almost natural groups («Astragalorum phalanges»). It was an important precondition to positioned approach of biological classification in botany, that «the true beginning and end of botany is the natural system» (PhB. Systemata. II. 77).

Peter Artedi and Carl Linnaeus – founding fathers of fish classification

The paper is a review of those aspects of the life and scientific activities of Artedi and Linnaeus which are immediately related to the creation of their classifications of fishes and scientific ichthyological nomen clature. A short comparative analysis of Artedi’s and Linnaeus’s publications is done to retrace the development of the fish classification from «Ichthyologia» by Artedi (1738) to 10th and 12th editions of «Systema Naturae» by Linnaeus (1758, 1766–1767).

Destiny of Carl Linnaeus’s collection

Collection of Carl Linnaeus is a historical and scientific treasure that was bought bt J.E. Smith from the Linnaeus’s widow and moved to Great Britain. Paper follows a destiny of the Linnaeus’s collection since the announcement of his will till present time.

Discontinuity and sporadicity as species features

Space and spatial dynamics of a taxonomic systems are described. Some species of antlions (Neuroptera: Myrmeleontidae) and other taxa of insects are hold up as an examples.

The systemic concept of formation and the problem of species

This concept of formation is quite similar to epigenetic one. The biological integrity is considered as an all-round concordance between constituents of structure, functioning, reacting and development, while an organism is seen as a dynamic-steady system of integrative relations between genome and phenome based on mutual influence. Any transformation or emergent self-organization within a living system is connected with the maintenance of its steady state. The adaptive constituent of the system’s steady state concerns to its fitness, and the structural constituent – to its steady-state structure. The only difference between ontogenesis and formation is that the object of the first one is an organism, while the object of the second one is a reproducing totality of organisms, which is elementary in the frame of typology and historically conditioned within a definite space. A species is considered as typologically indivisible and historically unrepeatable variant of steady relations between genome and phenome, which canalize the diversity of their own systemically permitted transformations. The «epigenetic» constituent of divergence between species is determined by difference in relations between phenome and genome, not by change of the latter’s compound. The «products» of epigenetic speciation distinctly differ in morphology and ecology, having a lack of post-zygotic reproductive isolation. The reproductive isolation between them is caused by ecological and ethological barriers. The genetic constituent of divergence between species is incompatibility between nonconspecific genomes, which more often is revealed in a form of hybrids sterility. The prevailing of the genetic constituent of interspecific divergence is caused by allopatrie in uniform environmental conditions. In such case, the genomic divergence is accompanied by invariable relations between genome and phenome, that determines morphological and ecological uniformity of cryptic species. The divergence between panmictic species more often includes both of the constituents in different ratio. Due to its universality, the systemic approach to the problem of species is also appropriate to apomictic organisms.

The pecularities of flowering plants speciation

Among the fundamental distinctions between plants and animals at the different levels of organization of the plant kingdom it is especially important distinctions on a population – species level of organization. This is a higher-order variety of racial structures or «eidologic» units typical of plants in connection with multiplicative processes as hybridization and specific features of biological isolating mechanisms of races (and species), polyploidy of different origin, agamic-and clonal types of asexual breeding etc. After the brief historical review of botanical works on these problems it is emphasized, what of diversity of plants the nature of micro-evolutionary transformations to populations has unequal basis (where frequently populations considerably hybrid breakdown, two ore more species etc.) and, actually, the speciation. Basic natural units of evolutionary process at plants are not only species (mono- or polytypical), but also it is long existing complexes of populations of the different species, interacting with each other (down to complex of syngameons uniting species of a genus or even different genera). The first who has definitely distinguished species and species-complexes was Wladimir Komarov. Now it is possible to distinguish at vascular plants some types of species (up to 7), and species-complexes (i.e. the basic eidological units, it is real evoluting in the nature). Consequently it is possible to discuss about the greater potentiality for evolutionary transformations for a vascular plants, than one at a vertebrate animals.

Pattern of morphological variety of angeospermous pollen aperture distribution and natural ordering of biological variety, or – what is a variety (problems of description and interpretation)

Beside typical pollen forms with symmetrical aperture arrangement, various deviant forms with less symmetrical arrangements have been observed in remote Angiospermous taxa. The pollen forms may be arranged in the continuous, geometrically regular and taxonomically unspecific transformational series. Continuous and regular pattern of the variety obtained can not be described by any way of the classification without complete destroying of the unity of this pattern due to the typological procedures (discretization). The paradox is considered: the variety described consists of discrete pollen grains (as the biological variety consists of mortal living bodies) i.e. has the discrete pattern, but the variety of pollen morphological characteristics have the continuous pattern. Problems of description of the pattern of the biological variety and ways of interpretation are discussed.

Molecular markers, phylogeography and search for the criteria for delimiting species

The search for practical criteria for delimiting species was always topical. The waves of species splitting and lumping always altered depending both from dominant species concept, fashion and methods applied. Starting from the end of the last century the description of species diversity is at the peak of splitting wave. The specific feature of current splitting wave is that it relies 1) on new wide array of characters – molecular markers with their universality and easy application; 2) wide application of molecular markers in its turn gave birth to new methodology – phylogenetic analysis penetrates to intraspecies level, new direction of studies – phylogeography (Avise et al., 1987) appears and explosively develops. Phylogeography very successfully fall on phylogenetic species concept using gene trees as the basis for delimiting species and this «tree-thinking» approach together with widely expanding studies on phylogeography lead to dramatic increase in species number practically in all groups of vertebrates. Unlike morphological characters, molecular markers are universal (occur in all or almost in all organisms) and genetic distances, therewith, at a first glance gave an universal metrics for delimiting species which could be applied to almost all groups. Thus remarkable and long-awaited perspective opens- systematics receive an universal tool for distinguishing and delimitation of species. However, this hope on universal criteria once again appeared to be false and all issues which rise while working with molecular markers are very similar to those one have using morphological approach. In other words application of molecular markers bring us back to old and well known issues major part of which currently has no solution and the feeling that biologists at last have gain a unit similar to exact sciences is very wrong.

Fractal and chaotic patterns in animal morphology

In the review the spatial organization of cells, cell ensembles, tissues and metazoan body are considered using the concepts of fractal geometry, topology and dynamic chaos theory. We investigated both the scenario of transition from chaos into order during self-organization of cells in vitro and the reverse scenario of transition from order to chaos in the fractal morphogenesis of metazoan cell systems. Chaotic features in animal morphology were identified and quantified. Fractal morphogenesis was studied using epithelial branching channels of gastrovascular system in the scyphomeduse Aurelia aurita and tracheal gill system in the mayfly larvae Siphlonurus immanis and Parameletus chelifer, as well as structures of colonial interna in rhizocephalan crustaceans Peltogasterella gracilis and Polyascus polygenea. It was shown that completely identical fractal patterns do not occur even within a single animal body with radial or bilateral symmetric, functionally equivalent repetitive modules. Fractal dimension was used to quantify the spatial complexity of neuron morphology in central nervous system of the fishes Pholidapus dybowskii, Oncorhyhchus keta and Oncorhyhchus masou. During ontogenesis of Oncorhyhchus masou the values of fractal dimension and linear morphometric indicators were rising in studied neuron groups. Probably biological morphogenesis with chaotic fractal regime had an advantage in evolution, providing morphofunctional variability, plasticity and adaptability to unpredictable environmental changes.

Discrete adaptive norms: mechanisms and role in evolution

In experiments with defensive morph formation in the clones of green algae Scenedesmus acutus, ciliate Euplotes aediculatus and crustacean Daphnia pulex was shown that actinomycin D (inhibitor of de novo transcription) inhibits both transformation of typical morph to the defensive one in the presence of enemies and the opposite transformation of the defensive morph to the typical one after removing of enemies or signals of their presence. From this it follows that each morph has the own gene program and forms capable to discrete adaptive norms formation have relatively more rich genome. The revision of the evolutional role of adaptive modifications were made.

The hierarchy and combinatorial space of characters in evolutionary systematics

In this paper, the Linnaean hierarchy is discussed as the taxonomic model for the evolutionary differentiation of the earth’s biota. This model allows us to understand why a number of ideas and arguments in the evolutionary theory are unfortunate. Among these is the idea of a linear ladder of nature («scala natura»). The arguments about species constancy versus species variability, natural species versus «artificial» higher taxa, a single type of animals versus multiple types, as well as about the connection versus disconnection of microevolution and macroevolution are meaningless. Two kinds of phylogeny are considered. The first reflects the appearance of new characters in the course of evolution, and their sequence provides the nesting hierarchy of groups. The second reflects the appearance of new character states and provides a basis for the diversity of taxa at each hierarchical level. The taxonomic hierarchy is an embodiment of the first kind of phylogeny and does not require the tracing of ancestors and descendants along the lines of character development. Character ranking and the improvement of a tentative taxonomic hierarchy with the help of a posteriori weighting of differences are discussed. The method of character weighting leads to the construction of prognostic combinatorial arrangements that can predict the existence of organisms with certain character state combinations at each hierarchical level. The drawbacks of cladistic methodology, especially the «synapomorphy principle», the «dichotomy principle», and the monophyly «definition» through the internal composition of groups, are noted. It is pointed out that these principles only lead to the creation of heterogeneous groups and wrong character ranking.

The species problem in biology – another look

The modern species problem in biology is defined as a contradiction between the need for general notion of the species having a unified contents in various branches of biology, and impossibility to reach it. Any species concept becomes biologically valid under conditions of a biologically sound basic theory, which defines what is the species in a general case and why and how does it come to existence. There a hierarchy of species concepts and definition exist, with a most general «ideal» concept (a kind of «species theory») belonging to its highest level of generality. No such concept is known to exist at the moment; one of its version could be elaborated within a theoretical framework of synergetic model of developing and structuring biota, with the species being an element of the biota’s structure. A number of particular contentwise species concepts belong to the middle level of that hierarchy, each corresponding to a particular aspect of consideration of the biota’s structure. Different middle-level species concepts and definitions could be valid for the different taxonomic groups and for the different tasks of exploration of that structure. Operational species concepts belong to the lower level of that hierarchy, which principal aim is to elaborate particular methods of recognition of particular species. To be biologically sound, they are to be consistent with certain middle-level concept(s). With the universal, biologically sound species concept being absent, it is the taxonomic species concept that is a classificatory unversalia. However, its biological contents is different in different groups of organisms. Judgments about species diversity are special kind of the «taxonomic hypotheses», which are dawn and tested within a framework of certain basic biological theory fixing certain aspect of consideration of the biota’s structure. Personal knowledge plays an important role in understanding of both the species problem in general and the ways it could be resolved in form of particular species concepts and definitions.

The version of the contemporary theory of evolutionary systematics

Integral theory of evolutional systematics is presented in this article for the first time in contemporary science. It became formed as the science about evolution of species’ diversity and methods of investigation of it. Here is defined its object, subject, aim and method. Theoretical evolutional systematics is separated from practical systematics. Three sections are included in its content: idiographical systematics, nomothetical systematics and phylogenetical cybernetics. Idiographical systematics includes theories of descriptions (= meronomy), classifications (= taxonomy) and reconstructions of phylogenesis (= phylonomy). Nomothetical systematics includes the laws of phylogenetics, postulates of systematics, axioms and theorems of evolutional systematics in a whole, forming deductive theoretical system of evolutional systematics (DTS ES). Status of laws is added to 21 conformities to natural laws of phylogenetics. Here are formulated 6 postulates of systematics. On the base of logical investigations of laws and postulates as statements, the laws of phylogenetics are represented in form of 6 axioms and 15 theorems. Postulates of systematics are considered as 6 axioms. DTS ES is represented in the paper on the base of analysis of connections between 12 axioms. Phylogenetical cybernetics includes interpretation of the theory on the some systemic and probabilistic models of species, their classification and reconstruction of phylogenesis, the examples are present in the article. It is divided on three sections of investigations : systemology, theory of control phylogenetical transformations and theory of information processes in phylogenesis. The sections of evolutional systematics are interpreted accordingly philosophical conception of the levels of scientific knowledge. Systematics and phylogenetics are considered as two aspects of evolutional systematics as united science, reflecting its onthology (= the laws of phylogenesis) and gnosiologyl (= postulates of systematics). This solution conforms to initial definition of evolutional systematics as the science of evolution of species’ diversity and methods of its investigation and conforms the contemporary darwinism.



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