THE SPECIES PROBLEM - ONGOING ISSUES Edited by Igor Ya. Pavlinov The Species Problem - Ongoing Issues http://dx.doi.org/10.5772/3313 Edited by Igor Ya. Pavlinov Contributors Richard Mayden, Kirk Fitzhugh, Igor Pavlinov, Jack W. Sites, Jr., Larissa Vasilyeva, Steven Stephenson, Richard Richards, Victor Shcherbakov, David N. Stamos, James Staley, Friedmann Vladimir Published by InTech Janeza Trdine 9, 51000 Rijeka, Croatia Copyright © 2013 InTech All chapters are Open Access distributed under the Creative Commons Attribution 3.0 license, which allows users to download, copy and build upon published articles even for commercial purposes, as long as the author and publisher are properly credited, which ensures maximum dissemination and a wider impact of our publications. After this work has been published by InTech, authors have the right to republish it, in whole or part, in any publication of which they are the author, and to make other personal use of the work. 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ISBN 978-953-51-0957-0 free online editions of InTech Books and Journals can be found at www.intechopen.com Contents Preface VII Section 1 Introductory 1 Chapter 1 The Species Problem, Why Again? 3 Igor Ya. Pavlinov Section 2 Conceptual Issues 39 Chapter 2 The Species Problem: A Conceptual Problem? 41 Richard A. Richards Chapter 3 Biological Species as a Form of Existence, the Higher Form 65 Victor Prokhorovich Shcherbakov Chapter 4 Defining ‘Species,’ ‘Biodiversity,’ and ‘Conservation’ by Their Transitive Relation 93 Kirk Fitzhugh Chapter 5 Transitioning Toward a Universal Species Concept for the Classification of all Organisms 131 James T. Staley Chapter 6 An Essentialistic View of the Species Problem 141 Larissa N. Vasilyeva and Steven L. Stephenson Chapter 7 Species, Trees, Characters, and Concepts: Ongoing Issues, Diverse Ideologies, and a Time for Reflection and Change 171 Richard L. Mayden Chapter 8 Conspecific Recognition Systems and the Rehabilitation of the Biological Species Concept in Ornithology 193 V. S. Friedmann Chapter 9 Species Delimitation: A Decade After the Renaissance 225 Arley Camargo and Jack Jr. Sites Section 3 Historical Issue 249 Chapter 10 Darwin’s Species Concept Revisited 251 David N. Stamos ContentsVI Preface Each branch of knowledge, if it is burdened with at least some portion of metaphysics and natur-philosophy, possesses a kind of “eternal problems”. They are both, “problems” and “eternal” because of the lack of trivial and unambiguous ultimate answers to the funda‐ mental questions such as “what” and “why” are there things of such and such kind in Na‐ ture. Any attempt of escaping them by putting and answering the question “how” and using some operational means, reduces science to technology of counting and measuring. In biology, one of the most fundamental burdens is the species problem; so in this sense, it is actually both the “problem” and “eternal”. This is reflected in the endless stream of pub‐ lications on the species subject, be they either (nature)philosophical issues of the “why” kind (why is there the species), or biological searches of the “what” kind (what are the spe‐ cies of particular organisms), or operational treatments of the “how” kind (how to count the species). In the last decades we have been witnessing appearance of several important landmark papers, monographs and collections on the species problem [1–16] stimulated by new rise of interest of the scientific community to the above old questions of “what” and “why” and occasionally of “how” regarding the species and forming together the problem of the same name. The present book provides another collection of papers on the species problem. It doesn’t pretend neither to be a kind of ‘landmark”, to reflect the current state of the problem in question nor to concentrate on some of its aspects. Rather, this book was initially designed as a forum for exposing ideas, which may provide a look at the species problem not fitting the “recognized” pattern(s). The book includes ten chapters with pretty high theoretical content which are pretty diverse in their subjects; they are divided more or less convention‐ ally between three sections, with most of them being placed in the “Conceptual Issues” sec‐ tion. The “Introductory” section opening the book includes the editor’s (Igor Ya. Pavlinov) chap‐ ter with a self-explaining title “The species problem, why again?“. Its focal point is consid‐ eration of the species problem from the standpoint of the modern non-classical science paradigm, with ontological relativity and subjective inherency being its central themes. Ac‐ cordingly, he concentrates on consideration of this problem in the context of three-parti‐ tioned cognitive situation, within which the entire species problem outlines a conceptual space of certain kind, with particular species concepts being its subspaces allowable to be treated as specific onto-epistemological models. It is stressed that the latter are ordered into a conceptual pyramid of various levels of generality, an “ultimate beginner” of which is supposed to be a kind of synergetic model of the evolving biota. Species ontological plural‐ ism is stressed, which follows from acknowledging gradual divergent evolution of the “specieshood” resulted in the latter’s different manifestations in different groups of organ‐ isms according to their particular life strategies. Richard A. Richards, in his chapter “The species problem: A conceptual problem?”, devel‐ ops some of just above ideas in somewhat more sophisticated manner. His main point is to clarify, whether the species problem is conceptual (theoretical) or empirical one, and his answer is decisively in favor of the first version. Richards consequently considers the prob‐ lem in question from standpoint of the conceptual framework (another term for the above conceptual space), his general attitude is principally monistic. So he asserts it that there is (or should be) a kind of a single general species concept corresponding to some “definition‐ al core” of that framework, with its different aspects belonging to the latter’s “descriptive periphery” corresponding to various particular species concepts. The main theme of Victor P. Shcherbakov’s chapter “Biological species as a form of exis‐ tence, the higher form” is rather naturphilosophical. He tries to explain an emergence and existence of the species of living beings, in its most general meaning, as a particular discrete unit at a particular level of generality within the hierarchically patterned Universe. He in‐ formally defined the species as a multiorganismic self-reproducing entity endowed with a kind of “substantive existence” due to specific interaction of its tokens (organisms). An ulti‐ mate conclusion of the chapter is that the species as a “higher form of existence” emerged in course of historical development of the living matter as an entity possessing, in contrast to particular organisms, a possibility to both change (to evolve) and to remain “itself” (to persist) potentially eternally. The chapter of Kirk Fitzhugh “Defining ‘species,’ ‘biodiversity,’ and ‘conservation’ by their transitive relations” concentrates around consideration of the species as a particular (with no special status) case of the taxon defined as an abductively inferred class of explanatory hypotheses (not ontological individuals) accounting for particular characters distributions among observed organisms. Such a basically epistemological concern of the species leads to a tentative definition of the latter as an explanatory account of occurrences of similar char‐ acter(s) among individuals by way of character origin and subsequent fixation. Interesting is Fitzhugh’s reconsideration of the biodiversity and conservation concepts based on his non-trivial understanding of the species. For him, biodiversity is a metaphor for the hy‐ potheses of taxa (species in particular) as surrogates for hypotheses of the past, proximate tokogeny; this makes the very notion of biodiversity redundant relative to the notion of taxa. Respectively, from such a standpoint, conservation does not “conserve” species or taxa, but ensures that circumscribed sets of organisms would continue tokogeny into the future. James T. Staley begins his chapter “Transitioning toward a universal species concept for the classification of all organisms” with a statement that development of a uniform species con‐ cept that applies to all organisms is one the most important goals in biology. He justly be‐ lieves that, for a species concept to be actually universal, it has to be applied to all and any kinds of organisms including prokaryotes. Taking the latter as a “standard” for considering various species concepts, he comes to a conclusion that the phylogenomic species concept could become accepted as such a sought universal species concept. Larissa Vasilieva’s & Steven L. Stephenson’s chapter “An essentialistic view of the species problem” consider the latter from a typological taxonomic standpoint. They try to re-vital‐ ize an essentialistic view of the species problem, emphasizing (with reference to E. Sober) PrefaceVIII that it is most compatible with the individual treatment of the species. Their general pre‐ sumption is that majority of the existing species concepts are based upon within-species processes and relationships and therefore are inconsistent. Vasilieva & Stephenson suggest to consider and to define species on the basis of relationships between intensions of taxa, which for them, as for typologists, are characters arranged hierarchically and defining as hierarchically arranged taxa, some of which (at certain level of hierarchy) are the species. Richard L. Mayden’s chapter gives an interesting and quite emotional consideration of sev‐ eral topics concerning misconceptions and misunderstandings of the species issues among theoreticians and “praticiens” dealing some or other ways with the species. These include confusions with theoretical vs empirical considerations, phylogenetic vs pure genetic con‐ siderations, etc. Mayden keeps on insisting that Evolutionary Species Concept is most gen‐ eral among existing and includes, as its particular case, the General Lineage Concept. Of special concern is a disintegration of the research community involved in theoretical dis‐ cussions and practical applications of the species issues, which may provide its own nega‐ tive impact on the biodiversity crisis; after all, if biodiversity is defined informally as a global “species pool”, then what have biodiversity experts to study and to conserve if theo‐ reticians cannot ever agree about the species? Vladimir S. Friedmann starts his chapter “Conspecific recognition systems and the rehabili‐ tation of the biological species concept in ornithology”, like some others in this book, with a kind of naturphilosophical consideration, now it is an acknowledgement of the biological matter being structured at supraorganismal level (putting aside any ecosystemic considera‐ tion), and then proceeds with seeking for mechanisms providing this structuredness. For him, they are basically elements of the “friend-foe” recognition system working at popula‐ tion level and accounting for both inner integrity and outer separateness of conspecific population systems in birds. This theoretical consideration is added with an empirical one, according to which particular species, in which the above recognition system is not studied, are to be delimitated based on morphological (in the largest sense) hiatuses. The chapter of Arley Camargo & Jack W. Sites “Species delimitation: A decade after the renaissance” seems to be, among contributions of this book, most fitting the mainstream of contemporary biology by considering a particular question of the species problem, namely species delimitation (SDL), at the genomic level. Their addressing to the above “how” kind of question involves some newest approach of finding discontinuties among populations using multi-locus coalescent-based method (MLCM). The authors expose in short history of the method in question, its advantages and shortages, and possible future developments. Though dealing with rather technical SDL approach, they conclude with pointing out its possible bearing on theoretical understanding of the species ontology being developed within a framework of the conception correlating this ontology with the speciation mecha‐ nisms. The “Historical Issues” section contains David N. Stamos’ chapter “Darwin’s species con‐ cept revisited”, which is of a special kind. He considers the species subject, unlike other contributors, in a historical aspect focusing on one but quite important episode in the re‐ cent history of the species concept, namely on its understanding by Charles Darwin. Sta‐ mos himself characterizes his approach as the “contextualist historical”, which presumes that emergence of the ideas, of whatever importance and level of generality in their per‐ spective evaluation, is to be considered not in an isolation from but within the local histori‐ Preface IX cal and, moreover, personal contexts. What he thinks to be true for the Darwin’s species concept, is much so for any other concepts being debated hotly in the contemporary biolo‐ gy. So I guess this chapter puts an important final point for the entire book stressing histor‐ ically delimited, and therefore particular, status of any solution of the “eternal” species problem. Igor Ya. Pavlinov Zoological Museum of Moscow State University, Russia PrefaceX [...]... at the present time, according to which the species in classification” becomes separated from the species in nature” and it is the latter that is now being considered as a focal point of the species problem [9-11] Respectively, dis‐ cussion of this problem should begin with consideration of the following key issues: What is the species problem? Why is it about just the species? Why is it just the problem? ... “secondary” species concepts, operational ones are nothing more than “tertiary” ones belonging to the lowest level of the conceptual pyramid Close to the previous one by its meaning is a division of the species concepts reflecting their belonging to the “first” and the “third” worlds of Popper (see 3.4), which are the species in 19 20 The Species Problem - Ongoing Issues Nature” and the species in theory”... eventually the species level of organization of the living matter, i.e defines the “spe‐ cies as a rank” From this, it is evident that one of the key issues in the species problem is that about the “specieshood”, namely, about that possible specific quality, which makes any species the species and distinguishes the latter from other units of the structured biota The main part of this issue is, whether a... and the species category problem [16, 28-29] In the terms adopted here, the species category is defined by the specieshood, while the species taxon is (quite roughly) defined by particular manifestation of the specieshood in particular groups of organisms One of the sources of the species problem is that biologists (and occasionally philosophers) put quite different questions analyzing the species. .. organization, functioning and evolution 15 16 The Species Problem - Ongoing Issues 3.5 In what senses are the species “real”? Within an intersection of the ontological and epistemological components of cognitive situa‐ tion of the species problem (see 3.2), theoretical issues concerning the species “modes of be‐ ing” are most important One of these involves ontology of the species units”, which consideration... beginner” for the species concepts Any of the conceptual pyramids of the species problem, in the ways of their construing con‐ sidered in the previous section, remains closed on itself However, in the terms of the above genus -species scheme supplemented with epistemologically interpreted incompleteness the orem (see 4), any kind of the species pyramid” should be built into a concept (theory) of the next... conventional species, phy‐ logroup, ecomorph, age phase, etc., and the species in its current common understanding is just “one the many” of these elements Accordingly, the tip of the conceptual pyramid of the species problem proper should be not any “ideal” species concept proper, but rather the general concept of the unit (element) of the biotic structure, a particular case of which is the species concept... which is the spe‐ cies notion as a theoretical construct At last, it will be shown that another “radical solu‐ tion” of the species problem may be just to acknowledge objective multiplicity of the “kinds of species of living beings, corresponding to which is subjective multiplicity of the species concepts 2 Whence the species problem Any cognitive problem is systemic by its nature, and the species problem. .. its strict sense; rather, the species in classification” is a judgment (hypothesis) about the species in Nature” put forward on the basis of some data at hands within the scientific context provided by particular species in theory” Thus, the species in The Species Problem, Why Again? http://dx.doi.org/10.5772/51960 classification” is a kind of connecting link between all three species realities”... regard, the species problem is quite comparable to the homology problem or to the gene problem; in each of them, respective unit, though uniformly called (the species, the homo‐ logue, the gene, respectively), are recognized and treated much differently in particular re‐ search programs Pretty curious seems to be a kind of “psychological” source of the species problem, i.e con‐ viction of the debate . species problem? Why is it about just the species? Why is it just the problem? [10, 12-13]. The Species Problem - Ongoing Issues4 2.1. What is the species problem Generally. regarding the species and forming together the problem of the same name. The present book provides another collection of papers on the species problem.