Abstract
Different concepts define species at the pattern-level grouping of organisms into discrete clusters, the level of the processes operating within and between populations leading to the formation and maintenance of these clusters, or the level of the inner-organismic genetic and molecular mechanisms that contribute to species cohesion or promote speciation. I argue that, unlike single-level approaches, a multi-level framework takes into account the complex sequences of cause-effect reinforcements leading to the formation and maintenance of various patterns, and allows for revisions and refinements of pattern-based characterizations in light of the gradual elucidation of the causes and mechanisms contributing to pattern formation and maintenance.
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Notes
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There are other ways of classifying species concepts, such as operational versus theoretical (Mayden 1999), prospective versus retrospective (O'Hara 1993), history-based versus character-based (Baum and Donoghue 1995), mechanistic versus historical (Luckow 1995), pattern- versus process-based (Cracraft 1983; Harrison 1998), class versus entity concepts (Reydon 2005).
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“An adaptive zone is some part of the resource space together with whatever predation and parasitism occurs on the group considered. It is a part of the environment, as distinct from the way of life of a taxon that may occupy it, and exists independently of any inhabitants it might have” (Van Valen 1976, 234). Only “minimally different” adaptive zones are taken into account in order to ensure that higher taxa do not count as ecological species.
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The description of the phenomenon varies from GCSC to PSC/GSC. The patterning captured by PSC/GSC has a temporal component (i.e., phylogeny); in contrast, the patterning captured by GCSC and diagnostic criteria associated with BSC focuses on the spatial/geographical aspect (i.e., diagnosable clusters in sympatry). However, the two are related aspects of the same phenomenon, namely the formation of spatial–temporal discrete clusters among organisms (Avise and Wollenberg 1997).
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Hudson and Coyne (2002, 1564) further argue that mechanism considerations are also relevant to genealogical species as well, on the grounds that the coalescence of genes alone does not guarantee that groups remain exclusive when isolated populations are once again able to exchange genes.
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Mayden makes a similar argument in reference to EvSC: “the non-operational Evolutionary Species Concept […] should serve as the theoretical concept appropriate for the category Species. As operational concepts, the remaining ideas have been incompatible with one another in their ability to encompass species diversity because each has restrictive criteria as to what qualifies as a species. However, the operational concepts can complement one another and do serve a vital role under the Evolutionary Species Concept as fundamental tools necessary for discovering diversity compatible with the primary theoretical concept” (1999, 95).
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Wiley (1978, 20) acknowledges a similar dependance vis-á-vis EvSC: “Separate evolutionary lineages (species) must be reproductively isolated from one another to the extent that this is required for maintaining their separate identities, tendencies, and historical fates.”
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Claims that the process-level is best suited to define species [e.g., (Ereshefsky 2010)] can be ruled out for the same reasons.
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Examples include the Hertz-Thomson characterization of cathode rays discussed by Achinstein (2001), modern nosological classifications whereby symptomatic descriptions encompass several distinct diseases defined in terms of etiology (main cause) and/or pathology (mechanism of the disease) [e.g., diabetes mellitus (Alberti and Zimmet 1998)], the splitting of the memory phenomenon into more and more specific kinds of memory (Roediger et al. 2002).
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Acknowledgments
I would like to thank Lindley Darden, Erika Milam, Eric Saidel, Pamela Henson, Joan Straumanis, the DC History and Philosophy of Biology group, as well as three anonymous reviewers and the editor for helpful comments and discussion. This work is supported by ‘Fonds de la recherche sur la société et la culture’, Québec, Canada [grant number 127231].
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Baetu, T.M. Defining Species: A Multi-Level Approach. Acta Biotheor 60, 239–255 (2012). https://doi.org/10.1007/s10441-011-9143-z
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DOI: https://doi.org/10.1007/s10441-011-9143-z