Species: Turning a Conundrum Into a Research Program
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View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by DigitalCommons@University of Nebraska University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Faculty Publications from the Harold W. Manter Laboratory of Parasitology Parasitology, Harold W. Manter Laboratory of 6-1999 Species: Turning a Conundrum Into a Research Program Daniel R. Brooks University of Toronto, [email protected] Deborah A. McLennan University of Toronto E. C. Bernard University of Tennessee Follow this and additional works at: https://digitalcommons.unl.edu/parasitologyfacpubs Part of the Parasitology Commons Brooks, Daniel R.; McLennan, Deborah A.; and Bernard, E. C., "Species: Turning a Conundrum Into a Research Program" (1999). Faculty Publications from the Harold W. Manter Laboratory of Parasitology. 309. https://digitalcommons.unl.edu/parasitologyfacpubs/309 This Article is brought to you for free and open access by the Parasitology, Harold W. Manter Laboratory of at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Faculty Publications from the Harold W. Manter Laboratory of Parasitology by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Journal of Nematology 31(2):117–133. 1999. © The Society of Nematologists 1999. Species: Turning a Conundrum into a Research Program1 Daniel R. Brooks and Deborah A. McLennan2 Abstract: The most appropriate ontological basis for understanding the role of species in evolutionary biology is the Evolutionary Species Concept. The ESC is not an operational concept, but one version of the Phylogenetic Species Concept is. Linking the ontology of species with the epistemological basis of actual biological studies requires that we specify both a discovery mode for identifying collections of organisms that we believe are evolutionary species, and a series of evaluation criteria for assessing those entities we have discovered. Simply naming a collection of specimens, no matter how strong one’s evolutionary beliefs, is not sufficient for declaring that evolutionary species have been discovered. All operational historical species concepts represent discovery modes with minimal evaluation criteria; all operational non-dimensional species concepts represent evaluation criteria that do not specify discovery modes. Thus, both categories of knowledge are necessary and neither is sufficient for assigning species status. This leads naturally to a hierarchical research program in historical ecology, beginning with phylogenetic analysis of a group of entities postulated to be evolutionary species, which provides a productive arena for our arguments about species concepts. Key words: Evolutionary Species Concept, historical ecology, Phylogenetic Species Concept, phylog- eny, speciation. “Why is not all nature in confusion, in- isms grouped naturally into discrete species, stead of the species being, as we see them, species into genera, genera into families, well defined?” (Darwin, 1872:161). and so on until all living things were in- cluded in a single classification of life. Pre- AShort History of the Darwinian taxonomists, like all natural phi- “Species Problem” losophers of the day, argued that the only “real” entities were those that had immu- It is not uncommon for major scientific table spatio-temporal existence. Because of concepts to have ambiguous definitions. In their unchangeable nature, such bits of re- Biology, one such term is “species.” This ality could be grouped into “classes” de- dilemma pre-dates Darwin (Stevens, 1992). fined by the fixed properties of their com- Initially, the “species” was simply a category ponents. Classic examples of such “real” in classifications, the least inclusive group of entities, which at the time were called “spe- organisms to be recognized formally. That cies,” are “hydrogen” and “gold.” Like the biosphere was organized into an in- these other natural kinds, biological species ternested hierarchy of species and groups of existed and conformed to a single hierarchi- species was thought to be self-evident. It was cal classification simply because it pleased simply the nature of the world that organ- the Creator for it to be so. In the absence of any causal principles explaining the taxo- Received for publication 20 November 1998. nomic hierarchy linking species together, 1 Symposium paper presented at the 37th Annual Meeting of the Society of Nematologists, 20–24 July 1998, St. Louis, MO. pre-Darwinian taxonomists relied on aes- 2 Centre for Comparative Biology and Biodiversity, Depart- thetics, intuition, and personal judgement ment of Zoology, University of Toronto, Toronto, Ontario M5S 3G5, Canada. to make decisions. E-mail: [email protected], mclennan@zoo. Darwin (1859) complicated things by sug- utoronto.ca Development of the perspective presented herein has been gesting that while organisms could indeed helped substantially by an enormous amount of published lit- be grouped naturally into species, these bio- erature appearing during the past 25 years, in addition to dis- cussions with Kare Bremer, John Collier, Joel Cracraft, Michael logical “species” were not immutable. Dar- Donoghue, Cyril Finnegan, Darryl Frost, William Fink, Vicki win’s belief in both the reality and the evo- Funk, Robert Geesink, David Hull, Arnold Kluge, Diedel Kor- net, Brian Maurer, Richard Mayden, Jack Maze, Brent Mishler, lutionary nature of biological species was Stan Salthe, Geoffrey Scudder, Jonathan Smith, Hans-Erik based on several empirical observations. Wanntorp, Quentin Wheeler, E. O. Wiley, and Rino Zandee. The authors especially thank Virginia Ferris for the invitation to First, comparing fossil and living species sug- participate in the symposium on species concepts. This study gested that the species populating the was supported by operating grants from the Natural Sciences and Engineering Research Council (NSERC) of Canada. planet today are not the same as those popu- This paper was edited by E. C. Bernard. lating the planet in the past. Second, the 117 118 Journal of Nematology, Volume 31, No. 2, June 1999 world today is not populated by a small num- Darwin’s comments: “Hence, in determin- ber of species widely distributed in many dif- ing whether a form should be ranked as a ferent habitats in many different parts of the species or a variety, the opinion of natural- world; different species live in different parts ists having sound judgement and wide expe- of the world. And third, all living and extinct rience seems the only guide to follow. We species fit into a single hierarchical classifi- must, however, in many cases, decide by a cation that looks like a genealogy. Using majority of naturalists, for well-marked and perhaps the original form of the “Wiley Cri- well-known varieties can be named which terion” (Brooks and McLennan, 1991), Dar- have not been ranked as species by at least win suggested that if it looked like a geneal- some competent judges” (Darwin, 1872). ogy, it was one. Such concerns still preoccupy the minds This conclusion led Darwin to his first of some taxonomists (Grant, 1985). In the theory—that all species on this planet were second half of the 20th century, evolutionary related to each other through a single com- taxonomy (Mayr, 1969) epitomized the pre- mon history of descent with modification: Darwinian view that taxonomy was partly sci- phylogeny. It is not mere happenstance that ence and partly art form, with a sizeable the only illustration Darwin ever placed in component of personal judgement and in- any edition of Origin of Species was a phylo- tuition involved in trying to justify taxo- genetic tree (Bowler, 1996). Darwin had cre- nomic decisions in terms of evolutionary ated a dual role for species in biology. In principles. Phenetics, or numerical tax- addition to being units of classification, spe- onomy (Sneath and Sokal, 1973; Levins, cies were now also units of evolutionary pro- 1979), by contrast, abandoned all efforts to cess; identifying and classifying species was provide a link between taxonomy and evo- not simply a matter of convenience, it was lution by advocating purely empirical classi- explanatory. From the statements about spe- fication procedures having no particular cies in Origin of Species, it is clear that Darwin connection with evolutionary principles. recognized this duality. Biologists have at- Hull (1965) presciently described taxonomy tempted to resolve this dualism in a variety at this time as the inheritor of 2,000 years of of ways during the past 140 years, but it still stasis resulting from the notion that all units exists. For example, Ereshevsky (1992) of classification had to have uniquely defin- stated “Biological systematists attempt to ing (and hence unchanging) characters, an provide a taxonomy of the world’s organic indication that the Darwinian revolution still diversity. Evolutionary biologists attempt to had not penetrated deeply into taxonomic explain why that diversity exists.” practice. Following the publication of Origin of Spe- Other biologists have dealt with Darwin’s cies, taxonomists, by and large, continued to dualism by attempting to replace the pre- concentrate on species as units of classifi- Darwinian taxonomic concept of species cation and tended to ignore evolutionary with one based strictly in evolutionary principles, invoking these principles only to mechanisms or by trying to eliminate the justify taxonomic decisions made follow- species concept altogether. Darwin wrote ing traditional