Where's the Species?

Comments on the Phylogenetic Species Concepts*

MARC ERESHEFSKY
Dept. of Philosophy University of Wisconsin Madison, W153706 U.S.A.

Dissatisfaction with the Biological Species Concept has for some time been an impetus for biologists to propose new and alternative species definitions (for example, Simpson's (1961) Evolutionary Species Concept and van Valen's (1976) Ecological Species Concept). More recently, proposals for two new species definitions, coming under the same name the Phylogenetic Species Concept, appear in the pages of this journal (Cracraft 1987, Mishler and Brandon 1987). I am sympathetic to the criticisms launched at the Biological Species Concept, especially those given by Mishler and Donoghue (1982) and Mishler and Brandon (1987). However, it is important to consider what the Phylogenetic Species Concepts have to offer in its place. Though I think these proposed concepts nicely avoid some of the difficulties which face the Biological Species Concept, I doubt that either concept offers any new positive description of what species are. I will begin my comments with Cracraft's formulation of the Phylogenetic Species Concept, then I will turn to Mishler and Brandon's version. After a critical discussion of the Biological Species Concept, Cracraft (1978) introduces his Phylogenetic Species Concept as follows:
.. biology is faced with formulating an alternative species definition. Although this has been a persistently troublesome task, the previous discussion suggests a deceptively simple solution: equate species with evolutionary units. Accordingly, species can be defined as an irreducible cluster of organisms, within which there is a parental pattern of ancestry and descent, and which is diagnosably distinct from other such clusters. Species are thus basal, differentiated taxa (p. 341; see also Cracraft 1983, p. 170).

In brief, Cracraft thinks we should define species as evolutionary units, and he takes evolutionary units to be the "smallest diagnosable clusters" (1983, p. 179) of organisms which are connected by parent-offspring relations. A question immediately arises from this description of species: What is a "smallest diagnosable cluster" of organisms, "diagnosably distinct from other such clusters"? According to Cracraft, there are two criteria which
Biology and Philosophy 4 (1989) 89-96. 1989 by KluwerAcademic Publishers.

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can be used for individuating such taxa: "(1) one or more diagnostic characters (of any kind as long as they are heritable), and (2) reproductive cohesion" (1987, pp. 341-342). Of these criteria, only the first is necessary; the second is useful "because the diagnostic characters of a species may be restricted to a single sex, ontogenetic stage or morph" (1987, p. 342). Thus species are groups of genealogically connected organisms which share at least one diagnostic character. Furthermore, that character may be, but need not be, the ability to interbreed, or following Paterson (1985), the ability to recognize each other as mates. If we take Cracraft's description of species literally, then it implausibly follows that every group of genealogically connected organisms which share a single distinctive heritable trait is a species. For example, according to Cracraft's definition, every time Dobzhansky and his co-workers successfully introduced a new and heritable mutation into an interbreeding group of Drosophila, they created a new species. And, according to Cracraft's definition, the group of organisms Homo sapiens consists of at least thirty different species (corresponding to the thirty races of Homo sapiens Mayr speaks of (1970, p. 396)) and probably many many more. While there may be reasons for recognizing these groups as some sort of taxonomic units, it seems implausible to assert that each is a distinct species. Furthermore, from the description given above, we see that a species is "defined as an irreducible cluster of organisms, within which there is a parental pattern of ancestry and descent, and which is diagnosably distinct from other such clusters" (Cracraft 1987, p. 341; my italics). Earlier in his paper Cracraft writes, "Given a definition of species, irreducibility implies that entities meeting this definition cannot be subdivided into other, smaller entities which themselves play the role of species within theories having them as their entities" (1987, pp. 333-334). If, as Cracraft asserts, the role of species in evolutionary theory is to be the most basal groups of genealogically connected organisms which share at least one distinctive trait, and a species cannot be subdivided into smaller such groups, then it follows that only groups of genealogically connected organisms which share a single distinctive trait are species. So, according to Cracraft's species concept and his notion of irreducibility, the group Homo sapiens, for example, cannot be a species because it can be subdivided into smaller genealogical clusters of organisms which share a distinctive trait. The same consideration applies to almost any other group of organisms which biologists usually take to be a species, or even a subspecies. On the one hand, Cracraft's description of species asserts, quite implausibly, that taxa below the level of race are species. On the other hand, it excludes most if not all groups of organisms which are usually taken to be species from being species. Cracraft may counter that my portrayal of what species are by his

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Phylogenetic Species Concept is not what he means by species. This may be the case since I suspect that Cracraft thinks that species are what many biologists now take to be subspecies (see Cracraft 1983, p. 173). Nevertheless, his description does not capture this. If that is what Cracraft takes species to be, then he needs to revise his description. Another criticism I have of Cracraft's Phylogenetic Species Concept concerns what his concept does not do. His definition is entirely a claim about what phylogenetic groups or patterns of organisms constitute species; it gives no information about the processes or causes which produce and maintain such groupings. Nevertheless, Cracraft writes,
One definition is preferable to another because it allows us to understand nature better, that is, within the context of theories and hypotheses about biological patterns and processes, that definition lends more coherence and explanatory extensibility to those conjectures (1987, p. 331).

Since Cracraft's definition of species makes no reference to any processes which make species what they are, I do not see how his definition can lend itself to any "explanatory extensibility" concerning species phenomena. To make this point clearer, consider the Biological Species Concept of Mayr (1970, p. 12): "Species are groups of interbreeding natural populations that are reproductively isolated from other such groups." Undoubtedly this definition of species has its problems, but at least it identifies processes which might help explain certain species phenomena. For instance, the processes of gene flow and reproductive isolation, if truly effective, can be cited to explain why species maintain their unity. And citing the lack of gene flow between the subpopulations of a species can provide an account of how speciation occurs. I agree with Cracraft (1987, p. 339) that a major problem with the Biological Species Concept is that the processes it specifies are simply not found in many groups of organisms thought to be species by many biologists. But at least that definition posits some processes which if present and truly effective could help explain certain species phenomena. Cracraft's Phylogenetic Species Concept, on the other hand, is only a pattern claim about species; according to that concept, species are the smallest diagnosable clusters of genealogically connected organisms. Undoubtedly stipulating that species are such clusters does imply which groups of organisms are and are not species. But this does not help explain, for example, how species maintain their unity or how speciation occurs. In other words, Cracraft's Phylogenetic Species Concept provides no insight into why species are species; or more specifically, it provides no insight into how species maintain and lose the unity which we perhaps somewhat vaguely associate with species. Providing some such insight into the nature of species seems to be an important desideratum of any species concept. Because Cracraft's definition fails to provide any such insight, it

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fails Cracraft's own criterion for a preferable species concept - it lacks "explanatory extensibility" (1987, p. 331). My final point concerning Cracraft's Phylogenetic Species Concept is not so much a criticism of the concept, but a criticism of how Cracraft argues for it. After reviewing the Biological Species Concept, Cracraft introduces his Phylogenetic Species Concept as follows: "the previous discussion suggests a deceptively simple solution: equate species with evolutionary units" (1987, p. 341). This would be a simple solution if we had a clear idea of what an evolutionary unit is. But, as Cracraft is well aware (1983, p. 163), there is a wide range of opinions on what groups of biological entities are evolutionary units. For example, Cracraft and Eldredge (1980, pp. 89-90) and Ghiselin (1987, p. 137) maintain that evolutionary units must be reproductive units.l Hull (1980, pp. 327-329) and Rosenberg (1985, p. 139) hold that evolutionary units are lineages: groups of organisms, and perhaps other biological entities, connected by parent-offspring relations. Mary Williams (1985, pp. 584-585) seems to think that evolutionary units are groups of genealogically connected organisms which are exposed to the same selection forces. And Bunge (1981, p. 284) asserts that evolutionary units are "biopopulations." Given the diversity of opinions among biologists and philosophers as to what an evolutionary unit is, equating species with evolutionary units is not a "deceptively simple solution" to the species problem. It is just replacing one problem with another problem. Let us turn to the Phylogenetic Species Concept suggested by Mishler and Brandon (1987). This concept was first introduced in Mishler and Donoghue (1982) and is summarized as follows:
A species is the least inclusive taxon recognized in a classification, into which organisms are grouped because of degree of monophyly (usually, but not restricted to, the presence of synapomorphies), that is ranked as a species because it is the smallest "important" lineage deemed worthy of formal recognition, where "important" refers to the action of those processes that are dominant in producing and maintaining lineages in a particular case (1987, p. 46).

This definition places two requirements on species. First, species must be monophyletic groups or lineages of organisms. Second, such lineages must be important in the sense that they are produced and maintained by such processes as (but not limited to) actual interbreeding, selective constraints and developmental canalization (Mishler and Brandon 1987, p. 37). Mishler and Brandon, following Mishler and Donoghue (1982), call the first requirement a "grouping" criterion, and the second a "ranking" criterion. They maintain that each criterion is necessary, and together they are sufficient, for a group of organisms to be a species. I have comments to make concerning both these criteria. Suppose we have a group of organisms and we know that they form a

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monophyletic group. The group thus passes the "grouping" requirement of Mishler and Brandon's definition of species. What, according to this definition, would make that group a species? From the above description, we see that it must be an "important" lineage. But what makes it an "important" lineage? According to the above description, it is those processes that are dominant in producing and maintaining such lineages. Thus an "important" lineage is a lineage which is produced and maintained by those processes which produce and maintain "important" lineages. I submit that the "ranking" criterion of the concept being promoted by Mishler and Brandon is circular; and because it is circular it does not give any way to distinguish species from 6ther monophyletic taxa. In other words, the Phylogenetic Species Concept of Mishler and Brandon does not tell us what species are, other than that they are monophyletic lineages. Mishler and Brandon might reply that their concept gives more information than that about species because it refers to those processes which may cause a monophyletic group to be a species. This is true, but this information still does not tell us what a species or an "important" lineage is. Here is an analogy: Suppose you do not know what chairs are and you want to learn; I tell you that a chair is something which can be made with a saw and hammer, with a metal cutter, or by weaving reeds together. I even demonstrate how each process can cause the existence of a chair by building chairs with each method. Do you now know what a chair is? I do not think so. Being told that each one of these processes, among others, can cause the'existence of a chair does not tell you what a chair is. You need to know what it is about these resultant objects, caused by various processes, which makes them chairs. For instance, you need to know that they are devices on which people can sit, or some such description of chairs. According to Mishler and Brandon's "ranking" criterion, species are monophyletic lineages produced and maintained by gene flow, selective constraints, developmental canalization, or some other process which produces and maintains species. Each one of these processes may be important in causing and maintaining the existence of particular species. But still, we do not know why the lineages caused and maintained by such processes are species. In other words, we do not know what it is about these lineages which makes them species. Since Mishler and Brandon's "ranking" criterion does not tell us why such lineages are species, other than that these lineages are produced and maintained by those processes which cause lineages to be species, it leaves us in the dark as to what species are. What about their "grouping" criterion? According to Mishler and Brandon's species concept, all species must be monophyletic. But, as Mishler and Brandon point out, the cladistic definition of monophyly

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(the one they support) is not usually applied at the species level (1987, pp. 46-47). To remedy this, they suggest the following definition of monophyly:
A monophyletic taxon is a group that contains all and only descendents of a common ancestor, originating in a single event (1987, p. 47).

Is it plausible to assume that all species satisfy this definition of monophyly? Suppose A is a distinctive taxon because of a cluster of unique synapomorphies which originated at event x (see figure 1). A is a large, 'healthy' species within which there is much interbreeding. Its members are exposed to a common selection regime and they contain similar homeostatic genotypes. Some years after event x, A gives rise to a peripheral isolate, B. The members of B interbreed, are exposed to their own selection regime and sooner or later give rise to their own unique set of synapomorphies at event y. A, meanwhile, is unaffected by the establishment of B. According to Mishler and Brandon's Phylogenetic Species Concept, B is a species. I have referred to A as a species, and I think that this is fairly uncontroversial. 2 The problem for Mishler and Brandon's species concept is that A does not satisfy their definition of monophyly: A does not contain all and only the descendants of event x; some of the descendants of event x are contained in B. Mishler and Brandon could reply that A is not a species. But this seems implausible, for they would have to assert that there are no cases of species which continue to exist after some of their members give rise to a new and different species. Nevertheless, suppose they follow Hennig (1966) and assert that no parental species survives a speciation event. In other words, suppose that A is not one but two species, one originating at event x and the other at event y. This maneuver, however, does not escape the present problem because the first species in A fails to satisfy Mishler and Brandon's definition of monophyly: the first species in A does not contain all and only the descendants of event x; some of the descendants
A B

Figure 1. A phylogenetic tree, where clusters of synapomorphies are shown as cross-bars. See text for discussion.

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of event x are in B and some are in the later species of A. Regardless of whether one adopts Hennig's approach or not, there seem to be obvious cases of species which do not satisfy Mishler and Brandon's definition of monophyly. There are thus species which do not satisfy the "grouping" criterion of their Phylogenetic Species Concept. As a result of this, Mishler and Brandon's species concept does not plausibly apply to all species. Both Cracraft (1987) and Mishler and Brandon (1987) include a discussion of the problems facing the Biological Species Concept. For the most part, each of their Phylogenetic Species Concepts avoids these problems. But what do these concepts have to offer in place of the Biological Species Concept? On the one hand, Cracraft's concept implausibly asserts that only the most basal groups of organisms, those genealogically connected and which share at least one distinctive heritable trait, are species. Furthermore, Cracraft's concept is merely a pattern claim about species, providing no further insight into the nature of species. On the other hand, Mishler and Brandon's concept implausibly asserts that all species are monophyletic. And even if we assumed that all species were monophyletic, their concept does not in a non-circular fashion distinguish species from other monophyletic taxa. Some may find the Phylogenetic Species Concepts a step in the right direction because these concepts avoid some of the problems of the Biological Species Concept. I agree. But both definitions require revision and supplementation if they are to be fully adequate. Adopting a popular American expression, I conclude that both definitions need to more adequately address the question in the title of this paper.
NOTES * The author thanks Joel Cracraft, Brent Mishler and David Hull for commenting on earlier drafts of this paper, and Elliott Sober for his constant help and encouragement. ' For Cracraft and Eldredge (1980, p. 89), reproductive units are groups of organisms "among which there is at least occasional sexual reproduction." Whereas, for Ghiselin, reproductive units are groups of organisms among which there is "reproductive competition" between their organisms (1974, p. 538). 2 One author who disagrees with this is Hennig (1966). See text below for a consideration of Hennig's approach.

REFERENCES Bunge, M.: 1981, 'Biopopulations, Not Biospecies, Are Species and Evolve', Behavioral
and Brain Sciences 4, 284-285.

Cracraft, J.: 1983, 'Species Concepts and Speciation Analysis', Current Ornithology 1, 159-187.

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Cracraft, J.: 1987, 'Species Concepts and the Ontology of Evolution', Biology and Philosophy 2, 329-346. Eldredge, N. and Cracraft, J.: 1980, Phylogenetic Patterns and the Evolutionary Process, Columbia University Press, New York. Ghiselin, M.: 1974, 'A Radical Solution to the Species Problem', Systematic Zoology 23, 536-544. Ghiselin, M.: 1987, 'Species Concepts, Individuality, and Objectivity', Biology and Philosophy 2, 127-143. Hennig, W.: 1966, Phylogenetic Systematics, University of Chicago Press, Urbana. Hull, D.: 1980, 'Individuality and Selection', Annual Review of Ecology and Systematics 11,311-332. Mayr, E.: 1970, Populations, Species, and Evolution, Harvard University Press, Cambridge. Mishler, B. and Donoghue, M.: 1982, 'Species Concepts: A Case for Pluralism', Systematic Zoology 31,491-503. Mishler, B. and Brandon, R.: 1987, 'Individuality, Pluralism, and the Phylogenetic Species Concept', Biology and Philosophy 2, 37-54. Paterson, H.: 1985, 'The Recognition Concept of Species', TransvaalMuseum Monograph 4, 21-29. Rosenberg, A.: 1985, The Structure of Biological Science, Cambridge University Press, New York. Simpson, G.: 1961, The Principles of Animal Taxonomy, Columbia University Press, New York. van Valen, L.: 1976, 'Ecological Species, Multispecies, and Oaks', Taxon 25,233-239. Williams, M.: 1985, 'Species are Individuals: Theoretical Foundations for the Claim', Philosophyof Science 52, 578-590.