he species category plays a central role in many areas of biology. In addition to systematics, the work of conservation biology, ecology, evolutionary biology, human biology, and molecular biology all ultimately involve species taxa and therefore depend upon some conception of what species are, how to delimit them, and how they persist over time. For example, species are indispensable in discussions concerning biological diversity and conservation. But it is hard to estimate the total number of extant species or to estimate the relative contributions of different groups of organisms to total species diversity without a sound conception of species (Claridge, Dawah & Wilson, 1997).

The reason that species are central to so many dif- ferent areas of biological inquiry involves their special status compared to other levels of the Linnaean hierar- chy. Subspecies, genera, families, orders, etc., are to some extent arbitrarily defined. But not species. Most biologists agree that species are real and not simply the by-product of human definition and classification schemes (Futuyma, 1986; Mayr, 1988). Instead, species are the result of complex historical and evolutionary processes: They come into existence, change, and then cease to exist. In short, species are the empirical traces left by the workings of evolution, and we cannot under- stand one without understanding the other.

Given that species are real and that they play an important role in many different areas of biology, it seems reasonable to expect biologists to agree on how to define the term \u201cspecies.\u201d After all, we use the term in everyday discourse with no problem: Children from var- ious cultures can readily identify species (Atran, 1999), non-specialists communicating in newspapers and other public forums readily use the term to discuss envi- ronmental issues, and biologists themselves use the term in their research.

Unfortunately, there is currently no consensus among biologists regarding the definition of the term \u201cspecies\u201d (Wilson, 1999). Instead, as we shall see shortly, there are anywhere from three to 22 definitions, and there is ongoing debate about which definition is the right one (Claridge, Dawah & Wilson, 1997). This, in a nutshell, is the species problem. As philosopher of biolo- gy David Hull summarizes the situation, \u201cWe are drown- ing in a sea of species concepts\u201d (Hull, 1999, p. 44).

Just labeling something a problem does not neces- sarily make it one. Why should it be a problem that there are a host of species definitions and no consensus among biologists? The worry is that the species catego- ry is such a basic element in the scheme of biology that not having a standard agreed-upon definition indicates something is wrong. Maybe we don\u2019t really understand species if we can\u2019t define the term, maybe there is some- thing fundamentally wrong with the conceptual frame- work of evolution that is hindering us from defining the term, or maybe, as some philosophers and biologists have begun to argue, there is something fundamentally wrong with the Linnaean hierarchy itself (Ereshefsky, 1999; Mishler, 1999).

What is the relevance of the species problem to the biology classroom? From the glimpse provided in the last few paragraphs, one might well conclude that the species problem is too hard, too complicated, or too controversial for introductory students to tackle. Nonetheless, I believe there are compelling reasons to thoughtfully and deliberately incorporate some ele- ments of the species problem into introductory courses.

First, biological science is not simply a heap of facts that need to be memorized. Biological facts are inter- connected and there is an overarching theoretical frame- work, evolution, that allows biologists to interpret, understand, and explain organic phenomena. However, because biological facts are interconnected, a poor grasp of key concepts will hinder students from under- standing other key concepts. So, for example, a superfi- cial understanding of species will hinder students from understanding evolution (Bishop & Anderson, 1990;

CARL CHUNG is Assistant Professor of Humanities at General College, University of Minnesota, Minneapolis, MN 55455; e-mail: chung023@umn.edu.

Halld\u00e9n, 1988). I believe that challenging students to grapple with some of the complexities inherent in the species problem will result in a more sophisticated con- ception of species and that this will help students understand other key concepts in biology.

Second, research shows that students come to their biology courses with a wide range of assumptions and misconceptions, especially about evolution (Lawson & Thompson, 1988; Bishop & Anderson, 1990; Lawson & Worsnop, 1992; Jensen & Finley, 1997; Moore, 2002). Such assumptions and misconceptions are hard to dis- lodge: Instead students learn to \u201ctalk the talk\u201d on exami- nations but then happily revert to more familiar ways of thinking after the course is over. To meet this challenge, biology teachers need to identify, analyze, and help stu- dents become aware of these assumptions and miscon- ceptions. In doing so, it will become clear that in trying to teach students evolution, biology teachers are in fact trying to teach students a whole new framework for understanding organic phenomena. Evolution is not sim- ply just another biological \u201cfact\u201d or \u201ctheory.\u201d Rather it is a worldview or paradigm that requires students to give up an alternative worldview or paradigm that they connect to traditional sources of authority (such as parents) and unexamined \u201ccommon sense.\u201d One way to help students begin to make such a transition is to start them thinking like a biologist\u2014not just learning passively what biologists say or think or do, but actively employing biological con- cepts on real problems. Again, I believe the complexities inherent in the ongoing debates surrounding the species problem offer one way to accomplish this.

What is the species problem? The species problem involves answering questions such as the following: What are species? Is there anything special about the species category that distinguishes it from other taxo- nomic categories? How do species taxa originate and how are they maintained? What role(s) should a species concept play in different areas of biology? Since the fate- ful publication of Darwin\u2019s On the Origin of Species in 1859, biologists have struggled to answer such ques- tions. The problem is not a lack of answers. Instead, as noted above, the problem is that the biological commu- nity cannot reach a lasting consensus on which answers it thinks are the right ones.

To illustrate this impasse, consider the following three attempts to identify and evaluate available species concepts. First, in 1942, Ernst Mayr discussed five such concepts in his ground-breaking book Systematics and

2. The practical species concept, based on the sub- jective judgment of any competent systematist (ibid., p. 115).

5. The biological species definition, based on the practical need of the taxonomist and incorporat- ing elements from the other concepts, reads as follows:

A species consists of a group of populations which replace each other geographically or eco- logically and of which the neighboring ones intergrade or interbreed wherever they are in contact or which are potentially capable of doing so (with one or more of the populations) in those cases where contact is prevented by geo- graphical or ecological barriers.

potentially interbreeding natural populations, which are reproductively isolated from other such groups.(ibid., pp. 119-120)

In his subsequent analysis, Mayr argues that the morphological approach based on overall similarity is outdated and unable to deal with intraspecific variation (ibid., pp. 109-111). He also argues that concept #2, #3, or #4 cannot serve as an adequate species concept by itself, thus leaving the biological species concept based on reproductive isolation as his clear choice.

But even Mayr\u2019s tireless and vociferous defense of the biological species concept for the last 50 plus years has not led to agreement. In 1997, David Hull and R. L. Mayden provided two additional analyses of available species concepts. Hull identifies seven workable species concepts: phenetic, biological, evolutionary, mate recog- nition, cohesion, monophyletic, and diagnostic (Hull, 1999, pp. 360-361). To evaluate them, Hull appeals to three general criteria that philosophers of science have stressed as important for scientific concepts: universality (or generality), applicability, and theoretical significance (ibid., p. 358). After evaluating each species concept against the criteria and assigning scores, Hull writes,

... the depressing result is that all seven species con- cepts score about the same. This outcome may well explain why the species controversy continues unabated. Even if everyone agreed on what counts as a good species concept, no one concept is clearly superior to all the others\u2014and not everyone agrees on what general criteria characterize a good species con- cept.(ibid., pp. 376-377)

Finally, a more complicated solution is offered by R. L. Mayden. In his analysis, Mayden identifies a whop- ping 22 species concepts (Mayden, 1997). Basically, Mayden accepts Hull\u2019s criteria as a basis for evaluation, with the addition of operationality as a fourth criterion. What complicates Mayden\u2019s discussion, however, is his use of a distinction between a single primary species concept and a host of secondary species concepts (this distinction, in a slightly different form, is due to Mayr, 1957; see Mayden, 1997). The primary concept is essen- tially theoretical; as such, it needs to be consistent with accepted knowledge of species and it needs to encom- pass all species phenomena. The secondary concepts then act as \u201cbridging concepts\u201d (ibid.) that allow biolo- gists to \u201crecognize entities compatible with [the primary concept\u2019s] intentions\u201d (ibid.). The secondary concepts are thus based upon specific aspects of species.

Mayden\u2019s conclusion is that only the evolutionary species concept, originally formulated by Simpson, can serve as the primary species concept. This concept defines a species as

... an entity composed of organisms which maintains its identity from other such entities through time and over space, and which has its own independent evo- lutionary fate and historical tendencies(Mayden,

All other species concepts are, for Mayden, second- ary concepts in the sense that they serve simply to sup- plement the primary evolutionary species concept with more specific details of species.

It is too early to determine whether Mayden\u2019s \u201cdefin- itive\u201d treatment of the species problem will convince tra- ditional critics of the evolutionary species concept. Critics may certainly dispute Mayden\u2019s reasons for adopt- ing the distinction between primary and secondary con- cepts in the first place, or his choice of the evolutionary species concept asthe primary concept, or even the assumption that there is only one primary concept instead of a collection of primary concepts. But then it seems possible that the species problem may simply reappear within the framework of Mayden\u2019s analysis.

For present purposes, the examples illustrate force- fully how three different attempts to address the species problem result in three different outcomes. Recent pub- lications on the species problem continue to offer multi- ple and divergent recommendations (Wilson, 1999). At present, then, there is no consensus on how to define the species category and there is no general agreement on an alternative resolution to the problem (but see Ghiselin, 1974; Ereshefsky, 1999; and de Quieroz, 1999 for addi- tional attempts at definitive solutions). Given this state of affairs, it is worth consideringwhy a conception of species that is both rigorous and generally acceptable to biologists is proving to be so elusive. By briefly reviewing

the advantages and shortcomings of the biological and the evolutionary species concepts, we can shed some light on what perpetuates the species problem.

The Biological & the
Evolutionary Species Concepts:
Pros & Cons

As noted earlier, the biological species concept emphasizes reproductive isolation as the crucial and defining property of a species (Mayr, 1942, 1963, 1993; Futuyma, 1986). As such, to determine whether two populations (or groups of populations) are distinct species, all one has to do is \u201cwait and see.\u201d If individuals from the populations interbreed, they belong to the same species; if individuals from the populations do not interbreed, they belong to different species. Because reproductive isolation amounts to the existence of bio- logical barriers to gene flow, the biological species con- cept is also said to define a species as a protected gene pool (Mayden, 1997, p. 391).

The biological species concept has a number of advantages that help account for its impressive \u201cstaying power\u201d (Coyne, 1994). For example, on the traditional morphological and more recent phenetic approaches, sibling species are hard to distinguish due to the empha- sis on overall similarity (Mayr, 1982). Conversely, such approaches would also tend to count as separate species those populations whose members exhibit any striking, systematic variation (ibid.). In each of these cases, how- ever, proponents claim that the biological species con- cept points the biologist in the right direction by stress- ing reproductive isolation, which in turn leads to a broad- er consideration of other biological factors such as eco- logical niche and mating behaviors (Mayr, 1982, 1993).

Another advantage is that the biological species concept readily connects to research programs in both micro- and macroevolutionary biology. It accomplishes this by (1) stressing that the species is the key unit of evolutionary change, and (2) providing an operational criterion (reproductive isolation) that can help inform specific research questions (Coyne, 1994; Gould, 1994). So, to cite one example, from the point of view of the biological species concept, \u201cThe origin of species is now seen as the origin of reproductive isolating mecha- nisms\u201d (Coyne, 1994, p. 22).

The problems facing the biological species concept, however, are also significant (for more complete lists of problems and references, see Coyne, 1994; Mayden, 1997; and de Queiroz, 1999). Two major problems arise due to the emphasis upon reproductive isolation. First, because the organisms under study can only be under- stood to form true species if they are sexually reproduc- ing, the biological species concept runs into trouble