The problem with the species problem

When Charles Darwin convinced the scientific community that species evolve, the long-held essentialist view of each species as fixed was rejected and a clear conceptual understanding of the term was lost. For the next century, a real species problem existed that became culturally entrenched within the scientific community. Although largely solved decades ago, the species problem remains entrenched today due to a suite of factors. Most of the factors that help maintain its perceived intractability have been revealed and logically dismissed; yet this is not widely known so those factors continue to be influential. It is time to recognize this false foundation and relegate the species problem to history.     

Genomics and the bacterial species problem

Whether or not bacteria have species is a perennially vexatious question. Given what we now know about variation among bacterial genomes, we argue that there is no intrinsic reason why the processes driving diversification and adaptation must produce groups of individuals sufficiently coherent in their genetic and phenotypic properties to merit the designation 'species' - although sometimes they might.     

Numerical approaches to the species problem

The use of numbers by systematists is not new. Measurements to describe individuals and formal taxa have been used since the beginnings of our science. But the advent of electronic computers now permits a much more accurate understanding of the phenotypic relationships within and among populations and taxa. Furthermore, estimates of cladistic relationship also are being attempted with the help of computers. Computers can increase our understanding of speciation, but this requires us to think intelligently about the meaning of their results.Presented at the symposium Speciation and the Species Concept during the XIIth International Botanical Congress, Leningrad, July 8, 1975.     

The mind of the species problem

The species problem is the long-standing failure of biologists to agree on how we should identify species and how we should define the word 'species'. The innumerable attacks on the problem have turned the often-repeated question 'what are species?' into a philosophical conundrum. Today, the preferred form of attack is the well-crafted argument, and debaters seem to have stopped inquiring about what new information is needed to solve the problem. However, our knowledge is not complete and we have overlooked something. The species problem can be overcome if we understand our own role, as conflicted investigators, in causing the problem.     

Long-term persistence of species and the SLOSS problem

The single large or several small (SLOSS) problem has been addressed in a large number of empirical and theoretical studies, but no coherent conclusion has yet been reached. Here I study the SLOSS problem in the context of metapopulation dynamics. I assume that there is a fixed total amount A(0) of habitat available, and I derive formulas for the optimal number n and area A of habitat patches, where n=A(0)/A. I consider optimality in two ways. First, I attempt to maximize the time to metapopulation extinction, which is a relevant measure for metapopulation viability for rare and threatened species. Second, I attempt to maximize the metapopulation capacity of the habitat patch network, which corresponds both with maximizing the distance to the deterministic extinction threshold and with maximizing the fraction of occupied patches. I show that in the typical case, a small number of large patches maximizes the metapopulation capacity, while an intermediate number of habitat patches maximizes the time to extinction. The main conclusion stemming from the analysis is that the optimal number of patches is largely affected by the relationship between habitat patch area and rates of immigration, emigration and local extinction. Here this relationship is summarized by a single factor zeta, termed the patch area scaling factor.     

The cladistic solution to the species problem

The correct explanation of why species, in evolutionary theory, are individuals and not classes is the cladistic species concept. The cladistic species concept defines species as the group of organisms between two speciation events, or between one speciation event and one extinction event, or (for living species) that are descended from a speciation event. It is a theoretical concept, and therefore has the virtue of distinguishing clearly the theoretical nature of species from the practical criteria by which species may be recognized at any one time. Ecological or biological (reproductive) criteria may help in the practical recognition of species. Ecological and biological species concepts are also needed to explain why cladistic species exist as distinct lineages, and to explain what exactly takes place during a speciation event. The ecological and biological species concepts work only as sub-theories of the cladistic species concept and if taken by themselves independently of cladism they are liable to blunder. The biological species concept neither provides a better explanation of species indivudualism than the ecological species concept, nor, taken by itself, can the biological species concept even be reconciled with species individualism. Taking the individuality of species seriously requires subordinating the biological, to the cladistic, species concept.     

The problem of species revisited

Hypothetical-deductive analysis is an alternative to both induction and deduction, the difference between which is considered less important. The hypothetical-deductive approach requires a scientist to abandon the synthetic theory of evolution in favor of the epigenetic theory. This latter treats species as a discrete level of biodiversity and, particularly, the one where sympatric biparental groups show reproductive independence. In the case of uniparental (asexual and parthenogenetic groups), as well as in bisexual allopatric groups, the species level of divergence is identifiable by analogy to sympatric biparental relatives. The biological and evolutionary species concepts are disproved: the former contradicts observation, and the latter is non-operational.     

Resolution of the species problem in African trypanosomes

There is a general assumption that eukaryote species are demarcated by morphological or genetic discontinuities. This stems from the idea that species are defined by the ability of individuals to mate and produce viable progeny. At the microscopic level, where organisms often proliferate more by asexual than sexual reproduction, this tidy classification system breaks down and species definition becomes messy and problematic. The dearth of morphological characters to distinguish microbial species has led to the widespread application of molecular methods for identification. As well as providing molecular markers for species identification, gene sequencing has generated the data for accurate estimation of relatedness between different populations of microbes. This has led to recognition of conflicts between current taxonomic designations and phylogenetic placement. In the case of microbial pathogens, the extent to which taxonomy has been driven by utilitarian rather than biological considerations has been made explicit by molecular phylogenetic analysis. These issues are discussed with reference to the taxonomy of the African trypanosomes, where pathogenicity, host range and distribution have been influential in the designation of species and subspecies. Effectively, the taxonomic units recognised are those that are meaningful in terms of human or animal disease. The underlying genetic differences separating the currently recognised trypanosome taxa are not consistent, ranging from genome-wide divergence to presence/absence of a single gene. Nevertheless, if even a minor genetic difference reflects adaptation to a particular parasitic niche, for example, in Trypanosoma brucei rhodesiense, the presence of a single gene conferring the ability to infect humans, then it can prove useful as an identification tag for the taxon occupying that niche. Thus, the species problem can be resolved by bringing together considerations of utility, genetic difference and adaptation.     

A (not-so-radical) solution to the species problem

What are species? One popular answer is that species are individuals. Here I develop another approach to thinking about species, an approach based on the notion of a lineage. A lineage is a sequence of reproducing entities, individuated in terms of its components. I argue that one can conceive of species as groups of lineages, either organism lineages or population lineages. Conceiving of species as groups of lineages resolves the problems that the individual conception of species is supposed to resolve. It has added the virtue of focusing attention on the characteristic of species that is most relevant to understanding their role in evolutionary processes, namely, the lineage structure of species.     

On the nature of the species problem and the four meanings of 'species'

Present-day thought on the notion of species is troubled by a mistaken understanding of the nature of the issue: while the species problem is commonly understood as concerning the epistemology and ontology of one single scientific concept, I argue that in fact there are multiple distinct concepts at stake. An approach to the species problem is presented that interprets the term 'species' as the placeholder for four distinct scientific concepts, each having its own role in biological theory, and an explanation is given of the concepts involved. To illustrate how these concepts are commonly conflated, two widely accepted ideas on species are criticized: species individualism and species pluralism. I argue that by failing to distinguish between the four concepts and their particular roles in contemporary biological theory, these ideas stand in the way of a final resolution of the species problem.