Online research in philosophy

Sexually active male haplochromine cichlid fishes possess pronounced yellow ovoid spots on the anal fm, which mimic eggs of the female and have therefore been called egg dummies (Wickler, 1962b).It is thought that divergence in egg dummy characteristics can considerably reduce gene flow and in this way may trigger off reproductive isolation. Two ways in which egg dummy divergence can develop are described. Both mechanisms may have been operating, at the same time and in the same area, in different species, or possibly even in a single species.

Historians and philosophers of science agree that Darwin had an understanding of species which led to a workable theory of their origins. To Darwin species did not differ essentially from ‘varieties’ within species, but were distinguishable in that they had developed gaps in formerly continuous morphological variation. Similar ideas can be defended today after updating them with modern population genetics. Why then, in the 1930s and 1940s, did Dobzhansky, Mayr and others argue that Darwin failed to understand species and speciation? Mayr and Dobzhansky argued that reproductively isolated species were more distinct and ‘real’ than Darwin had proposed. Believing species to be inherently cohesive, Mayr inferred that speciation normally required geographic isolation, an argument that he believed, incorrectly, Darwin had failed to appreciate. Also, before the sociobiology revolution of the 1960s and 1970s, biologists often argued that traits beneficial to whole populations would spread. Reproductive isolation was thus seen as an adaptive trait to prevent disintegration of species. Finally, molecular genetic markers did not exist, and so a presumed biological function of species, reproductive isolation, seemed to delimit cryptic species better than character-based criteria like Darwin’s. Today, abundant genetic markers are available and widely used to delimit species, for example using assignment tests: genetics has replaced a Darwinian reliance on morphology for detecting gaps between species. In the 150th anniversary of The Origin of Species , we appear to be returning to more Darwinian views on species, and to a fuller appreciation of what Darwin meant.

Pierre Trémaux’s 1865 ideas on speciation have been unjustly derided following his acceptance by Marx and rejection by Engels, and almost nobody has read his ideas in a charitable light. Here we offer an interpretation based on translating the term sol as “habitat”, in order to show that Trémaux proposed a theory of allopatric speciation before Wagner and a punctuated equilibrium theory before Gould and Eldredge, and translate the relevant discussion from the French. We believe he may have influenced Darwin’s revision to the third edition of the Origin on rates of evolution, and suggest that Gould’s dismissal of Trémaux is motivated by concern that others might think punctuated equilibrium theory was tainted by a connection with Trémaux.

This paper shows that species are individuals with respect to evolutionary theory in the sense that the laws of the theory deal with species as irreducible wholes rather than as sets of organisms. 'Species X' is an instantiation of a primitive term of the theory. I present a sketch of a proof that it cannot be defined within the theory as a set of organisms; the proof relies not on details of my axiomatization but rather on a generally accepted property of speciation; hence the same argument should work for any axiomatization which captures this generally accepted property of speciation.

The purpose of this paper is to test the contemporary concept of biological species against some of the problems caused by treating species as spatiotemporally extended entities governed by criteria of persistence, identity, etc. After outlining the general problem of symmetric division in natural objects, I set out some useful distinctions (section 1) and confirm that species are not natural kinds (section 2). Section 3 takes up the separate issue of species definition, focusing on the Biological Species Concept (BSC). Sections 4 and 5 examine the matter of species identity over space and time respectively, as determined by the BSC. Both gradualistic and punctuated equilibrium models of speciation are discussed. In section 6 I argue that the BSC fails to determine adequate criteria for dealing with certain kinds of speciation. Section 7 moves speculatively beyond the BSC to a brief examination of alternatives.

Simple models for the evolution of qualitative multistate traits are considered, in which the traits are permitted to evolve in time-dependent versus speciation-dependent fashion. Of particular interest are the means and variances of distances for these traits in evolutionary phylads characterized by different rates of speciation, when alternative characters are neutral with respect to fitness, and when the total number of observable characters is limited to small values. As attainable character states are increasingly restricted, mean distance (D) in a phylad decreases, regardless of whether evolution is a function of time or of rate of speciation. The ratio of mean distances in species-rich and species-poor phylads of comparable evolutionary age (DR/DP) remains near one when differentiation is proportional to time, even when attainable character states are severely restricted. DR/DP also nears one as a result of restricting character states when differentiation is proportional to rate of speciation, but the effect is not severe unless the number of character states is very small and the probability of change per speciation very large. These and other results are discussed with reference to available data sets on qualitative multistate traits.

The crucial phase of speciation is argued to be the evolution of mating cross-incompatibility (prezygotic incompatibility) between the genotypes distinguishing the prospective species populations. Based on this idea, a single-locus model of speciation is presented, which is shown to be biologically plausible and may help to settle the controversy as to the biological significance of single-locus modes of speciation. The model involves three alleles, two of which characterize in homozygous state the prospective species populations and in heterozygous state their hybrids. The third allele represents a mutant which is equivalent to one of the first two alleles with the exception that it inhibits mating with carriers of the third allele. This third allele is fixed in one population and immigrates into a second population which contains the mutant inhibiting matings with members of the former population. Migration in the reverse direction does not occur. Proceeding from a widely applicable concept of fitness and mating preference it is shown that postzygotic incompatibility (hybrid or heterozygote disadvantage) alone suffices to trigger evolutionary replacement of the extant mating relations in the population receiving immigrants by any arbitrary degree of prezygotic incompatibility. This corroborates Wallace's hypothesis and emphasizes the potential biological relevance of speciation by reinforcement (parapatric speciation) at single gene loci.

Speciation is an aspect of evolutionary biology that has received little philosophical attention apart from articles mainly by biologists such as Mayr (1988). The role of speciation as a terminus a quo for the individuality of species or in the context of punctuated equilibrium theory has been discussed, but not the nature of speciation events themselves. It is the task of this paper to attempt to bring speciation events into some kind of general scheme, based primarily upon the work of Sergey Gavrilets on adaptive landscapes, using migration rate, or gene flow, as the primary scale, and concluding that adaptive and drift explanations are complementary rather than competing. I propose a distinction between intrinsic and extrinsic selection, and the notion of reproductive reach and argue that speciation modes should be discriminated in terms of gene flow, the nature of selection maintaining reproductive reach, and whether the predominant cause is selective or stochastic. I also suggest that the notion of an adaptive “quasispecies” for asexual species is the primitive notion of species, and that members of reproductively coherent sexual species are additionally coadapted to their mating partners.

The cladistic species concept proposed by Ridley (1989) rests on an undefined notion of speciation and its meaning is thus indeterminate. If the cladistic concept is made determinate through the definition of speciation, then it reduces to a form of whatever species concept is implicit in the definition of speciation and fails to be a truly alternative species concept. The cladistic formalism advocated by Ridley is designed to ensure that species are monophyletic, that they are objectively real entities, and that they are individuals. It is argued that species need not be monophyletic in order to be real entities, and that ancestor-descendant relations are not the only relations that confer individuality on entities. The species problem is recast in terms of a futile quest for a definition of that single kind of entity to which the term species should uniquely apply.

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.