Theory of speciation (18TH CENTURY)

Also called geographic speciation, this theory is most often associated with Ernst Mayr (1904- ), an evolutionary biologist at Harvard University (although many other biologists have theorized on the subject).

It asserts that new species arise among sexually reproducing organisms because geographic isolation enables a small subgroup to diverge genetically from the larger, established population.

This isolated subgroup may diverge owing to particular environmental pressures, mutations or GENETIC DRIFT. For this subgroup to become a new species, its unique characteristics must allow it to survive in its isolated location, and its gene pool must be protected from mixing with that of other species.

Also see: ALLOPARAPATRIC SPECIATION, ALLOPATRIC SPECIATION, COHESION SPECIES CONCEPT, GENETIC REVOLUTION, PARAPATRIC SPECIATION, SALTATION SPECIATION, species essentialism, STASIPATRIC SPECIATION, SYMPATRIC SPECIATION

Source:
M Ereshefsky, The Units of Evolution: Essays on the Nature of Species (Cambridge, Mass., 1992)

Historical background

In addressing the question of the origin of species, there are two key issues: (1) what are the evolutionary mechanisms of speciation, and (2) what accounts for the separateness and individuality of species in the biota? Since Charles Darwin’s time, efforts to understand the nature of species have primarily focused on the first aspect, and it is now widely agreed that the critical factor behind the origin of new species is reproductive isolation.^[5] Next we focus on the second aspect of the origin of species.

Darwin’s dilemma: Why do species exist?

In On the Origin of Species (1859), Darwin interpreted biological evolution in terms of natural selection, but was perplexed by the clustering of organisms into species.^[6] Chapter 6 of Darwin’s book is entitled “Difficulties of the Theory.” In discussing these “difficulties” he noted “Firstly, why, if species have descended from other species by insensibly fine gradations, do we not everywhere see innumerable transitional forms? Why is not all nature in confusion instead of the species being, as we see them, well defined?” This dilemma can be referred to as the absence or rarity of transitional varieties in habitat space.^[7]

Another dilemma,^[8] related to the first one, is the absence or rarity of transitional varieties in time. Darwin pointed out that by the theory of natural selection “innumerable transitional forms must have existed,” and wondered “why do we not find them embedded in countless numbers in the crust of the earth.” That clearly defined species actually do exist in nature in both space and time implies that some fundamental feature of natural selection operates to generate and maintain species.^[6]

Effect of sexual reproduction on species formation

It has been argued that the resolution of Darwin’s first dilemma lies in the fact that out-crossing sexual reproduction has an intrinsic cost of rarity.^{[9][10][11][12][13]} The cost of rarity arises as follows. If, on a resource gradient, a large number of separate species evolve, each exquisitely adapted to a very narrow band on that gradient, each species will, of necessity, consist of very few members. Finding a mate under these circumstances may present difficulties when many of the individuals in the neighborhood belong to other species. Under these circumstances, if any species’ population size happens, by chance, to increase (at the expense of one or other of its neighboring species, if the environment is saturated), this will immediately make it easier for its members to find sexual partners. The members of the neighboring species, whose population sizes have decreased, experience greater difficulty in finding mates, and therefore form pairs less frequently than the larger species. This has a snowball effect, with large species growing at the expense of the smaller, rarer species, eventually driving them to extinction. Eventually, only a few species remain, each distinctly different from the other.^[9][10][12] The cost of rarity not only involves the costs of failure to find a mate, but also indirect costs such as the cost of communication in seeking out a partner at low population densities.

African pygmy kingfisher, showing coloration shared by all adults of that species to a high degree of fidelity.^[14]

Rarity brings with it other costs. Rare and unusual features are very seldom advantageous. In most instances, they indicate a (non-silent) mutation, which is almost certain to be deleterious. It therefore behooves sexual creatures to avoid mates sporting rare or unusual features (koinophilia).^[15][16] Sexual populations therefore rapidly shed rare or peripheral phenotypic features, thus canalizing the entire external appearance, as illustrated in the accompanying illustration of the African pygmy kingfisher, Ispidina picta. This uniformity of all the adult members of a sexual species has stimulated the proliferation of field guides on birds, mammals, reptiles, insects, and many other taxa, in which a species can be described with a single illustration (or two, in the case of sexual dimorphism). Once a population has become as homogeneous in appearance as is typical of most species (and is illustrated in the photograph of the African pygmy kingfisher), its members will avoid mating with members of other populations that look different from themselves.^[17] Thus, the avoidance of mates displaying rare and unusual phenotypic features inevitably leads to reproductive isolation, one of the hallmarks of speciation.^{[18][19][20][21]}

In the contrasting case of organisms that reproduce asexually, there is no cost of rarity; consequently, there are only benefits to fine-scale adaptation. Thus, asexual organisms very frequently show the continuous variation in form (often in many different directions) that Darwin expected evolution to produce, making their classification into “species” (more correctly, morphospecies) very difficult.^{[9][15][16][22][23][24]}

Modes

Comparison of allopatric, peripatric, parapatric and sympatric speciation

All forms of natural speciation have taken place over the course of evolution; however, debate persists as to the relative importance of each mechanism in driving biodiversity.^[25]

One example of natural speciation is the diversity of the three-spined stickleback, a marine fish that, after the last glacial period, has undergone speciation into new freshwater colonies in isolated lakes and streams. Over an estimated 10,000 generations, the sticklebacks show structural differences that are greater than those seen between different genera of fish including variations in fins, changes in the number or size of their bony plates, variable jaw structure, and color differences.^[26]

Allopatric

During allopatric (from the ancient Greek allos, “other” + patrā, “fatherland”) speciation, a population splits into two geographically isolated populations (for example, by habitat fragmentation due to geographical change such as mountain formation). The isolated populations then undergo genotypic or phenotypic divergence as: (a) they become subjected to dissimilar selective pressures; (b) they independently undergo genetic drift; (c) different mutations arise in the two populations. When the populations come back into contact, they have evolved such that they are reproductively isolated and are no longer capable of exchanging genes. Island genetics is the term associated with the tendency of small, isolated genetic pools to produce unusual traits. Examples include insular dwarfism and the radical changes among certain famous island chains, for example on Komodo. The Galápagos Islands are particularly famous for their influence on Charles Darwin. During his five weeks there he heard that Galápagos tortoises could be identified by island, and noticed that finches differed from one island to another, but it was only nine months later that he reflected that such facts could show that species were changeable. When he returned to England, his speculation on evolution deepened after experts informed him that these were separate species, not just varieties, and famously that other differing Galápagos birds were all species of finches. Though the finches were less important for Darwin, more recent research has shown the birds now known as Darwin’s finches to be a classic case of adaptive evolutionary radiation