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Given the extraordinary diversity of life on the planet there must be mechanisms for speciation     : the formation of two species from one original species. Darwin envisioned this process as a branching event and diagrammed the process in the only illustration found in On the Origin of Species ( [link] a ). Compare this illustration to the diagram of elephant evolution ( [link] b ), which shows that as one species changes over time, it branches to form more than one new species, repeatedly, as long as the population survives or until the organism becomes extinct.

Image (a) shows a sketch of lines branching into a tree shape. At the bottom are 11 vertical lines labeled A through L. These then are branched out as they move up across the page through fourteen rows labeled with Roman numerals. Some branches make a straight line from the bottom row to the top row, others keep branching out further at each row, and some are straight partway through the rows until they connect to an existing branch or form no connection and instead stop. The top four rows each consists of a single line from a branch tip (there are 6 branch tips at row XI) to one of 15 individual final designations. Illustration B shows the evolution of modern African and Asian elephants from a common ancestor, the Palaeomastodon. The Palaeomastodon was similar to modern elephants; however, it was smaller and had a long nose instead of a trunk. Side branches of the elephant evolutionary tree gave rise to mastodons and mammoths. The mammoth is more closely related to modern elephants than the mastodon.
The only illustration in Darwin's On the Origin of Species is (a) a diagram showing speciation events leading to biological diversity. The diagram shows similarities to phylogenetic charts that are drawn today to illustrate the relationships of species. (b) Modern elephants evolved from the Palaeomastodon , a species that lived in Egypt 35–50 million years ago.

For speciation to occur, two new populations must be formed from one original population and they must evolve in such a way that it becomes impossible for individuals from the two new populations to interbreed. Biologists have proposed mechanisms by which this could occur that fall into two broad categories. Allopatric speciation (allo- = "other"; -patric = "homeland") involves geographic separation of populations from a parent species and subsequent evolution. Parapatric speciation     (para- = "beside"; -patric = "homeland") involves speciation occurring between populations that share a border, and may occasionally interbreed with each other. Sympatric speciation (sym- = "same"; -patric = "homeland") involves speciation occurring within a parent species remaining in one location.

Biologists think of speciation events as the splitting of one ancestral species into two descendant species. There is no reason why there might not be more than two species formed at one time except that it is less likely and multiple events can be conceptualized as single splits occurring close in time.

17.4d allopatric speciation

A geographically continuous population has a gene pool that is relatively homogeneous. Gene flow, the movement of alleles across the range of the species, is relatively free because individuals can move and then mate with individuals in their new location. Thus, the frequency of an allele at one end of a distribution will be similar to the frequency of the allele at the other end. When populations become geographically discontinuous, that free-flow of alleles is prevented. When that separation lasts for a period of time, the two populations are able to evolve along different trajectories. Thus, their allele frequencies at numerous genetic loci gradually become more and more different as new alleles independently arise by mutation in each population. Typically, environmental conditions, such as climate, resources, predators, and competitors for the two populations will differ causing natural selection to favor divergent adaptations in each group.

Isolation of populations leading to allopatric speciation can occur in a variety of ways: a river forming a new branch, erosion forming a new valley, a group of organisms traveling to a new location without the ability to return, or seeds floating over the ocean to an island. The nature of the geographic separation necessary to isolate populations depends entirely on the biology of the organism and its potential for dispersal. If two flying insect populations took up residence in separate nearby valleys, chances are, individuals from each population would fly back and forth continuing gene flow. However, if two rodent populations became divided by the formation of a new lake, continued gene flow would be unlikely; therefore, speciation would be more likely.

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Source:  OpenStax, General biology part i - mixed majors. OpenStax CNX. May 16, 2016 Download for free at http://legacy.cnx.org/content/col11749/1.5
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