The scientific study of speciation — how species evolve to become new species — began around the time of Charles Darwin in the middle of the 19th century. Many naturalists at the time recognized the relationship between biogeography (the way species are distributed) and the evolution of species. The 20th century saw the growth of the field of speciation, with major contributors such as Ernst Mayr researching and documenting species' geographic patterns and relationships. The field grew in prominence with the modern evolutionary synthesis in the early part of that century. Since then, research on speciation has expanded immensely.
The language of speciation has grown more complex. Debate over classification schemes on the mechanisms of speciation and reproductive isolation continue. The 21st century has seen a resurgence in the study of speciation, with new techniques such as molecular phylogenetics and systematics. Speciation has largely been divided into discrete modes that correspond to rates of gene flow between two incipient populations. Current research has driven the development of alternative schemes and the discovery of new processes of speciation.
Early history
Charles Darwin introduced the idea that species could evolve and split into separate lineages, referring to it as specification in his 1859 book On the Origin of Species. It was not until 1906 that the modern term speciation was coined by the biologist Orator F. Cook. Darwin, in his 1859 publication, focused primarily on the changes that can occur within a species, and less on how species may divide into two. It is almost universally accepted that Darwin's book did not directly address its title. Darwin instead saw speciation as occurring by species entering new ecological niches.
Darwin's views
Controversy exists as to whether Charles Darwin recognized a true geographical-based model of speciation in his publication On the Origin of Species. In chapter 11, "Geographical Distribution", Darwin discusses geographic barriers to migration, stating for example that "barriers of any kind, or obstacles to free migration, are related in a close and important manner to the differences between the productions of various regions [of the world]". F. J. Sulloway contends that Darwin's position on speciation was "misleading" at the least and may have later misinformed Wagner and David Starr Jordan into believing that Darwin viewed sympatric speciation as the most important mode of speciation. Nevertheless, Darwin never fully accepted Wagner's concept of geographical speciation.
The evolutionary biologist James Mallet maintains that the mantra repeated concerning Darwin's Origin of Species book having never actually discussed speciation is specious. The claim began with Thomas Henry Huxley and George Romanes (contemporaries of Darwin's), who declared that Darwin failed to explain the origins of inviability and sterility in hybrids. Similar claims were promulgated by the mutationist school of thought during the late 20th century, and even after the modern evolutionary synthesis by Richard Goldschmidt. Another strong proponent of this view about Darwin came from Mayr. Mayr maintained that Darwin was unable to address the problem of speciation, as he did not define species using the biological species concept. However, Mayr's view has not been entirely accepted, as Darwin's transmutation notebooks contained writings concerning the role of isolation in the splitting of species. Furthermore, Many of Darwin's ideas on speciation largely match the modern theories of both adaptive radiation and ecological speciation.
Darwin's dilemmas
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 a critical factor behind the origin of new species is reproductive isolation. Darwin also considered the second aspect of the origin of species.
Darwin was perplexed by the clustering of organisms into species. 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.