Speciation

Concept

One of the defining characteristics of a species is its reproductive isolation: the fact that among animals and plants that reproduce sexually, it is impossible for members of two different species to mate and produce fertile offspring. Speciation is the process whereby a single species develops over time into two distinct, reproductively isolated species. It is one of the key evolutionary processes and is responsible for the diversity of life that exists on Earth. In the following essay we explore not only the basic facts of speciation and biological diversity but also an example of adaptive radiation, in the form of the wide range of species within the mammalian order.

How It Works

Species and Speciation
The concept of species, discussed in the article devoted to that subject, is an extraordinarily complex one. Owing to limitations of space, that essay only hints at the many details, the competing schools of thought, and the varying definitions of species. Likewise, in the present context, it is possible to examine the concept of speciation only in the most cursory fashion. In addition to consulting the essay on Species for more information, the reader is encouraged to review the article on Taxonomy.
Taxonomy is the area of the biological sciences devoted to the identification, nomenclature, and classification of organisms according to apparent common characteristics. It uses a wide array of specialized rankings for grouping animals, but only seven of them are essential to most biology students. These seven, known as the obligatory hierarchy, are kingdom, phylum, class, order, family, genus, and species. In the case of mammals, it is also useful to refer to subphylum, which in this case is Vertebrata (see the classification of humans in Species), but for the most part it is enough for the beginning student to attain at least some mastery of the obligatory ranks.
Note that species is the most specific of these ranks, which is fitting, because species and specific come from the same Latin root, specie, or "kind." Nonetheless, it is difficult to define species beyond a reference to its place among the categories of the obligatory taxonomy. According to the biological species concept, discussed briefly in Species, a species is any population of individual organisms capable of mating with one another and producing fertile offspring in a natural setting. This is far from the only definition, however.

Interspecific Mating

Occasionally, it is possible to produce an infertile hybrid, such as a mule, which is created by the mating of a male donkey and a female horse, or a hinny, the product of the less common union between a male horse and a female donkey. The infertility is due to genetic disorders that arise when mating takes place between distinct species, and even this imperfect product is possible only by mating two species that are very closely related. Donkeys and horses, for instance, both belong to family Equidae, which makes them very closely connected.

In the taxonomic ranking of humans, this would be equivalent to a human mating with a fellow hominid, or member of family Hominidae. If the long-extinct genus Australopithecus were still around, it is not inconceivable that humans could mate with them and produce at least sterile offspring. Of course, it is unlikely that many humans would want to mate with Australopithecus, the most famous example of which was named "Lucy" after the Beatles' song "Lucy in the Sky with Diamonds." Standing about 3.5-5 ft. (1-1.5 m) tall, Australopithecus was very close in appearance to a modern ape who lived about four million years ago.

All of humans' close relatives are extinct, and today our nearest relatives are members of the order Primates: apes, monkeys, and marsupials. It is impossible to imagine a human mating with one of these animals and producing offspring of any kind. Likewise, it is extremely unlikely that a horse or donkey could mate with a tapir or rhinoceros, which are about as distant in relation to them as other primates are to us. (These species all belong to the order Perissodactyla, herbivorous mammals possessing either one or three hoofed toes on each hind foot.

The Problem of Defining Species

Although the biological species concept is accepted widely, it has its shortcomings, not least of which is the fact that not all species reproduce sexually. Although sexual reproduction is the case with a wide array of animals and even plants, quite a few organisms reproduce by some asexual means: for example, single-cell organisms reproduce by splitting.
Among the competing definitions of species is the phenetic (or morphological) species concept, which relies in part on common sense. According to the phenetic species concept, a species is the smallest possible population of organisms that consistently and continually remains distinct and distinguishable by ordinary methods of observation. There are also a variety of definitions that fall under the heading "phylogenetic species concepts," all of which maintain in one way or another that taxonomic classifications should incorporate the most widely recognized hypotheses regarding the evolutionary lines of descent that produced the organisms in question.

The Process of Speciation

Clearly, there is no hard and fast definition of species, but in general terms, everyone who has some familiarity with the concept has at least a basic knowledge of what does and does not qualify as a species. We will leave finer distinctions to trained taxonomists and other biologists and move on to a fact regarding which there is no disagreement: a wide array of species exists in the world today. Some estimates calculate the number of species in the five kingdoms—animals, plants, monerans, protista, and fungi (see Taxonomy for a very brief identification of each)—at about 1.5 million.
This is only the number of identified species, however. Other figures, based on the probable numbers of unidentified species in the world, put the sum total in the tens of millions. Whatever the case, it is obvious that over the course of evolutionary history (discussed in Evolution and Paleontology), there has been a widespread adaptive radiation—that is, a diversification of species as a result of specialized adaptations by particular populations of organisms.

Speciation events are described as either allopatric or sympatric. Allopatric ("different places") speciation occurs when a population of organisms is divided by a geographic barrier, a great example being the division of squirrel species caused by the formation of the Grand Canyon (see Evolution). Another example is the speciation of the black-throated green warbler, which today consists of one species in the eastern United States, along with three others in the western part of the country. Some scientists speculate that there may once have been a single species of black-throated green warbler, whose population was split by the formation of a glacier during the Pleistocene epoch. The latter was the period of the last ice age, which ended about 10,000 years ago, but the end of the ice age was a slow process. It may be that glaciers, formed in the latter part of that time, helped to separate what became three different western species.
Species share the same gene pool, or the sum of all genetic codes possessed by members of that species. The isolation of two populations slowly results in differences between gene pools, until the two populations are unable to interbreed either because of changes in mating behavior or because of incompatibility of the DNA between the two populations. (Deoxyribonucleic acid, or DNA, contains genetic codes for inheritance. See Genetics for more on this subject.) More rare than allopatric speciation, sympatric ("same place") speciation happens when a group of individuals becomes reproductively isolated from the larger population of the original species. This type of speciation typically results from mutation, or alterations in DNA that result in a genetic change.

Studies of three-spined sticklebacks, a variety of freshwater fish, in British Columbia have revealed what appears to be a fascinating example of sympatric speciation. Evolutionary biologist Dolph Schluter and others have discovered that the region contains two species of stickleback, one with a large mouth that feeds on large prey close to shore, the other with a small mouth that feeds on plankton in open water. Both species jointly inhabit five different lakes. Through DNA analysis, scientists have determined that the lakes were colonized independently by common marine ancestors, meaning that the process of sympatric speciation between the two varieties had to have occurred independently at least five times. This seems to indicate a situation of competition for resources that favored stickleback species at either extreme of size, as opposed to those of medium size and medium-sized mouths.

Rate of Evolutionary Change
Closely tied to speciation is the rate of evolutionary change, or the speed at which new species arise. This is a long process, one that is usually not observable within a human lifetime or even the span of many lifetimes, though bacteria at least have shown some evolutionary change in their growing resistance to antiobiotics (see Infection). DNA analysis (see Genetics and Genetic Engineering for more about DNA) has been used to examine the rate of evolutionary change. To perform such analysis, it is necessary first to determine the percentage of similarity between the organisms under study: the greater the similarity, the more recently the organisms probably diverged from a common stock. Data obtained in this manner then must be corroborated by information obtained from other sources, such as the fossil record and comparative anatomy studies.
At certain times the rate of evolutionary change can be very rapid, leaving little fossil evidence of intermediate forms, a phenomenon known as punctuated equilibrium. This is contrasted with phyletic (that is, evolutionary) gradualism. Of course, the term rapid in this context is relative, since we are talking about vast spans of time. Life on Earth has existed for about 3,000 million years, and the fossil record goes back some 1,000 million years. This is the case, in part, because to leave fossilized remains, an organism must have "hard parts" that can become mineralized to turn into fossils. (
Real-Life Applications

The Diversity of Mammals
One of the most interesting examples of speciation is that which has produced the vast array of species, including humans, that fall within the mammalian class. Mammals began evolving before the dawn of the Cenozoic era about 65 million years ago. The Cenozoic era, which started with a catastrophic event that brought about the mass extinction of the dinosaurs and the end of the Mesozoic era (see Paleontology), is truly the age of the mammal. Just as dinosaurs dominated the Mesozoic, today the world belongs to mammals as to no other class of creature.
Since its humble beginnings in the shadow of the dinosaurs, class Mammalia has undergone a massive radiation to the point that today some 4,625 species of mammal, in about 125 families and 24 orders, are recognized. (That number is changing, as noted later in the context of elephants.) This diversity is tied closely to mammals' enormous mobility, which facilitated their spread throughout the world. Aside from much less complex life-forms, such as arachnids and insects (see Parasites and Parasitology), mammals are believed to be distributed more widely throughout the world than any other comparable taxonomic grouping. Insects may be the most diverse of all animal classes, with numbers of species that may be many times greater than the number of mammals, but considering mammals' much-greater level of physical development and complexity, the diversity of their species is astounding.

Mammals' Early Evolution
In the next section we list the orders of mammals and give very brief descriptions of each. The purpose here is not to provide anything like a comprehensive discussion but rather to illustrate the enormous range of species in a class that includes anteaters, dolphins, humans, elephants, and bats. The fact that all these diverse creatures, and many more, emerged from a common evolutionary lineage is almost as amazing as the fact that this common ancestor was a reptile.
Mammals are believed to have come from the reptilian order Therapsida, which emerged during the Triassic period (from about 245 to 208 million years ago) in the early part of the Mesozoic era. Over the course of many millions of years, these creatures began to develop a number of mammal-like qualities—in particular, endothermy, or the ability to maintain internal temperature regardless of environmental conditions. In other words, these cold-blooded creatures became warm-blooded. This evolutionary process was as complex as it was lengthy. Nor was there a clean break with the past—no moment when the therapsids faded away or when it would have been clear that mammals had taken the place of their reptilian ancestors. Rather, in what must have been a fascinating taxonomic situation, for many millions of years, species that combined aspects of both reptiles and mammals walked the earth.