speciation

n.
The evolutionary formation of new biological species, usually by the division of a single species into two or more genetically distinct ones.

[SPECI(ES) + -ATION.]

speciate spe'ci·ate' v.
speciational spe'ci·a'tion·al adj.

speciation

For more information on speciation, visit Britannica.com.

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. We discuss this group, along with all other mammalian orders, later in this essay.)

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. (See Paleontology for more on these subjects.)

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.

Mammalian Orders

The listing of the 20 orders of living mammals that follows is arranged not alphabetically but in the probable order in which these groups evolved. (This is not to imply that the process was orderly or linear; it was not.) Very few dates are given, simply because there is much dispute in most cases. Numbers of species within each order are also a subject of debate among taxonomists, and therefore these numbers are not always precise.

In the essay, Species, there is a taxonomic listing of the obligatory ranks for humans; included within that listing is a short description of the kingdom (Animalia), phylum (Chordata), and subphylum (Vertebrata) to which mammals belong. Mammal itself is defined as a vertebrate (an animal with a spinal column) that feeds its young from special milk-secreting glands, termed mammae, located on the mother's body. Mammals are warm-blooded or endothermic, meaning that their internal temperatures remain relatively stable, and their bodies usually are covered with hair. They have other distinguishing characteristics as well, such as a relatively large cranium (skull) with a hinged lower jaw attached to it.

Monotremes

Order Monotrema consists of primitive, egg-laying mammals spread throughout parts of the region known as Oceania, which includes Australia, New Zealand, and islands of the southeastern Pacific. The habitat of this order lies specifically in Australia, Tasmania, and New Guinea. Monotremes, as they are called, are distinguished further by the fact that their mammary glands are without nipples, that teeth are present only in the young, and that adults have horny beaks.

The monotremes illustrate the fact that to be constituted as an order or family, a taxon, or taxonomic group, need not have large numbers. The entire order consists of a single existing species, the duck-billed platypus, which constitutes a family of its own, and two species of echidnas, creatures that look like a cross between a platypus and a porcupine. (The "porcupine" look comes from the fact that their bodies are covered in spines, or spiky protrusions.)

Marsupials

The marsupials, or order Marsupialia, include two other famous animal citizens of Oceania: the kangaroo and its close relative, the wallaby. Marsupials' young are poorly developed at birth and must continue to grow while attached to their mothers' nipples. For this reason, they must remain close to the mother, and therefore natural selection for marsupials favored those strains in which females possess a pouch bearing four teats.

Immediately after birth, the young marsupial (a kangaroo baby is called a joey) installs itself in the mother's pouch. Given this situation, marsupials can support only one offspring a year and thus are not given to the large litters that characterize another order, Carnivora, which we discuss later. Kangaroo offspring remain in the pouch until the age of about 7-10 months, by which time the mother has conceived again; the female kangaroo goes into heat just a few days after giving birth. The embryonic kangaroo remains in a state of dormancy, or arrested development, until the older sibling has left the pouch.

The marsupial order (some authorities call it a superorder, with numerous subordinate orders) consists of some 240 species. The greatest number of these, including many species of kangaroo, wallaby, wombat, and koala, are found exclusively in Australia. Some 70 additional species are scattered across parts of Oceania, including Australia, Tasmania, New Guinea, and smaller islands. There are an additional 70 species in the Americas, including four species of the genus Didelphis—the large American opossum, better known in the southern United States as possums.

Why the preponderance of marsupials in Australia and Oceania as a whole? The reason lies in Earth's geologic history, which has seen regular collisions and divisions of the continents, which are even today shifting slowly under our feet. It appears that prior to about 70 million years ago, at a time when most of Earth was united in a single supercontinent called Pangaea, marsupials originated in what is now North America and migrated to the land masses that became Australia and Oceania. Because the bulk of marsupial species remained on Australia and nearby areas when Pangaea began to break apart, marsupials underwent much greater speciation there than in North America.

Xenarthrans, Insectivores, Scandentia, and Dermoptera

Known variously as xenarthrans and edentates, members of order Xenarthra either lack teeth or have very small ones. Evolutionary development has adapted the forward limbs of these creatures for digging or for holding on to the branches of trees. Included in this order are some 30 species of sloth, anteater, and armadillo. Sloths are herbivores, armadillos are omnivores (i.e., they eat plants and small animals), and anteaters, as their name would suggest, are hard-core insectivores.

The term insectivore can refer to any organism that lives by eating insects, but it is also the name for members of the order Insectivora, which includes shrews, hedgehogs, moles, and various other, less well known groups. Some 400 species, of which about 300 are shrews in a single family (Soricidae), make up this order. Not only their diet but also their pointed snouts and rodent-like appearance distinguishes this group. Many, but not all, are diggers, like the xenarthrans. Like all mammals, they have the pentadactyl limb (an appendage with five digits, like the human arm and hand)—in their case, a foot with five toes.

Order Scandentia, which is identical with the family Tupaiidae, or tree shrews, is sometimes grouped with order Insectivora. Despite their name, tree shrews, of which there are five genera and between 15 and 19 species, may live either on the ground or in the trees. Squirrel-like in appearance, they have strong claws on all their toes and are excellent climbers.

Another very small order of tree-dwellers is Dermoptera, which consists of just two species of flying lemur. Found in Indonesia and the Philippines, these creatures are equipped with skin flaps adapted for gliding. This aspect of their morphological makeup calls to mind the "flying" squirrel, but their nocturnal habits are more like those of lemurs (discussed later, with other primates); hence their common name.

Chiropterans

Among the most fascinating of mammalian orders is Chiroptera, better known as bats. This order, which consists of about 900 species in some 175 genera, is the only group of truly flying mammals, as opposed to the "flying" lemurs we just discussed. Yet they, too, have the pentadactyl limb, only in their case the forelimb has been adapted as a wing. Among the intriguing features of bats is their use of acoustic orientation, or echolocation, to find their way through the dark caves and nocturnal exteriors that make up their world. Contrary to popular belief, they are not blind, but they do have very small eyes, simply because vision is not important for bat navigation. (See Migration and Navigation for more about this subject.)

As befits an order with such a wide array of species, bats run the gamut with respect to their eating habits. The majority of bat species are insectivores that consume many thousands times their weight in insects each year. Many others are fruit bats, important members of the ecosystems they occupy, because they consume fruit and spread seeds, helping assist in seed dispersal. (Some bats also aid in pollination; see Reproduction). Then there are the three species of vampire bat, which are largely responsible for bats' unfortunate reputation with humans.

Members of subfamily Desmodontinae, species of vampire bat include Diaemus youngi and Diphylla ecaudata. However, the best-known variety is the common vampire bat, Desmodus rotundus, which is native to an area that stretches from the southwestern United States to the northwestern third of South America. As one would suspect of a creature named vampire, they live off blood, which they suck at night from birds or other mammals, including humans. These creatures are livestock pests and strike fear in humans both because of the imaginary association with vampires and for the quite real threat of contracting rabies from them. The vast majority of bats, however, are creatures that cause no harm to humans and often are unfairly persecuted as the result of human prejudices. Even in the case of the vampire bat, there is a strong possibility that it may one day help save human lives. Scientists have discovered that the saliva of Desmodus rotundus is better than any other known substance for keeping blood from clotting; therefore, vampire bat saliva may one day be adapted for use in treating heart attacks and strokes.

Primates

The order of humans, Primates, falls approximately in the middle of the mammalian class in terms of evolutionary order. This is an interesting aspect of speciation, evolution, and taxonomy: even though humans themselves are the most advanced of all creatures, it is not a logical necessity that we should come from the most recently evolved order. In fact, the opposite would seem to be the case. To produce a species whose intelligence dwarfs that of all other animals, the line of descent should be a long one. Where primates are concerned, that is certainly the case. The oldest primate samples date back some 75 million years, or long before the end of the Mesozoic.

Because it is from primates that humans draw their lineage, more has been written about primate evolution than on that of all other mammalian orders combined. The subject is such a vast one that we will not attempt to approach it here, except to encourage the reader to study in more detail elsewhere the process by which the human lineage emerged from order Primates, family Hominidae, and genus Homo.

Primates consist of two broad groups, suborders Prosimii and Anthropoidea. The first, the prosimians, includes five families (or six, since tree shrews are sometimes included) of lemurs, lorises, and tarsiers. The other suborder, known as the higher primates, encompasses another six families: marmosets and tamarins; South American monkeys other than marmosets; African and Asian monkeys; lesser apes, or siamangs and gibbons; great apes, or orangutans, gorillas, and chimpanzees; and humans, both living and extinct. (Most orders contain extinct members, but for the most part they are not discussed here.)

Most primates are tree dwellers, and among the approximately 230 species, there is enormous variation in eating habits. Many lemurs are insectivores, while great apes tend to be fruit eaters. Quite a few are omnivores, though no primate other than humans is known for eating large mammals, such as cows, sheep, and pigs. The pentadactyl limb (an appendage with five digits) is a significant feature for primates, which alone have the advantage of the opposable thumb for grasping. Humans and a few other primate species are also the only animals with four limbs who are not only capable of standing upright but also function best in this way.

Carnivores

As with insectivore , carnivore is a name both for an eating preference—in this case, meat—and for members of a primate order, Carnivora. Most members of this extraordinarily varied group eat meat, including, in some cases, the "meat" of insects. Bears and some other species are omnivorous, meaning that they also eat plants, and hyenas and jackals are classic examples of detritivores, or animals who feed on the remains of other creatures.

The distinction between detritivore and carnivore relates not to the materials each consumes but to their place in the food web. Rather than consume live creatures, hyenas and jackals feed on the carcasses of dead ones. Usually these creatures are artiodactyls (discussed later), such as antelopes, which have been killed by other carnivores—big cats, such as the lion or cheetah. After the big cats have fed on the fleshy parts of the prey, hyenas come to consume the flesh that remains, and they are followed by jackals and vultures, swoop in to pick the bones. These detritivores help process the remains of formerly living things, which ultimately return to the soil. (See Food Webs for more on this subject.)

Clearly, all members of order Carnivora eat meat, though in different ways and sometimes in combination with fruit or other vegetation. Natural selection has equipped them for this purpose with sharp claws and teeth. Carnivora includes some 270 species grouped into ten families, listed here:

• Canidae (dogs, wolves, jackals, and foxes)
• Felidae (cats)
• Hyaenidae (hyenas)
• Mustelidae (skunks, mink, weasels, badgers, and otters)
• Otariidae (eared seals)
• Odobenidae (walrus)
• Phocidae (earless seals)
• Procyonidae (raccoons)
• Ursidae (bears)
• Viverridae (mongooses and civets).

Note that Felidae is a particularly varied family of some 36 species: lions, lynxes, tigers, leopards, and even ordinary domesticated cats. Thirty-five species belong to a single subfamily, Felinae, which is native to most parts of the world other than Australia, Madagascar, most oceanic islands, and, of course, Antarctica. The last species, the cheetah (Acinonyx jubatus), is segregated into another subfamily, Acinonychinae, primarily because this cat, native to Africa and southwest Asia, is a daytime hunter, unlike its nocturnal cousins.

Cetaceans and Sirenians

Orders Cetacea and Sirenia include the majority of aquatic mammals, as opposed to the many amphibious mammals, such as seals, sea lions, sea elephants, and walruses, that belong variously to families Otariidae, Odobenidae, and Phocidae of the order Carnivora. Cetaceans include whales, dolphins, and porpoises, while sirenians are made up of just three species of manatee and one of dugong. Sirenians are large, friendly creatures that inhabit the Atlantic coast and tributary rivers (manatee) or the Indian and Pacific coastlines (dugong), but cetaceans are much more familiar.

With cetaceans, two questions, one specific and one general, often arise. The answer to the first of these, "What is the difference between a dolphin and a porpoise?," is that a porpoise is smaller and more chubby and has a blunt snout, whereas a dolphin has a beaklike snout. Some taxonomists and marine biologists put porpoises in the same family as dolphins, whereas others treat them as two different families. The more general, and much more important, question is "Why are these mammals living in the water?" In fact, life itself first appeared in the sea, so perhaps the question should be "Why or how did anything start living on land?" The transition from water to land took place long before the age of the dinosaurs, much less the emergence of mammals, but later, some mammals began to return to the water, probably about 70 million years ago.

Certainly, there is no question that cetaceans are mammals, a fact first recognized by Aristotle (384-322 B.C.), a Greek thinker who is noted not only as one of the greatest philosophers of all time but also as the father of the biological sciences. (See Taxonomy for more about Aristotle's contributions.) As Aristotle observed, whales and dolphins bear live young and suckle them with milk-producing glands; their bodies have hair, albeit only very small strands; and they possess lungs, breathing air through a blowhole.

As evidence of their terrestrial, or land-based, origins, consider that a whale fetus possesses the remnants of four limbs, each with five fingers (the pentadactyl limb), like any land mammal. Adult whales and dolphins have the streamlined, fishlike morphological appearance that is necessary for life underwater, but their resemblance to fishes is of the superficial, analogous variety discussed in Taxonomy. They have maintained and modified key terrestrial features; for example, a blowhole atop the head—one in dolphins, two in whales—replaces the nostrils, and thus the passageways for food and air are completely separate. This differs from the situation with most terrestrial mammals, which take in food and air through the same opening.

Proboscideans, Perissodactyls, and Hyraxes

Moving from the largest aquatic mammals, the whales, to the largest terrestrial variety, we come to the order Proboscidea, which includes elephants. Our discussions of most orders in class Mammalia have illustrated particular aspects of taxonomy and speciation, and so it is with proboscideans, which give evidence of the many species from the past that are gone forever. The order is a large one, with three suborders and some 300 species, but anyone who searches for most of those species will search in vain. All but three species are extinct. These three are the Asian elephant (Elephas maximus) and two varieties of African elephant (Loxodonta africana or African savanna elephant and Loxodonta cyclotisare, the African forest elephant). Until 2001, taxonomists believed that there were only two living species of elephant. (See Taxonomy for more on this subject.)

Another 16 species belong to the order Perissodactyla, herbivores whose hind feet bear either one or three hoofed toes. Included among perissodactyls is another large animal from the grasslands of Africa and tropical Asia: the rhinoceros. The group also encompasses donkeys, zebras, and tapirs, but by far the most important in human terms is Equus caballus, the domesticated horse. Described by the French zoologist Comte Georges de Buffon (1707-1788) as "the proudest conquest of man," the horse was domesticated (adapted so as to be useful and advantageous for humans) some 6,000 years ago. Nonetheless, feral or wild horses remain an important subspecies.

Order Hyracoidea also consists of hoofed mammals: seven species of hyrax, a primarily herbivorous creature native to Africa and the far southwestern extremities of Asia. Hyraxes are sometimes lumped in with pikas under the term rock rabbit, but, in fact, pikas are lagomorphs, a group we discuss later. To further the confusion, hyraxes are probably the animal called a coney in the Bible, even though there is an animal called a cony (no "e") that is actually a lagomorph. This is just one of many examples of confusion resulting from the complexities of the animal world and humans' attempts to name and classify its members.

Tubulidentates

For most orders, the lowercase adjectival name (e.g., primates, carnivores, insectivores, and so on) is commonly used. On the other hand, the names hyracoidean and tubulidentate are seldom used for more obscure groups, such as Hyracoidea and Tubulidentata. If any order of mammal is obscure, it is Tubulidentata, which emerged some 60 million years ago and which consists of a single species: the African ant bear (Orycteropus afer). The latter creature is better known by the name aardvark, which in the Afrikaans language means "earth pig."

Aardvarks at first glance might seem to belong with anteaters, sloths, and armadillos in the order Edentata, and that is what taxonomists thought for a long time. The latter half of the name tubulidentate, however, suggests the area of differentiation: the teeth. Aardvarks' teeth are unique among those of all mammals. Viewed from the top, the aardvark's jawbone is V-shaped, with the teeth midway along either side of the V. The teeth themselves are not fixed to the jaw but rest in the flesh attached to it, and instead of being covered with enamel, they are protected with a cementlike substance. The substance comes from tubules that run under the teeth.

Artiodactyls

Like many other mammalian orders we have discussed, members of order Artiodactyla are ungulates, or hoofed animals. Whereas perissodactyls have odd-numbered toes, artiodactyls have even-numbered toes—either two or four—on each foot. This large group, consisting of some 220 species, comprises a wide variety of well-known species in nine families. Among them are cows, pigs, sheep, goats, deer, antelope, bison, camels, giraffes, hippopotamuses, and numerous less well known varieties, such as okapi, pronghorn, peccaries, and deerlike chevrotains.

The camel family is particularly widespread geographically, including as it does many varieties—the llama, alpaca, and vicuña of South America—whose home is far from the habitats in the Near East that are associated with the camel. In this family is what may be a previously undiscovered species, whose existence the British Broadcasting Corporation (BBC) reported in early 2001. Living on a former nuclear weapons testing range in a remote region of Chinese central Asia, these creatures drink saltwater, which in itself is an unusual characteristic.

Although it is extraordinarily hardy, even by the standards of camels, the central Asian camels are threatened; fewer than 1,000 remain. As the BBC reported, this makes them more endangered than the more well known giant panda. The creatures survived nuclear testing in the area, which ceased in 1996, but they continue to be threatened by much less spectacular varieties of explosive: dynamite and land mines, planted by hungry locals. John Hare of the Wild Camel Protection Foundation told the BBC, "We found land mines put by the saltwater springs. So when the camels come to drink, they step on them. Bang! They are blown to pieces and picked up as meat."

As to whether the camels constitute a separate species, the molecular geneticist Olivier Hanotte told the BBC: "There are two possibilities here. One is that the domestic camel was bred from these wild ones some time back in history. The second is that the domestic camel we see today was bred from another species that has disappeared. This would mean that these wild camels are a totally separate species." As of early 2002 the camels' fate, both practically and taxonomically, remained undecided.

Pholidota/Pangolins

Order Pholidota consists of seven species of scaly anteater, or pangolin. Members of this order are normally called pangolins, rather than an adjectival form of Pholidota. The word pangolin comes from a Malay term meaning "rolling over," a reference to the fact that when it is threatened, the animal curls into a little ball. As with the aardvark, members of this order once were grouped with Edentata but now are considered a separate order. Pangolins are also like aardvarks in thesense that their evolutionary relationship toother mammals is not clear.

Rodents, Lagomorphs, Andmacroscelideans

Rodents, or members of order Rodentia, are familiar to us as both pests and pets as well as aids to research through their use as test subjects in laboratories. They are also the most abundant of all mammalian orders: about one-fourth of all families, 35% of all genera, and 50% of all living species of mammal are rodents. The group consists of some 2,205 species, among them mice, rats, squirrels, beavers, gophers, and porcupines. The name rodent comes from the Latin rodere, meaning "to gnaw," and, indeed, the defining characteristic of rodents is their chisel-like upper front teeth.

Whereas rats typically are despised creatures, mice (distinguished from rats simply because they are smaller) often are considered cute—that is, if the mouse in question is a pet or a laboratory mouse, rather than a pest chewing up the insulation or electrical wiring in someone's house. The fact that rodents are so often pests and pets arises in part from rodents' close association with humans. This is a distinction in itself, since few mammal orders manage to live successfully in such close proximity to humans. Not only do squirrels often live around human dwellings, but other species (for better or worse) often enter structures where humans live or work. Particularly notorious in this regard are black rats (Rattus rattus) and Norway rats (R. norvegicus), which are just two of some 500 rat species.

The two remaining orders of mammal also are composed of small, furry creatures. Lagomorphs, or members of the order Lagomorpha, are small mammals with large upper incisors (front teeth) but no canines or eyeteeth and with molars that lack roots. The 80-odd species of lagomorphs include rabbits, hares, and their lesser-known cousin the pika, or mouse-hare. The difference between rabbits and hares relates to their conditions at birth: rabbits are furless, blind, and helpless, whereas hares are furry, have open eyes, and are capable of hopping within minutes.

Finally, 28 species of elephant shrew, or jumping shrew, make up the order Macroscelidea, a collection of species known for their long, flexible, sensitive snouts. Some authorities group macroscelideans with order Insectivora, whereas others place them in another order, Mentophyla, with tree shrews. The latter often have been placed variously in orders Scandentia, Insectivora, or Primates, indicating that many areas of mammalian taxonomy remain in dispute.

Where to Learn More

Boxhorn, Joseph. "Observed Instances of Speciation." Talk. Origins (Web site). <http://www.talkorigins.org/faqs/faq-speciation.html>.

Kirby, Alex. "'New' Camel Lives on Salty Water." British Broadcasting Corporation (Web site). <http://news.bbc.co.uk/hi/english/sci/tech/newsid_1156000/1156212.stm>.

"Mammalia." Animal Diversity Web, The University of Michigan Museum of Zoology <http://animaldiversity.ummz.umich.edu/chordata/mammalia.html>.

Mammal Species of the World (MSW). Smithsonian National Museum of Natural History, Department of Systematic Biology—Vertebrate Zoology (Web site). <http://www.nmnh.si.edu/msw/>.

Marks, Jonathan. Human Biodiversity: Genes, Race, and History. New York: Aldine de Gruyter, 1995.

Norton, Bryan G. The Preservation of Species: The Value of Biological Diversity. Princeton, NJ: Princeton University Press, 1986.

Patent, Dorothy Hinshaw. The Challenge of Extinction. Hillside, NJ: Enslow Publishers, 1991.

——. Biodiversity. Illus. William Muñoz. New York: Clarion Books, 1996.

Schilthuizen, Menno. Frogs, Flies, and Dandelions: Speciation—The Evolution of New Species. New York: Oxford University Press, 2001.

Speciation (Web site). <http://www.ultranet.com/~jkimball/BiologyPages/S/Speciation.html>.

UCMP Hall of Mammals, University of California, Berkeley, Museum of Paleontology (Web site). <http://www.ucmp.berkeley.edu/mammal/mammal.html>.

---

The process by which new species of organisms evolve from preexisting species. It is part of the whole process of organic evolution. The modern period of its study began with the publication of Charles Darwin's and Alfred Russell Wallace's Theory of Evolution by Natural Selection in 1858, and Darwin's On the Origin of Species in 1859.

Belief in the fixity of species was almost universal before the middle of the nineteenth century. Then it was gradually realized that all species continuously change, or evolve; however, the causative mechanism remained to be discovered. Darwin proposed a mechanism. He argued that (1) within any species population there is always some heritable variation; the individuals differ among themselves in structure, physiology, and behavior; and (2) natural selection acts upon this variation by eliminating the less fit. Thus if two members of an animal population differ from each other in their ability to find a mate, obtain food, escape from predators, resist the ravages of parasites and pathogens, or survive the rigors of the climate, the more successful will be more likely than the less successful to leave descendants. The more successful is said to have greater fitness, to be better adapted, or to be selectively favored. Likewise among plants: one plant individual is fitter than another if its heritable characteristics make it more successful than the other in obtaining light, water, and nutrients, in protecting itself from herbivores and disease organisms, or in surviving adverse climatic conditions. Over the course of time, as the fitter members of a population leave more descendants than the less fit, their characteristics become more common.

This is the process of natural selection, which tends to preserve the well adapted at the expense of the ill adapted in a variable population. The genetic variability that must exist if natural selection is to act is generated by genetic mutations in the broad sense, including chromosomal rearrangements together with point mutations. See also Genetics; Mutation.

If two separate populations of a species live in separate regions, exposed to different environments, natural selection will cause each population to accumulate characters adapting it to its own environment. The two populations will thus diverge from each other and, given time, will become so different that they are no longer interfertile. At this point, speciation has occurred: two species have come into existence in the place of one. This mode of speciation, speciation by splitting, is probably the most common mode. Two other modes are hybrid speciation and phyletic speciation; many biologists do not regard the latter as true speciation.

Many students of evolution are of the opinion that most groups of organisms evolve in accordance with the punctuated equilibrium model rather than by phyletic gradualism. There are two chief arguments for this view. First, it is clear from the fossil record that many species persist without perceptible change over long stretches of time and then suddenly make large quantum jumps to radically new forms. Second, phyletic gradualism seems to be too slow a process to account for the tremendous proliferation of species needed to supply the vast array of living forms that have come into existence since life first appeared on Earth. See also Animal evolution; Population genetics; Species concept.

---

Speciation is the process by which new species of organisms arise. Earth is inhabited by millions of different organisms, all of which likely arose from one early life-form that came into existence about 3.5 billion years ago. It is the task of taxonomists to decide which out of the multitude of different types of organisms should be considered species. The wide range in the characteristics of individuals within groups makes defining a species more difficult. Indeed, the definition of species itself is open to debate.

Concepts of Species

In the broadest sense, a species can be defined as a group of individuals that is "distinct" from another group of individuals. Several different views have been put forward about what constitutes an appropriate level of difference. Principal among these views are the biological-species concept and the morphological-species concept.

The biological-species concept delimits species based on breeding. Members of a single species are those that interbreed to produce fertile off-spring or have the potential to do so. The morphological-species concept (from the ancient Greek root "morphos," meaning form) is based on classifying species by a difference in their form or function. According to this concept, members of the same species share similar characteristics. Species that are designated by this criteria are known as a morphological species.

Organisms within a species do not necessarily look identical. For example, the domestic dog is considered to be one species, even though there is a huge range in size and appearance among the different breeds. For naturally occurring populations of organisms that we are much less familiar with, it is much more difficult to recognize the significance of any character differences observed. Therefore deciding what characteristics should be used as criteria to designate a species can be difficult.

Speciation Mechanisms: Natural Selection and Genetic Drift

Before the development of the modern theory of evolution, a widely held idea regarding the diversity of life was the "typological" or "essentialist" view. This view held that a species at its core had an unchanging perfect "type" and that any variations on this perfect type were imperfections due to environmental conditions. Charles Darwin (1809-1882) and Alfred Russel Wallace (1823-1913) independently developed the theory of evolution by natural selection, now commonly known as Darwinian evolution.

The theory of Darwinian evolution is based on two main ideas. The first is that heritable traits that confer an advantage to the individual that carries them will become more widespread in a population through natural selection because organisms with these favorable traits will produce more offspring. Since different environments favor different traits, Darwin saw that the process of natural selection would, over time, make two originally similar groups become different from one another, ultimately creating two species from one. This led to the second major idea, which is that all species arise from earlier species, therefore sharing a common ancestor.

When so much change occurs between different groups that they are morphologically distinct or no longer able to interbreed, they may be considered different species; this process is known as speciation. A species as a whole can transform over time into a new species (vertical evolution) or split into more separate populations, each of which may develop into new species (adaptive radiation).

Modern population geneticists recognize that natural selection is not the only factor causing genetic change in a population over time. Genetic drift is the random change in the genetic composition of a small population over time, due to an unequal genetic contribution by individuals to succeeding generations. It is thought that genetic drift can result in new species, especially in small isolated populations.

Isolating Mechanisms

Whether natural selection and genetic drift lead to new species depends on whether there is restricted gene flow between different groups. Gene flow is the movement of genes between separate populations by migration of individuals. If two populations remain in contact, gene flow will prevent them from becoming separate species (though they may both develop into a new species through vertical evolution).

Gene flow is restricted through geographic effects such as mountain ranges and oceans, leading to geographic isolation. Gene flow can also be prevented by biological factors known as isolating mechanisms. Biological isolating mechanisms include differences in behavior (especially mating behavior), and differences in habitat use, both of which lead to a decrease in mating between individuals from different groups.

When geographic separation plays a role in speciation, this is known as allopatric speciation, from the Greek roots allo, meaning separate, and "patric," meaning country. In allopatric speciation, natural selection and genetic drift can act together.

For example, imagine a mud slide that causes a river to back up into a valley, separating a population of rodents into two, one restricted to the shady side of the river, the other to the sunny side. Because coat thickness is a genetically inherited trait, eventually, through natural selection, the population of animals on the cooler side may develop thicker coats. After many generations of separation, the two groups may look quite different and may have evolved different behaviors as well, to allow them to survive better in their respective habitats. Genetic drift may occur especially if either or both populations remain small. Eventually these two populations may be so different as to warrant designation as different species.

It is also possible for new species to form from a single population without any geographic separation. This is known as "ecological" or "sympatric" (from the Greek root sym, meaning same) speciation, and it results in ecological differences between morphologically similar species inhabiting the same area. Sympatric speciation can occur in flowering plants in a single generation, due to the formation of a polyploid. Polyploidy is the complete duplication of an organism's genome, for example from n chromosomes to 4n. Even higher multiples of n are possible. This increase in a plant's DNA content makes it reproductively incompatible with other individuals of its former species.

Bibliography

Futuyma, Douglas J. Evolutionary Biology, 3rd ed. Sunderland, MA: Sinauer Associates, 1998.

Mayr, Ernst. Evolution and the Diversity of Life: Selected Essays. Cambridge, MA: Belknap Press, 1976.

—R. John Nelson

What is speciation?

Speciation is the process by which new species are formed. This occurs when populations become separated from the rest of the species. At this point the isolated group will respond independently to natural selection until the population becomes reproductively isolated. The group is then considered a new species.

Previous question:	What are different ways of defining the term "species"?
Next question:	How can speciation occur?

---

This entry is from Wikipedia, the leading user-contributed encyclopedia. It may not have been reviewed by professional editors (see full disclaimer)