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.

Biologists group allopatric processes into two categories: dispersal and vicariance. Dispersal is when a few members of a species move to a new geographical area, and vicariance is when a natural situation arises to physically divide organisms.

Scientists have documented numerous cases of allopatric speciation taking place. For example, along the west coast of the United States, two separate sub-species of spotted owls exist. The northern spotted owl has genetic and phenotypic differences from its close relative: the Mexican spotted owl, which lives in the south (Figure 18.14).

Figure 18.14 The northern spotted owl and the Mexican spotted owl inhabit geographically separate locations with different climates and ecosystems. The owl is an example of allopatric speciation. (credit northern spotted owl: modification of work by John and Karen Hollingsworth; credit Mexican spotted owl: modification of work by Bill Radke)

Additionally, scientists have found that the farther the distance between two groups that once were the same species, the more likely it is that speciation will occur. This seems logical because as the distance increases, the various environmental factors would likely have less in common than locations in close proximity. Consider the two owls: In the north, the climate is cooler than in the south; the types of organisms in each ecosystem differ, as do their behaviors and habits; also, the hunting habits and prey choices of the southern owls vary from the northern owls. These variances can lead to evolved differences in the owls, and speciation likely will occur.

Adaptive Radiation

In some cases, a population of one species disperses throughout an area, and each population finds a distinct niche or isolated habitat. Over time, the varied demands of their new lifestyles lead to multiple speciation events originating from a single species. This is called adaptive radiation because many adaptations evolve from a single point of origin; thus, causing the species to radiate into several new ones. Island archipelagos like the Hawaiian Islands provide an ideal context for adaptive radiation events because water surrounds each island which leads to geographical isolation for many organisms. The Hawaiian honeycreeper illustrates one example of adaptive radiation. From a single species, called the founder species, numerous species have evolved, including the six shown in Figure 18.15.

Figure 18.15 The honeycreeper birds illustrate adaptive radiation. From one original species of bird, multiple others evolved, each with its own distinctive characteristics.

Notice the differences in the species’ beaks in Figure 18.15. Evolution in response to natural selection based on specific food sources in each new habitat led to evolution of a different beak suited to the specific food source. The seed-eating bird has a thicker, stronger beak which is suited to break hard nuts. The nectar-eating birds have long beaks to dip into flowers to reach the nectar. The insect-eating birds have beaks like swords, appropriate for stabbing and impaling insects. Darwin’s finches are another example of adaptive radiation in an archipelago.

Link to Learning

Click through this interactive site to see how island birds evolved in evolutionary increments from 5 million years ago to today.

Name three examples of adaptive radiation, and provide a brief statement about each one.

1. Domestic dogs – There are over 300 distinct dog breeds. Cows – There are over 800 cow breeds recognized worldwide. Domestic cats – Cats have changed drastically in just a few thousand years.
2. Whales and fish – Although it has been roughly 400 million years since fish and mammals diverged, whales and fish are morphologically similar. Birds and butterflies – Although the common ancestor between vertebrates and insects lived even longer ago than the common ancestor between whales and fish, birds and butterflies both developed flight. Rabbits and kangaroos – Although it has been over 150 million years since their divergence, rabbits and kangaroos both developed powerful jumping legs.
3. Hawaiian silverswords-There are about 30 species evolved from one parent species. Madagascar lemurs-Their common ancestor likely arrived to Madagascar over 60 million years ago. Hawaiian fruit fly-There are 500 species of fruit fly from one parent species.
4. Beetles – There are about 350,000 species of beetles that we know of. Birds – There are almost 10,000 species of birds in existence. Frogs – There are almost 5,000 species of frogs worldwide.

Sympatric Speciation

Can divergence occur if no physical barriers are in place to separate individuals who continue to live and reproduce in the same habitat? The answer is yes. The process of speciation within the same space is called sympatric speciation; the prefix sym means same, so sympatric means same homeland in contrast to allopatric meaning other homeland. A number of mechanisms for sympatric speciation have been proposed and studied.

One form of sympatric speciation can begin with a serious chromosomal error during cell division. In a normal cell division event chromosomes replicate, pair up, and then separate so that each new cell has the same number of chromosomes. However, sometimes the pairs separate and the end cell product has too many or too few individual chromosomes in a condition called aneuploidy (Figure 18.16).

Visual Connection

Figure 18.16 Aneuploidy results when the gametes have too many or too few chromosomes due to nondisjunction during meiosis. In the example shown here, the resulting offspring will have 2n+1 or 2n-1 chromosomes

In your own words, define aneuploidy.

1. Aneuploidy is a mutation that causes the genome of a cell to break down.
2. neuploidy is the condition of having too many or too few chromosomes as a result of an error during cell division
3. Aneuploidy is the condition of having an extra copy of every chromosome in the genome.
4. Aneuploidy is a prezygotic barrier in which the gametes of two distinct species will not fertilize each other.

Polyploidy is a condition in which a cell or organism has an extra set, or sets, of chromosomes. Scientists have identified two main types of polyploidy that can lead to reproductive isolation of an individual in the polyploidy state. Reproductive isolation is the inability to interbreed. In some cases, a polyploid individual will have two or more complete sets of chromosomes from its own species in a condition called autopolyploidy (Figure 18.17). The prefix auto- means self, so the term means multiple chromosomes from one’s own species. Polyploidy results from an error in meiosis in which all of the chromosomes move into one cell instead of separating.

Figure 18.17 Autopolyploidy results when mitosis is not followed by cytokinesis.

For example, if a plant species with 2n = 6 produces autopolyploid gametes that are also diploid (2n = 6, when they should be n = 3), the gametes now have twice as many chromosomes as they should have. These new gametes will be incompatible with the normal gametes produced by this plant species. However, they could either self-pollinate or reproduce with other autopolyploid plants with gametes having the same diploid number. In this way, sympatric speciation can occur quickly by forming offspring with 4n called a tetraploid. These individuals would immediately be able to reproduce only with those of this new kind and not those of the ancestral species.

The other form of polyploidy occurs when individuals of two different species reproduce to form a viable offspring called an allopolyploid. The prefix allo- means other (recall from allopatric): Therefore, an allopolyploid occurs when gametes from two different species combine. Figure 18.18 illustrates one possible way an allopolyploid can form. Notice how it takes two generations, or two reproductive acts, before the viable fertile hybrid results.

Figure 18.18 Alloploidy results when two species mate to produce viable offspring. In the example shown, a normal gamete from one species fuses with a polyploidy gamete from another. Two matings are necessary to produce viable offspring.

The cultivated forms of wheat, cotton, and tobacco plants are all allopolyploids. Although polyploidy occurs occasionally in animals, it takes place most commonly in plants. Animals with any of the types of chromosomal aberrations described here are unlikely to survive and produce normal offspring. Scientists have discovered more than half of all plant species studied relate back to a species evolved through polyploidy. With such a high rate of polyploidy in plants, some scientists hypothesize that this mechanism takes place more as an adaptation than as an error.

Reproductive Isolation

Given enough time, the genetic and phenotypic divergence between populations will affect characters that influence reproduction: If individuals of the two populations were to be brought together, mating would be less likely, but if mating occurred, offspring would be nonviable or infertile. Many types of diverging characters may affect the reproductive isolation, the inability to interbreed, of the two populations.

Reproductive isolation can take place in a variety of ways. Scientists organize them into two groups: prezygotic barriers and postzygotic barriers. Recall that a zygote is a fertilized egg: The first cell of the development of an organism that reproduces sexually. Therefore, a prezygotic barrier is a mechanism that blocks reproduction from taking place; this includes barriers that prevent fertilization when organisms attempt reproduction. A postzygotic barrier occurs after zygote formation; this includes organisms that don’t survive the embryonic stage and those that are born sterile.

Some types of prezygotic barriers prevent reproduction entirely. Many organisms only reproduce at certain times of the year, often just annually. Differences in breeding schedules, called temporal isolation, can act as a form of reproductive isolation. For example, two species of frogs inhabit the same area, but one reproduces from January to March, whereas the other reproduces from March to May (Figure 18.19).

Figure 18.19 These two related frog species exhibit temporal reproductive isolation. (a) Rana aurora breeds earlier in the year than (b) Rana boylii. (credit a: modification of work by Mark R. Jennings, USFWS; credit b: modification of work by Alessandro Catenazzi)

In some cases, populations of a species move or are moved to a new habitat and take up residence in a place that no longer overlaps with the other populations of the same species. This situation is called habitat isolation. Reproduction with the parent species ceases, and a new group exists that is now reproductively and genetically independent. For example, a cricket population that was divided after a flood could no longer interact with each other. Over time, the forces of natural selection, mutation, and genetic drift will likely result in the divergence of the two groups (Figure 18.20).

Figure 18.20 Speciation can occur when two populations occupy different habitats. The habitats need not be far apart. The cricket (a) Gryllus pennsylvanicus prefers sandy soil, and the cricket (b) Gryllus firmus prefers loamy soil. The two species can live in close proximity, but because of their different soil preferences, they became genetically isolated.

Behavioral isolation occurs when the presence or absence of a specific behavior prevents reproduction from taking place. For example, male fireflies use specific light patterns to attract females. Various species of fireflies display their lights differently. If a male of one species tried to attract the female of another, she would not recognize the light pattern and would not mate with the male.

Other prezygotic barriers work when differences in their gamete cells, eggs and sperm, prevent fertilization from taking place; this is called a gametic barrier. Similarly, in some cases closely related organisms try to mate, but their reproductive structures simply do not fit together. For example, damselfly males and females of different species have differently shaped reproductive organs. If one species tries to mate with another, their body parts simply do not fit together (Figure 18.21).

Figure 18.21 The shape of the male reproductive organ varies among male damselfly species, and is only compatible with the female of the same species. Reproductive organ incompatibility keeps each species reproductively isolated.

In plants, certain structures aimed to attract one type of pollinator simultaneously prevent a different pollinator from accessing the pollen. The tunnel through which an animal must access nectar can vary widely in length and diameter, which prevents the plant from being cross-pollinated with a different species (Figure 18.22).

Figure 18.22 Some flowers have evolved to attract certain pollinators. The (a) wide foxglove flower is adapted for pollination by bees, while the (b) long, tube-shaped trumpet creeper flower is adapted for pollination by hummingbirds.

When fertilization takes place and a zygote forms, postzygotic barriers can prevent reproduction. Hybrid individuals in many cases cannot form normally in the womb and simply do not survive past the embryonic stages. This is called hybrid inviability because the hybrid organisms simply are not viable. In another postzygotic situation, reproduction leads to the birth and growth of a hybrid that is sterile and unable to reproduce offspring of their own; this is called hybrid sterility.

Habitat Influence on Speciation

Sympatric speciation may also take place in ways other than polyploidy. For example, consider a species of fish that lives in a lake. As the population grows, competition for food also grows. Under pressure to find food, suppose that a group of these fish had the genetic flexibility to discover and feed off another resource that was unused by the other fish. What if this new food source was found at a different depth of the lake? Over time, those feeding on the second food source would interact more with each other than the other fish; therefore, they would breed together as well. Offspring of these fish would likely behave as their parents: feeding and living in the same area and keeping separate from the original population. If this group of fish continued to remain separate from the first population, eventually sympatric speciation might occur as more genetic differences accumulated between them.

This scenario does play out in nature, as do others that lead to reproductive isolation. One such place is Lake Victoria in Africa, famous for its sympatric speciation of cichlid fish. Researchers have found hundreds of sympatric speciation events in these fish, which have not only happened in great number, but also over a short period of time. Figure 18.23 shows this type of speciation among a cichlid fish population in Nicaragua. In this locale, two types of cichlids live in the same geographic location but have come to have different morphologies that allow them to eat various food sources.

Figure 18.23 Cichlid fish from Lake Apoyeque, Nicaragua, show evidence of sympatric speciation. Lake Apoyeque, a crater lake, is 1,800 years old, but genetic evidence indicates that the lake was populated only 100 years ago by a single population of cichlid fish. Nevertheless, two populations with distinct morphologies and diets now exist in the lake, and scientists believe these populations may be in an early stage of speciation.