7 Chapter 7: Speciation

Mason Tedeschi and Lisa Limeri

Learning Objectives

By the end of this section, students will be able to…

  • Describe the relationship between micro- and macro-evolution.
  • Describe how allopatric speciation or sympatric speciation can occur within a lineage.
  • Explain 1) why genetic isolation and genetic divergence lead to speciation, and 2) the roles that gene flow, natural selection, genetic drift, and mutation can play in speciation.

Speciation

All of the extraordinary diversity of life on the planet arose from what was at one time a single common ancestor. Over time, new specie have arisen through 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 in On the Origin of Species (Figure 7.1). Compare this illustration to the diagram of elephant evolution (Figure 7.1), 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 X I; 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.
Figure 7.1 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 today 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 form 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. Sympatric speciation  (sym- = “same”; -patric = “homeland”) involves speciation occurring within a parent species remaining in one location.

Reading Question #1

What are the two types of speciation?

A. Evolutionary and non-evolutionary
B. Allopatric and sympatric
C. Adaptive and non-adaptive
D. Radiation and linear

Allopatric Speciation

A geographically continuous population typically has a gene pool that is relatively homogeneous. Gene flow, the movement of alleles across a species’ range due to migration, is relatively free because individuals can move and then mate with individuals in their new location. Thus, an allele’s frequency at one end of a distribution will be similar to the allele’s frequency at the other end. When populations become geographically discontinuous, it prevents alleles’ free-flow. 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 increasingly 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 creating 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 organism’s biology 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 a new lake divided two rodent populations 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 spotted owl subspecies exist. The northern spotted owl has genetic and phenotypic differences from its close relative: the Mexican spotted owl, which lives in the south (Fig 7.2).

Figure 7.2 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 further 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.

Reading Question #2

Allopatric speciation can occur due to…

A. Dispersal
B. Vicariance
C. Sympatric speciation
D. A and B
E. A, B, and C

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. We call the process of speciation within the same space sympatric. The prefix “sym” means same, so “sympatric” means “same homeland” in contrast to “allopatric” meaning “other homeland.” Scientists have proposed and studied many mechanisms.

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 extra sets of chromosomes in a condition that we call polyploidy (Fig 7.3).

Figure 7.3 Aneuploidy results when the gametes have too many or too few chromosomes due to nondisjunction during meiosis. In this example, the resulting offspring will have 2n+1 or 2n-1 chromosomes.

Which is most likely to survive, offspring with 2n+1 chromosomes or offspring with 2n-1 chromosomes?

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 that we call autopolyploidy (Fig 7.4). 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 7.4 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 that this plant species produces. 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 that we call 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 that we call an allopolyploidy. The prefix “allo-” means “other” (recall from allopatric): therefore, an allopolyploid occurs when gametes from two different species combine. Figure 7.5 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 7.5 Alloploidy results when two species mate to produce viable offspring. In this example, 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. 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.

Sympatric speciation may also take place in ways other than polyploidy. For example, consider a fish species that lives in a lake. As the population grows, competition for food increases. Under pressure to find food, suppose that a group of these fish had the genetic flexibility to discover and feed off another resource that other fish did not use. What if this new food source was located 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 7.6 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.

Illustrations show two species of cichlid fish which are similar in appearance except that one has thin lips, and one has thick lips.
Figure 7.6 Cichlid fish from Lake Apoyeque, Nicaragua, show evidence of sympatric speciation. Lake Apoyeque, a crater lake, is 1800 years old, but genetic evidence indicates that a single population of cichlid fish populated the lake only 100 years ago. 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.

Question #3

Sympatric speciation can occur due to…

A. Polyploidy
B. Populations adapting to use different parts of the habitat
C. Dispersal and vicariance
D. A and B
E. A, B, and C

Adaptive Radiation

In some cases, a population of one species disperses throughout an area, and each 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. We call this 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, the founder species, numerous species have evolved, including the six in Figure 7.7.

Figure 7.7 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 7.7. 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: The Origin of Birds

Watch this video to see how scientists use evidence to understand how birds evolved.

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 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 ability 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 an organism’s development 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, which we call temporal isolation, can act as a form of reproductive isolation. For example, two frog species inhabit the same area, but one reproduces from January to March; whereas, the other reproduces from March to May (Fig 7.8).

Figure 7.8 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 same species’ other populations. We call this situation 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, natural selection forces, mutation, and genetic drift will likely result in the two groups diverging.

Behavioral isolation occurs when the presence or absence of a specific behavior prevents reproduction. For example, male fireflies use specific light patterns to attract females. Various firefly species 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. We call this a gametic barrier.

Similarly, in some cases closely related organisms try to mate, but their reproductive structures simply do not fit together, which is called mechanical isolation. For example, damselfly males of different species have differently shaped reproductive organs. If one species tries to mate with the female of another, their body parts simply do not fit together. (Fig 7.9).

Figure 7.9 The shape of the male reproductive organ varies among male damselfly species, and is only compatible with the female of that species. Reproductive organ incompatibility keeps the 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 cross-pollinating with a different species (Fig 7.10).

Illustration a shows a bee collecting pollen from a bright purple foxglove flower. The bee's body fits inside the bell like flower. Illustration B shows a hummingbird drinking nectar from a long tube like trumpet creeper flower.
Figure 7.10 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. We call this hybrid inviability because the hybrid organisms simply are not viable. In another postzygotic situation, reproduction leads to hybrid birth and growth that is sterile. Therefore, the organisms are unable to reproduce offspring of their own. We call this hybrid sterility.

Reading Question #4

Which of the following reproductive isolation mechanisms are prezygotic? Select all that apply.

A. Behavioral isolation
B. Hybrid infertility
C. Mechanical isolation
D. Hybrid inviability
E. Gametic barrier

Reconnection

Speciation occurs over a span of evolutionary time, so when a new species arises, there is a transition period during which the closely related species continue to interact. After speciation, two species may recombine or even continue interacting indefinitely. Individual organisms will mate with any nearby individual with whom they are capable of breeding. We call an area where two closely related species continue to interact and reproduce, forming hybrids, a hybrid zone. Over time, the hybrid zone may change depending on the fitness of the hybrids and the reproductive barriers (Fig 6.11). If the hybrids are less fit than the parents, speciation reinforcement occurs, and the species continue to diverge until they can no longer mate and produce viable offspring. If reproductive barriers weaken, fusion occurs and the two species become one. Barriers remain the same if hybrids are fit and reproductive: stability may occur and hybridization continues (Fig 7.11).

Figure 7.11 After speciation has occurred, the two separate but closely related species may continue to produce offspring in an area called the hybrid zone. Reinforcement, fusion, or stability may result, depending on reproductive barriers and the relative fitness of the hybrids.

If two species eat a different diet but one of the food sources is eliminated and both species are forced to eat the same foods, what change in the hybrid zone is most likely to occur?

Hybrids can be either less fit than the parents, more fit, or about the same. Usually hybrids tend to be less fit; therefore, such reproduction diminishes over time, nudging the two species to diverge further in a process we call reinforcement. Scientists use this term because the hybrids’ low success reinforces the original speciation. If the hybrids are as fit or more fit than the parents, the two species may fuse back into one species (Fig 7.11). Scientists have also observed that sometimes two species will remain separate but also continue to interact to produce some individuals. Scientists classify this as stability because no real net change is taking place.

Reading Question #5

When populations come back into contact after a period of reproductive isolation, which of the following is not a possible outcome?

A. Cycling
B. Reinforcement
C. Fusion
D. Stability

Varying Rates of Speciation

Scientists around the world study speciation, documenting observations both of living organisms and those found in the fossil record. As their ideas take shape and as research reveals new details about how life evolves, they develop models to help explain speciation rates. In terms of how quickly speciation occurs, we can observe two current patterns: gradual speciation model and punctuated equilibrium model.

In the gradual speciation model, species diverge gradually over time in small steps. In the punctuated equilibrium model, a new species undergoes changes quickly from the parent species, and then remains largely unchanged for long periods of time afterward (Fig 7.12). We call this early change model punctuated equilibrium, because it begins with a punctuated or periodic change and then remains in balance afterward. While punctuated equilibrium suggests a faster tempo, it does not necessarily exclude gradualism.

Figure 7.12 In (a) gradual speciation, species diverge at a slow, steady pace as traits change incrementally. In (b) punctuated equilibrium, species diverge quickly and then remain unchanged for long periods of time.

The primary influencing factor on changes in speciation rate is environmental conditions. Under some conditions, selection occurs quickly or radically. Consider a species of snails that had been living with the same basic form for many thousands of years. Layers of their fossils would appear similar for a long time. When a change in the environment takes place—such as a drop in the water level—a small number of organisms are separated from the rest in a brief period of time, essentially forming one large and one tiny population. The tiny population faces new environmental conditions. Because its gene pool quickly became so small, any variation that surfaces and that aids in surviving the new conditions becomes the predominant form.

References

Adapted from Clark, M.A., Douglas, M., and Choi, J. (2018). Biology 2e. OpenStax. Retrieved from https://openstax.org/books/biology-2e/pages/1-introduction

definition

License

Icon for the Creative Commons Attribution-NonCommercial 4.0 International License

Introductory Biology 2 Copyright © 2023 by Mason Tedeschi and Lisa Limeri is licensed under a Creative Commons Attribution-NonCommercial 4.0 International License, except where otherwise noted.

Share This Book