2/9/12

Species Concepts and Variation Within Species

Species Concepts and Variation Within Species

NOTE: These are lecture notes for Biology 391, Organic Evolution, at The University of Tennesee at Martin. Anyone outside of UT Martin wishing to use these notes or to contact me for additional information should first read the information obtained by clicking here. Goals: introduce the different concepts that biologists have of what a species is, and introduce some of the forms of variation within species that can make it difficult to define species. Related Textbook Material: Freeman and Herron (2001) Chapter 12 Lab Manual Questions over this material are in Lab Manual Chapter XIII

The Lecture:
In the lectures so far, throughout the term, we have studied how evolution occurs within a species and we have studied phylogenies, which show the history of speciation, but we have not yet looked at how speciation occurs -- how one species becomes divided into more than one new species. Speciation will be the focus of the next few lectures. Before we can study speciation, we need to learn what a species is. Defining a species turns out to be somewhat difficult. Currently, different biologists have developed a variety of different definitions of species. At a conference I went to recently someone had a list of over thirty definitions. We won't learn all of those! Instead, we will study why it can be difficult to define species and then focus on two of the main current definitions of species. We care about what a species is for several reasons, both in terms of understanding evolution and for more practical reasons. For the purpose of understanding evolution, many biologists argue that a species is a fundamental unit of evolution -- that evolution of characteristics occurs within a species so that a species can be thought of as the unit that is evolving. Different species then evolve independently from one another. The formation of new species is a major evolutionary event because it splits a group that was evolving as a unit into more than one separately evolving units. Practically, defining species can have legal implications with regard to the Endangered Species Act. Whether or not we take action to conserve a particular group may depend on whether or not it is considered to be a separate species from other, similar groups. This can affect the future of these organisms and the economics of the people who live in the area where these organisms occur. Historically, before people started studying evolution, when species were thought to be unchangeable, people defined species simply as all individuals of a certain type, and would describe a type specimen of each species and categorize everything that looked like that type as belonging to the same species and everything different as belonging to other species. To some extent, this is still done -- biological museums keep "type specimens" of species to help categorize them, but it is recognized that there is a major problem with this kind of definition of species. We can't define species based on one description of what they should look like because individuals within species vary. We have seen that there is variation within species; it is the basis for evolution within species. To understand the problems we may have in defining species, we need to consider some of the different ways

www.utm.edu/departments/biology/rirwin/391/391Species.htm

1/5

2/9/12

Species Concepts and Variation Within Species

in which species can vary. Some of these kinds of variation are: Polymorphism refers to variation within a population where different individuals in the population have distinctly different structures, colors, biochemistry, etc. but clearly belong to the same population since they reproduce with one another. Geographic variation refers to variation over geography -- that is, populations in different areas look different from each other. There may not be any obvious pattern to such variation but sometimes there is. One common pattern of geographic variation is clinal variation. Clinal variation refers to a gradual change in some feature across geography. For example, if you go from south to north in the northern hemisphere you frequently find that populations of mammals are smaller in the south and gradually as you go north you find that within a species the individuals are larger and larger. Such a pattern of gradual change is called a cline so this form of variation is called clinal variation. Hybrid zones are geographic areas where two distinctly different forms of organism contact each other and interbreed. Generally, outside this zone, the two forms retain distinct differences from one another but within the zone intermediate forms between the two occur as a result of interbreeding. For example, a kind of woodpecker, flickers, show a hybrid zone in the United States. Western forms are called "red-shafted flickers" because the undersides of their wing and tail feathers (the "shafts") are red; eastern forms are called "yellow-shafted flickers" because the undersides of their wing and tail feathers are yellow. Flickers occur from the west coast to the east coast of the United States; throughout the west they are red-shafted and throughout the east they are yellow shafted. In the middle of the country there is an area just a few hundred miles wide where the two forms both occur and where they reproduce with each other. There, some flickers have red shafts, some yellow, and some have some red and some yellow. There are also DNA markers that have been identified such that all eastern forms are fixed for one DNA form and all western forms are fixed for a different DNA form but in the hybrid zone both forms occur.

Hybrid zones raise some interesting evolutionary questions. Since the two forms can reproduce with each other, we would expect gene flow to occur from one form to the other. If gene flow occurs, we would expect the traits from one form to spread into the other form. We observe, however, that the forms on the two sides of the hybrid zone retain fixed differences. How is this possible? There are several hypotheses that could explain the maintenance of stable hybrid zones -- that is, explain how it is possible for a hybrid zone to exist over time without there being spread of the characteristics of the form on one side of the zone into the form on the other side of the zone. Two of these hypotheses are as follows: 1. The environment may be such that hybrids (organisms produced by reproduction of the two main forms) have high fitness in the area of the hybrid zone but low fitness outside zone. If this is the case, then hybrids that move out of the zone will have low fitness and selection against them may counteract the effects of gene flow out of the hybrid zone. 2. The environment where the hybrid zone occurs may be very poor quality habitat for either form or for hybrids. As a result, individuals may move into the zone from either side, but will not reproduce well enough within the zone to produce many individuals that move out of the zone. If this is the case, there will be very little gene flow out of the zone and fixed differences between the forms on either side of the zone can be maintained. Biologists generally agree that we do not want to categorize the different forms in a polymorphic population

www.utm.edu/departments/biology/rirwin/391/391Species.htm

2/5

2/9/12

Species Concepts and Variation Within Species

as different species, because they reproduce with each other and offspring of the same parents could have different forms. We generally do not want to call different populations that show clinal variation different species since the populations change gradually from area to area and individuals can reproduce with individuals in slightly different neighboring populations, so there would be no clear place to draw a line to distinguish species and overall the different populations can clearly affect one another by reproducing with one another. But what about the different forms on either side of a hybrid zone? On one hand, they retain distinct differences from one another so it does not appear that they are affecting each other evolutionarily; based on this argument, people would consider them as belonging to different species. On the other hand, they reproduce with each other within the hybrid zone, and their offspring are healthy and fertile, so it appears that they are very similar and that they could affect each other, so based on this argument people would consider them as belonging to the same species. Examples such as this show why it can be difficult to categorize groups as species. We will discuss some of the main concepts of what a species is and consider how they would categorize groups from different examples as either the same or as different species. Biologists differ on whether or not they view species as fundamental evolutionary units. Some biologists argue that evolution occurs primarily within populations. There is evidence in a number of species that different populations evolve fairly independently from one another. If that is the case then it may not be important how we define species; it is determining what groups make up the populations within which evolution is occurring that is important. Other biologists point out that populations do affect each other, genetically -- gene flow occurs -- and argue that we want to define species as the groups that are evolving, the fundamental units of evolution. These are the biologists who really care about species concepts. They do not, however, all agree on which concept of species is the best. Two of the main concepts of species are as follows: 1. The biological species concept (BSC): categorizes species as groups of actually or potentially interbreeding organisms that are reproductively isolated from other such groups. In other words, organisms are considered to belong to the same species if they can potentially reproduce with each other and have offspring that are healthy and fertile. Otherwise, they are considered to belong to different species. 2. The phylogenetic species concept (PSC): categorizes a species as the smallest group that can be diagnosed with a derived character state that clearly separates it from other such groups. In other words, if you have a group of organisms that all share a derived character state and that can not be subdivided into smaller groups based on shared derived character states then you have a species. Different species have different derived character states from each other. Both these concepts of species try to identify the groups that will evolve as independent evolutionary units. The biological species concept is based on the idea that if gene flow can occur between groups (because they can reproduce and produce healthy, fertile offspring) then these groups can affect each other genetically and should be grouped into a single species. The phylogenetic species concept is based on the idea that we can tell which groups are in fact affecting each other genetically by their characteristics -- if they share a unique derived characteristic not found in other groups, then they are not spreading this characteristic to other groups; if they do not have the derived characteristics found in other groups, then the other groups are not affecting them. There are potential problems with each concept that are solved by the other concept. These follow: The biological species concept can not categorize asexual species (since they do not reproduce with other individuals at all) or fossil species (since we can not tell what fossils could have reproduced with what other

www.utm.edu/departments/biology/rirwin/391/391Species.htm

3/5

2/9/12

Species Concepts and Variation Within Species

fossils now that they're all dead.) The phylogenetic species concept, since it is based on characteristics other than reproduction, provides a basis for classifying such species. The biological species concept lumps into a single species some groups that are very ecologically and structurally different and that are apparently evolving separately. For example, there is a group of plants on the Hawaiian islands called the silverswords: some are vines, some are shrubs, some are small plants, some are trees. All forms can apparently reproduce with all other forms, but the different forms are clearly distinct from each other and evolving independently. They appear to be different species, but the biological species concept would categorize all as belonging to the same species because they can reproduce with each other. In contrast, the phylogenetic species concept classifies them as different species because each group has its own unique derived characteristics. Similarly, the biological species concept would categorize two groups that are different from each other but form a hybrid zone, like the flickers described above, as belonging to the same species despite their distinctive differences, while the phylogenetic species concept would categorize them as different species because of their distinctive differences. The phylogenetic concept may thus better reflect, in these cases, groups that are evolving independently from each other. The biological species concept may group organisms that are not each others closest relatives into a species. For example, in a group of birds called orioles, two forms, the Baltimore oriole and the Bullock's oriole, can reproduce with each other and produce healthy, fertile offspring. Phylogenetic analysis of these orioles, however, suggests that Bullock's orioles are more closely related to other species of oriole, with which they can not interbreed, than to Baltimore orioles. This kind of situation can arise because the ability to interbreed is primitive within these species -- originally, they belonged to the same species and reproduced with each other. More recently, some species have derived traits such that they can't reproduce with other groups, but some retain the primitive ability to reproduce with each other. Species categorized based on the phylogenetic species concept should avoid this problem. They are categorized based on derived characteristics and these, as you learned in the lectures on systematics, reflect phylogenetic relatedness. So the phylogenetic concept should group organisms into species in which all members are more closely related to each other than to other species. The phylogenetic species concept thus reflects evolutionary history better than does the biological species concept. There are also problems with the phylogenetic species concept, though. The phylogenetic species concept may divide species into groups based on characteristics that do not have any clear biological relevance and appear to be very minor traits. The phylogenetic concept may divide life up into an inconveniently large number of species. Probably the most serious problem with the phylogenetic species concept is that it may divide groups into species that are not going to remain separate species because the groups can still reproduce with each other. Thus, the species defined by the phylogenetic species concept may not be permanent -- they may merge together over time. The biological species concept avoids these problems. Now that we've considered the pros and cons of these concepts, it is important to note that most of the time they agree on what they define as species. Any time two groups are so different that they can not reproduce to produce healthy fertile offspring, both concepts consider them to be separate species. The biological concept considers them separate species because the ability to reproduce is the criterion this concept uses to group organisms into the same species. The phylogenetic concept considers them separate species because if two groups can not reproduce with each other then they must have some different derived traits that prevent them from reproducing. So the two concepts only differ where there are groups of organisms that can reproduce with each other and produce healthy, fertile offspring but have distinctly different derived traits. When this occurs, the biological concept lumps these organisms into the same species, and the phylogenetic concept splits them into separate species. This occurs among very closely related groups that may just be in the process of speciating. It is to be expected that as speciation occurs we will find some

www.utm.edu/departments/biology/rirwin/391/391Species.htm

4/5

2/9/12

Species Concepts and Variation Within Species

groups that are hard to categorize as still belonging to one species or having divided into two species. The arguments about species concepts reflect this. Now that we have some idea (or different possible ideas) of what a species is, we can look at the process of speciation. We'll do that in the next lecture. Click here to return to the index of lectures

www.utm.edu/departments/biology/rirwin/391/391Species.htm

5/5