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    Speciation

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    Most biologists agree that a species is the smallest arily independent unit. Evolution occurs when mutation, tion, gene flow, and drift operate on populations separately. The challenge is to establish practical criteria for determining independence is in effect.

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    Speciation

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    Most biologists agree that a species is the smallest arily independent unit. Evolution occurs when mutation, tion, gene flow, and drift operate on populations separately. The challenge is to establish practical criteria for determining independence is in effect.

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    Most biologists agree that a species is the smallest arily independent unit. Evolution occurs when mutation, tion, gene flow, and drift operate on populations separately. The challenge is to establish practical criteria for determining independence is in effect.

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    129 views22 pages

    Speciation

    Uploaded by

    Κατερίνα Μέτσιου
    Most biologists agree that a species is the smallest arily independent unit. Evolution occurs when mutation, tion, gene flow, and drift operate on populations separately. The challenge is to establish practical criteria for determining independence is in effect.

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    584 PARTIV. The History of Life Species consiet of interbreeding populations that evolve independently of other populations and to a system for naming and classifying the diversity of life that acca flects the evolutionary history of organisms, These goals have been difficule to achieve. To understand why, iis imp realize that most biologists agree on what a species is: It is the smallest ei arily independent unit. Evolutionary independence occurs when mutta tion, gene flow, and drift operate on populations separately. Evolution changes in allele frequencies, and species form a boundary for the spread of As a result, different species follow different evolutionary trajectories this definition, itis clear that the essence of speciation is lack of gene flo Although this definition sounds straightforward, it is often difficult into practice. The challenge is to establish practical criteria for identiyiag populations are actually evolving independently: To drive this point homey sider the three most important “species concepts” currently in use. Ea thrce agrees that species are evolutionarily independent units that are i lack of gene flow, but each employs a different criterion for determining dependence is actually in effect. The Biological Species Concept Under the biological species concept, the criterion for identifying eveh independence is reproductive isolation, Specifically, if populations of org not hybridize, or fail to produce fertile offspring when they do, then the productively isolated and considered good species. The biological species @ has been the textbook definition of a species since Ernst Mayr 1942. is used in practice by many zoologists and is the in the Endangered Species Act, which is the flagship biodiversity lepisatiom United States Reproductive isolation is clearly an appropriate criterion for i species because it confirms lack of ary independence in organisms that reproduce sexually Althoug compelling in concept and useful in some situations, it i oft For example, if nearby populations do not actually overlap, we have m knowing whether they are reproductively isolated. Instead, biologist hl make subjective judgments to the effect that, "If these populations were in the future, we believe that they ate already that the not interbreed, so we will name them different species.” In these cise designations cannot be tested with data. Furthermore, the biological spe ‘cept can never be tested in fossil forms, is irrelevant to asexual populiil Box 15.1), nd is dificult to apply in the many plant groups where hybrid between strongly divergent populations is routine, The Phylogenetic Species Concept Systematists are biologists who are responsible for classifying the diversify A growing number of syscematists and evolutionary biologists are pron alternative to the biological species concept called the phylogenetic 4 cept—abo known as the genealogical species concept. This approach a criterion for identifying species called monophyly. Monophyletic gra defined as taxa that contain all of the known descendants ofa single com ccestor (Figure 15.2). A taxon, in turn, is any named group of organism population to a species or genus to an order or phylum. 586 PARTIV The History of Life Figure 18:2. Monophyltc groups The tara ltled A on Snter st etniaget cap eprentinyiannenlanneg bhyogenete ree se i meron ete hat monopyete cys eyoasiaiaarnpuaratcesel dep eaia peep tomer sasray aa gl pia [ that are circled Under the phylogenetic species concept, species are identified by estima phylogeny of closely related populations and finding ‘groups. Or tuce like this species form the tips. For example ifthe ta bee in Figure 15.2 represent populations (as opposed to genera or families or other highe level groups), then they are the smallest monophyletic grouped aly tree and represent distinct species. In contrast, if populations cannot be tinguished in a phylogeny by unique, derived characters, then they will like the populations designated B, E, and G in Figure 15.2. The popula make up these clusters would be considered part of the same species The rationale behind the phylogenetic species concept is that tit el distinguish populations on a phylogeny if the populations have been iso erms of gene flow and have diverged genetically and/or morphological another way, to be called separat phylogenetic species, populations mi been evolutionarily independent long en gh for the diagnostic tris synapomorphies to emer The appeals of this approach are that it eal plied to any type of organism—sexually reproducing, asexually reproduc fossil—and that it is testable: Species are named on the basis of statistic icant differences in the traits used to estifnate the phylogeny. The chal this criterion into practice.As Chapter Hil cated, it takes a significant amount of time, money, and arefil analysis 1 6 evolutionary relationships. Asa result, well-supported, carefully constructed pl nies are available for only a handful of groups thus fr. In addition, iis widely nized that institut the phylogenetic species concept could easly double number of named species and might create a g ns at deal of confusion if names and species identities are changed. Propc rents of the concept are noth by the later prospect, however. They respond by say the number of named species did occur, itis necessary to reflect biological ail The Morphospecies Concept Paleontologists define species on thi basis of morphological difference fossils, When rigorous tests of reproductive isolation or well-estimated pi nies are lacking, as they usually are, botanists and zoologists working of Chapter 15. Mechanisms of Speciation 587 ame. The great advantage of the morphospecies concept is that it Pplicble. Bur when ic is not applied carefully, species definitions can and idiosyncratic, In che worst-case scenario, species designa are not comparable WS have to work around other restrictions when identifying species that differed in color or the anatomy of soft tissues cannot { Neither can populations thac are similar in morphology but divergent in traits like songs, temperature or drought tolerance, ouuship displays. Whether living or fossil, populations ike these Blic species. The adjective cryptic is appropriate because groups le independent of one another appear to be members of the same fon morphological similarity. pecies Concepts: Two Case Histories Probably unrealistic to insise on a single, all-purpose criterion for ies (Endler 1989), the major species eoncepts that have been pro~ Bilictive when applicd in appropriate situations. Consider, for exam= advantages and Bt ellors to apply more than one species concept have improved rae Berry ia the mont abundant group of ocear-dweling “== formed efforts to preserve Afvican elephants in Marine Copepods Sinall crustaceans that are extremely abundant in the world’ Ges Eurytemora affins, for example, is only 1-2 mm long, but is the GA! grazing animal in many of the world’s langest estuaries (Figure jin turn, are ecosystems that form where rivers flow into the sea Bend to be nutrient rich, estuaries are among the most productive BaFine environments. In many cases, fish that spend their jivenile fon Eryemora and other small inhabitants of estuarine environ adult lives in the open ocean fins s Found along the coasts of Asia, Europe, and North Amer Analyses based on the morphospecies concept had grouped all of into the same species. To test this hypothesis, Carol Eunmi Lee E fins from a wide array of locations throughout the Nor assess whether some of the 38 populations in her sample rep fe species under the biological species concept, she tested individuals populations for the ability to mate and produce fertile offipring. To question with the phylogenetic species concept, she sequenced diused similarities and differences in the base sequences observed in fons to estimate their evolutionary relationships. BThe phylogeny clearly showed that a leat eight indepenclent species hic area (Figure 15,3b), These results by the results of the mating tests, which showed that populations Phylogenetic species are unable to produce fertile offipring and are Jy isolated. The punchline is that species diversity in copepods is Greater than previously thought. The study ako suggests several inter~ fallow-up research. Are the eight newly recognized species adapted to ofthe environments they occupy? If'so, are there any interesting Chapter 15 Mechanisms of Speciation 589 fr the species that these copepods feed on and the fish that feed on In this case, employing more than one species concept is helping biolo- ¢ diversity and organize research on its conseq Species of Elephant Live in Africa? ll, the world’s elephants have been classified as two species: Affican afc) and Asian (Elephas maximus). Recent morphological analyses challenge this view, however, by pointing out that African elephants that foresehabitars versus savanna or grassland habitats have distinctive mor- fl features (Figure 15.4a). But because the populations that inhabit the Bist types don't interact regularly, individuals rarely have a chance to in- {Asa result, it has been difficult to assess whether forest and savanna ele- ly as separate species under the biological species concept. fy the situation, Alfred Roca and co-workers (2001) applied the phy Fe species concept. They began by collecting tissue samples from 195 ele fin21 populations throughout central and south Africa. They then isolated Flom the tissues, sequenced four genes from each individual, and used the ggdata to estimate which populations were closely or more distantly relat- phylogeny that resulted from this analysis clearly showed that forest and flephants qualify as distinct phylogenetic species (Figure 15.4b). The lis proposed naming the forest elephants Laxodomta cyelotis and retaining Bie [.afriana for the savanna/grassland populations, result has a sense of urgency b nany elephant populations in Africa Btning due to habitat loss and illegal hunting. Based on this work, itis clear Remervation programs should be focused on preserving both forest and sa Populations as distinct entities [n this case, employing ceria for ing species proved to be a productive approach in clarifying both conser- fand evolutionary issues. Mechanisms of Genetic Isolation that several tools are available for identifying species, we can turn our at- Bi now to the question of how species form. Classically, speciation has been thesized to be a three-stage process: an initial step that isolates populations, a Bisep that results in divergence in traits such as mating system or habitat Bhd 2 final step that produces reproductive isolation. According to this the isolation and divergence steps were thought to take place over time Boccur while populations were located in diffe ygraphic areas. The ipise was hypothesized to occur when these diverged populations came favo physical contact—an event known as secondary contact. gent research has shown that the isolation and divergence steps that initi~ Kcntion frequently take place at the same time and in the same place. In fr, it appears likely that in a significant number of speciation events or G majority, the third phase never occurs. Even so, the isolation/diver- econdary contact hypothesis provides a useful framework for analyzing ciation takes place. is section we consider the first step in speciation—genetic isolation. We'll Bt how physical separation or changes in chromosome complements can re- fg flow between populations. Once gene flow is dramatically reduced or Reliable criteria for Identifying species are essential for preserving biodiversity. The epeciation process begine when gene flow is disrupted and populations become genetically isolated. 590 PARTIV. The History of Life Asian forest Atrican savanna lophants elephants Figure 15.4 Elephant diversity (a) In West Africa, elephants that ive in forest habitats (lft) have mil logical characterstis that distinguish them from savanna-dwlling elephants (ight) from west. central al Africa, (b) This evolutionary tree indicates that forest dwelling elephants area distinct phylogenetic spac Cases, evolutionary independence begins and speciation is underway. [nS 15.3 we ask how genetic drift, natural selection, and sexual selection act i tations and cause genetically isolated populations to diverge. In Section 15M consider what happens if secondary contact occurs. Physical Isolation as a Barrier to Gene Flow Chapter 6 introduced models showing that gene flow tends to homogesit frequencies and reduce the differentiation of populations. Using the exit Chapter 15 Mechanisms of Speciation 591 Bakes from mainland and island habitats in Lake Erie, we also introduced. fofa balance becween gene flow and natural selection. Recall that exp. iad shown a selective advantage for unbanded snakes on island habitats jon of banded forms from the m: inland occurs regularly, and banded and unbande. forms subsequently interbreed, the island popula- fiinot completely diverge from mainland forms. Mig fed alleles for bancdedness, fend populations ation continually in- even though selection tended to eliminace them Konsider a thought experiment: What would happen if lake currents ina way that effectively stopped the migration of banded forms from the Bit the islands? Gene flow between the two populations would end and Gation-

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