THE DOMAINS OF LIFE 15.14 Arranging life into kingdoms is a work in progress In the next unit, we examine the that have populated Earth since - life first arose over 3.5 billion years i. help us trace the evolutionary rela- tionships among the organisms. Phylogenetic trees are hypothe- ses about evolutionary history Like all hypotheses, they are revised, or in some cases com- rejected, in accordance ew evidence. Molecular sys- tematies and cladistic analy combining to remodel phylog netic trees and challenge conver tional clasifiations, even at the kingdom level Over the years, many schemes eas have been proposed for clasiyi organisms into kingdoms, A two- Fighte 18.148. The fve kingdom clssitcation scheme kingdom system that divided all organisms into plants and animals was first proposed by Linnaeus and was popular for over 200 years. But it was beset with prob- Jems. Where do prokaryotes fit such a system? Where do unicellu- lar, photosynthetic protists that \ rove like animals belong? And what about the Fung? In 1969, American ecologist Robert H, Whittaker argued elfec- tively for a ive-kingdom system that places prokaryotes inthe kingdom Monera (Figure 15.14), Organisms of the other four king dom all consist of eukaryotic cell Kingdoms Plantue, Fungi, and Animalia consist of multicellular eukaryotes that generally differ in structure, development, and modes of nutrition. Plants make their own food by photosynthesis, FIQUe 15.148. The theedomain assifcaton scheme Fungi live by decomposing the remains of other organisms and absorbing small. organic It is important to keep in mind that d molecules, Most animals live by ingesting food and digesting schemes, including the five-kingdom system, are nat it within their bodies. nature, but human constructions. The five-kingdom Earliest ‘organisms The kingdom Protista, containing all eukaryotes that do was one attempt to classify the diversity of if ints not fit the definition of plant, fungus, or animal, is a taxo- that is useful and reflective of evolutionary histo nomie grab bag in the five-kingdom system. Most protists last decade, molecular studies have highlighted seria are unicellular, but Whittaker’ kingdom Protista also in the five-kingdom system, and most systematst i includes relatively simple multicellular organisms that are replacing it with classification schemes that are mote believed to be direct descendants of unicellular protsts with these new data. 32 UNIT TET Concepts of Evolutionure 15.14B shows the three-domain system, an alternative to the five-kingdom system that has gained sup. port from many biologists in recent years. Whereas the five Kingdom system reflects the concept that there are two fimdamentally different groups of living organisms— prokaryotes and eukaryotes—this newer scheme recognizes three basic groups: two domains of prokaryotes, Bacteria and Archaea, and one domain of eukaryotes, called Eukarya, Bacteria and Archaea differ in a number of important strac- tural, biochemical, and funetional features, which we will discuss in Chapter 16. Molecular and cellular evidence supports the phyloge- ‘etic hypothesis outlined in Figure 15.14B—that two lin- ‘ages of prokaryotes (the bacteria and the archaea) diverged very early in the evolutionary history of life. Molecular evi- tence also suggests that the archaea are more elosely related tb eukaryotes than to the bacteria. Thus, Figure 15.14B indicates that the lineage of the domain Eukarya diverged fom Archaea alter the archaeans and bacteria diverged. ‘The color coding in Figures 15.144 and 15.14B high- Juhi the relationship between the five-kingdom system and the three-domain system. Notice that all the eukaryotic Kingdoms are included in the domain Eukarya. As we dis- Chapter Review HAPTER SUMMARY Jirth History and Macroevolution (Introduction-15.9) Desi record dacuments macroevolution, the main events inthe his ini. Majo transitions in life-forms separ geologic ers: smaller fing divide eras nto periods. Life begun inthe Precambrian era, at of which aninals frst appeared. During the Paleozic era, plans nal diversified and moved ooto land. During the Mesoane era ths and cone- bearing plants dominated the land. Extinction of the ess arked the beginning ofthe Cenoaoie era, during which mam: a Bowering plants proliferate sata indictes te relative ages of ei . The sequence of fos in at species, Radiometric fi, which measures the decay of radioactive isotopes, can gauge the gs offs (152) CGntnental cri the slow, incessant movement of Fath’ erastal ssa the hot mantle, hs nlenced the distribution of ongunisms and palace the bistony of hie stot near the end of the Permian period, forming the super: tnentl dri rout the com et Page, This continental merger altered coastlines an climate lege extintions. Early in the Mesozole era, separation af the jets eased the sation and diversification of organisms (5.2). ses, the movements of Earths erstal pats, ae also assoc sh okanoes and earthquakes (15-1 ie end ofthe Cretaceous perind, many ife-forms disappeared age dinosaurs. These mass extinctions may have reslted ym the impact of an asteroid or from aa increase in voleanic rey mass extinction reduced the diversity of fe, but each was dys rebound in diversity: Mammals filed the voi left by the 1 (155). Dave features, such asthe hair and mammary glands of mar uy enable particular organisms to prosper after mass extinctions Js ao called exaptations) that have evolved in one environ cess further in the next chapter, the number of kingdoms and their evolutionary relationships are subjects of ongoin research and debate. For example, nucleic acid evidence indicates that kingdom Protista, as defined in the five- kingdom system, contains several groups of eukaryotes that are so phylogenetically distinct that they should be classified in separate kingdoms. ‘A strong consensus favors replacing the five-kingdom sys tem, and we follow the domain scheme outlined in Figure 15.148 in this textbook. However, what is most important to understand is that defining the higher categories of clssfi- cation (the kingdoms and domains of life) will always be a ‘work in progress, The most important function of a classifi cation system is to serve as a focal point for discussion about hhow to make clasifcation fit phylogeny. Web/CD Activity 15D Classification Schemes EB] = comparin the vrkngdom sytem wth ho tro: domain system, how many ofthe kingdoms fll into domain Eukarya? nog ‘mental context may beable to perform new funetions when conditions ‘change (15.6), Major acaptatons may arise rapidly mutations occur in certan genes that contol early events in an organism's development ?Paedoanorphoss, the retention of juvenile characteristics in the adult seents to ave payed key ole in human evolution "Bvoevo" ia reatarch fl that combines eolionary and developmental biology (15.7) The fossil record seems to reveal tends inthe evoltion of species aml line such a a grad sizeinerease inthe evolution of horses. An evolutionary trend may ceflect unequal speciation or unequal survival of species on a branching evolutionary tee (15.8). Phylogeny, which ean he depicted by piylogenetic tees, isthe evolutionary history of a group of organisms (15.9). Systematics and Phylogenetic Biology (15.10-15.13) Reconstructing pylogeny i par of systematics, the study of b Aliversity and clasifcation. Taxnomist assign a bxo-part name, oF bino tal to each species, The fst name, the gen ted covers a group of re sees. The second name refers toa species within the genus. Genera are grouped into progressively larger categories: family, order cas, phy Tum, king and domain, Taxonomists often debate the particular placement of organisms in ete fores reflect evolutionary relationships (15.10)- res as they steve to make their ete omologons structures tar evidence that organisms have evolved from a common ancestor In Contrast, nouhomlogs (analogous) similarities are evidence that organ isms fom different ewoationaryTineages have undergone convergent ‘volation Their resemblances have rested from Ising in similar env ronments, not from common ancestry (15.11). Sysematit increasingly tse molecular techniques (especially DNA and RNA sequence analyses) to clas organisms and develop phylogenetic hypotheses (15.12). The sg of systematics sto make classification reflect evolutionary history (plnlogeny). Using homologons features to compare organisms, clastic analysis attempts to deine monophyletic taxa, groups consisting of an Chapter 15 ‘Tracing Evolutionary History 313