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    Intro to Zoology - Evolution, Hickman

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    Zoology Lecture Chapter 6

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    Intro to Zoology - Evolution, Hickman

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    of 34
    Sede Prete igo CHAPTER A Legacy of Change Life's history is a legacy of perpetual change. Despite the apparent per- ‘manees ofthe atl word, ching charter lhings on earth tndin he univer. Ears ok trata eord the ever socal Change tat we cll organic evolution. Cues indsof animals and plants hve forshd and disappeared ving bend a spre fossil Fecordof tir existence. Many. bt nota have lft ving descendants thabear some resemblance them "We observe i's changes and meas them in many ways On shor eroltonary timescale, we se changes in the equencies of di fen pencils win population. Evolutionary changes in the ‘ave reqencis oh an colored maths octred within a Single man lite inthe poli towns of india England, The Organic Evolution ‘ormation of new species and dramatic changes in organismal fom, 25 illustrated by the evolutionary diversification of Hawatian birds. equies long timescales covering 100,000 to | millon years. Major evolution ary trends and episodic mass extinctions occur on even larger timescale, covering tens of million of years. The fossil record of horses though the past SO million years shows a series of different species replacing ‘older ones though time and ending with the horses alive today. The fossil record of marine invertebrates shows episodic mass extinctions separated by intervals of approximately 26 million years. Because every feature of life as we know it today isa product of evolution, biologists consider organic evolution the keystone of all bio- logical owiedge. 100 PARTTWO Continity and Evolution of Animal ite ‘Chapter 1, we introduced Darwinian evolutionary theory a the ‘dominant paradigm of biology. Charles Robert Darwin and Alfred Russel Wallace (Figure 6.1) first established evolution as «8 powerful scientific theory. Today the reality of organic evolution ‘can be denied only by abandoning reason. As the noted English biol- ‘gist Sir Julian Huxley wrote, “Charles Darwin effected the greatest, ‘of allevolutions in human thought, greater than Einstein's or Freud's, ‘or even Newion’s, by simultaneously establishing the fact and dis- ‘covering the mechanism of organic evolution.” Darwinian theory helps us to understand both the genetics of populations and long- term trends inthe fosil record. Darwin and Wallace did not origi- nate the basic idea of organic evolution, which has an ancient history. We review the history of evolutionary thinking a it led to Darwin's ‘theory, evidence supporting it, and changes tothe theory that have produced our modern Synthetic Theory of evolution 6.1 ORIGINS OF DARWINIAN EVOLUTIONARY THEORY Pre-Darwinian Evolutionary Ideas Early Greek philosophers, notably Xenophanes, Empedocles, and Aristotle, recorded an early idea of evolutionary change. They rec- ‘ognized fossils as evidence for former life that they believed had beeen destroyed by natural catastrophe, Despite their inquiry, the Greeks failed to establish an evolutionary concept. Opportunity for ‘evolutionary thinking became even more restricted as a Titeally interpreted biblical account ofthe earth's creation became accepted as tenet of faith. Archbishop James Ussher (mid-seventeenth cen- tury) declared the year 4004 cE as the date of life's creation, Evolutionary views were considered rebellious and heretical, but they refused to die, The French naturalist Georges Louis Buffon (1707 to 1788) stressed the influence of environment on the Figure 6.1. Founders of the theory of evolution by natural selocton , Chas Robert Darwin (1809 to 182). B, Are Russel Wallace (1823 to 1913} n 1895. Darwin and Wallace Independently developed the same theory. Aleter and essny fom Waface written to Darwin in 1858 spurred Darwin into writing On the Origin of Species, published in 1858, ‘modifications of animal form. He also extended the age of the earth to 70,000 years. Lamarckism: The First Scientific Explanation of Evolution French biologist Jean Baptiste de Lamarck (1744 to 1829; Figure 6.2) authored the frst complete explanation of evolution in 1808, the year of Darwin's birth. He made a convincing case that fossils were ‘remains of extinct animals. Lamarck’s proposed evolutionary mech- anism, inheritance of acquired characteristics, was engagingly simple: organisms, by striving to meet the demands of theirenviron- ‘ments, acquite adaptations and pass them by heredity to their off spring. According to Lamarck, the giraffe evolved its long neck because its ancestors lengthened their necks by sireching to obtain, {ood and then passed the lengthened neck to their offspring. Over ‘many generations, these changes accumulated to produce the long, necks of modern giraffes. We call Lamarck’s concept of evolution transformational, ‘because it claims that as individual organisms transform their char acteristics through the use and disuse of body parts, heredity makes corresponding adjustments to produce evolution, We now reject transformational theories because genetic studies show that tats acquired by an organism during is lifetime, such as strengthened ‘muscles, are not inherited by offspring. Darwin's evolutionary the- ‘ory differs from Lamarck’s in being a variational theory, based in ‘genetic differences that occur among organisms within a population, Evolution occurs at the level of the population, and it includes ‘changes across generations inthe organismal characteristies that pre- vail in the population. Darwin argued that organisms possessing. hereditary characteristics that conferred an advantage for survival reproduction would leave more offspring than would other organ- ‘sms, thereby causing the characteristics most favorable to the sur- vival and reproductive success of their bearers to accumulate in populations across generations. Charles Lyell and Uniformitarianism ‘The geologist Sir Charles Lyell (1797 to. 1875: Figure 6.3) established in his Principles of Geology (1830 to 1833) the principle of uniformitarianism. ‘Uniformitaranism encompasses two important princi ples that guide scientific study of the history of nature: (A) that the laws of physies and chemistry have not changed throughout the history ofthe earth, and (2) that fret geological evens occured by natural processes similar to those observed today. Lyell showed that nat- Fura forces, ating over long periods of time, could HF explain the formation of fossil-bearing rocks. For cal natal enn of cor p27) rr miniferans (p. 238), and molluses (p. 333) accumulate FE on the sea floor, they form sediments of calcium car J bonate that eventually become compressed into lime- © stone. Lyell's geological studies led him to conclude that the earth's age must be measured in hundreds of nillions of years. Measured rates of sedimentation are uch t00 slow to have produced earth's sedimentary rock formations in a shorter period of time. These wwembhe.com/hickmanips] 7e © sone Soc Figure 6.2 Jean Baptiste do Lamarck 174 o 1828), French naturals. wo offered the frst selene explanation of evoluon Lamerk’s hypothesis tha evolton proceeds by inhetance of ecaured _horocerisies has ber rejected and eplced by neo-Dorwinion theories CCHAPTER® Organic Evaltion Xu Huon Archive images Figure 6.3. si Chares ty (787 to 1875), Engh geologist and fiend of Darn His book rncples of Geology great influenced Darwin duting Dan's formative period Figure 6.4. theyeer voyage of IMS. Beogle. principles were important for discrediting miraculous and supernat- ural explanations of the history of nature and replacing them with scientific explanations. Lyell also stressed that geological changes. ‘occur primarily in small increments whose gradual accumulation through time builds the earth’s major geological formations, and he argued further that such changes have no inherent tendency to occur in any particular direction. Positions of mountains and seas would. ‘gradually change through time, for example, but the earth's surface would have no directional tendency toward becoming more moun- tainous or more inundated. Both ofthese claims left important marks ‘on Darwin's evolutionary theory. Darwin’s Great Voyage of Discovery “After having been twice driven back by heavy southwestern gales, Her Majesty's ship Beagle, a ten-gun brig, under the command of Captain Robert FitzRoy, R.N., sailed from Devonport on the 27th of December, 1831." Thus began Charles Darwin's account of the his toric five-year voyage of the Beagle around the world (Figure 6.4) Darwin, not quite 23 years old, had been asked to accompany Captain FitzRoy on the Beagle, a small vessel only 90 feet in length, ‘which was about to depart on an extensive surveying voyage to South America and the Pacific Ocean (Figure 6.5). It was the 102 PARTTWO Continuity and Evoltion of Arima Life Figure 6.5. che cousin. Ene Wess ‘wos posted by Com beginning of the most important scientific voyage ofthe nineteenth ‘century. ‘During the voyage (1831 to 1836), Darwin endured seasickness and the erratic companionship of Captain FitzRoy, but Darwin's youthful physical strength and early training as a naturalist equipped him for his work. The Beagle made many stops along the coasts of South America and adjacent islands. Darwin made extensive collec- tions and writen accounts ofthe fauna and flora ofthese regions. Hee unearthed numerous fossils of animals long extinct and noted ‘the resemblance between fossils ofthe South American pampas and the known fossils of North America. In the Andes he encountered seashells embedded in rock at 13,000 feet. He experienced a severe ‘earthquake and watched mountain torrents that relentlessly wore away the earth. These observations, and his reading of Lyell’s Principles of Geology during the voyage, strengthened his convi tion that natural forces could explain the geological features of the earth In mid-September of 1835, the Beagle arrived atthe Galspagos Islands, a voleanie archipelago straddling the equator 600 miles west of Beuador (Figure 6.6), The fame of the islands stems from their oceanic isolation and rugged voleanic terran, Circled by capri- cious currents, surrounded by shores of twisted lava bearing skeletal brushwood baked by the equatorial sun, inhabited by strange rep- tiles and by convicts stranded by the Ecuadorian government, the islands had few admirers among mariners. By the middle of the seventeenth century, the Spaniards called these islands “Las Islas ‘Galspagos"—the tortoise islands. The giant tortoises, used for food first by buccaneers and later by American and British whalers, seal- ers, and ships of war, were the islands' principal attraction. At the time of Darwin's visit, the tortoises already were heavily exploited. During the Beagle's five-week visit to the Galspagos, Darwin ‘documented the unique character of the Galépagos plants and ani mals, including the giant tortoises, marine iguanas, mockingbirds, and ground finches. Darwin later described these studies as the “ori- gin of all my views.” 1 Derwin and HM. Beagle. A, Darwin in 1840, four years ater the Beagle retumed to England, and ¢ year ater hs marioge to hs "The HMS, Beagle sl n Beagle Channel, Terra de! Fuego, onthe southern tip of South Amica Martens. one oftwo oficial artists onthe voyage ofthe Beagle, "822, The watercelor i i i Figure 6.6 The Galipagos lands wowed from ther ofa voleano, Darwin was struck by the fact that, although the Galépagos Islands and the Cape Verde Islands (visited earlier inthis voyage of the Beagle) were similar in climate and topography, Galfpagos plants and animals resembled most closely those of the South ‘American mainland, and they were entirely different from the African-derived forms of the Cape Verde Islands. Exch Galipagos Island often contained a unique species that nonetheless resembled forms on other Galipagos Islands, In short, Galapagos life must have ‘originated in continental South America and then undergone modifi ‘ation in the various environmental conditions of the different islands, He concluded that living forms were neither divinely created ‘nor immutable; they were, infact, products of along history of evo- lutionary change. ‘On October 2, 1836, the Beagle returned to England, where Darwin conducted most of his scientific work (Figure 6.7). Most of Darwin's extensive collections had preceded him thee, as had note- books and diaries kept during the cruise. Darwin's journal, published three years after the Beagle’s return to England, was an instant Figure 6.7 Dans study at orm House in Ken Englan, preserved today much as Rwes wen Dense vate On the Orig of Species success and required two additional printings within the frst year Darwin late revised his journal as The Voyage of the Beagle, a trav- logue of continuing popularity. ‘The main product of Darwin’s voyage, his theory of evolution, would continue to develop for more than 20 years after the Beagle's return. In 1838, he “happened to read for amusement” an essay on populations by T: R, Malthus (1766 to 1834), who stated that animal and plant populations, including homan populations, have the epro- ductive capacity to increase beyond the capacity of environmental resources to support them. Darwin already had been gathering infor- ‘mation on artificial selection of animals under domestication, Darwin was especially fascinated by artificial breeds of pigeons. ‘Many pigeon breeds differed so much in appearance and behavior that they would be considered different species if found in nature, yet, all had clearly been derived from a single wild species, the rock pigeon (Columba livia. After reading Malthus's article, Darwin realized that a process of selection in nature, a “strugle for exis- tence” because of overpopulation, could be a powerful force for ‘evolution of wild species. Darwin allowed the idea to develop in his own mind, writing private, trial essays in 1842 and 1844. Finally in 1856, he began to ‘assemble his voluminous data into a work on the origin of species. He expected to write four volumes, a very big book, “as perfect as ean make it.” However, his plans took an unexpected turn. In 1858, he received a manuscript from Alfred Russel Wallace (1823 to 1913), an English naturalist in Malaya with whom he corre- sponded. Darwin was stunned to find that in a few pages, Wallace ‘summarized the main points of the natural selection theory on which Darwin had been working fortwo decades. Rather than withholding his own work in favor of Wallace's as he was inclined to do, Darwin, ‘was persuaded by two close friends, the geologist Lyell and the bot- Aanist Hooker, to publish his views in brief statement that would ‘appear together with Wallace's paper inthe Journal of the Linnean ‘Society. Portions of both papers were read to an unresponsive audi- fence on July 1, 1858. For the next year, Darwin worked urgently to prepare an “abstract” of the planned four-volume work. This book was pub- lished in November 1859, withthe ttle On the Origin of Species by ‘Means of Natural Selection, or the Preservation of Favoured Races (CHAPTER 6 Orgone Evolution 103 in the Straggle for Life. Toe 1250 copies ofthe first printing sold the first day! The book instantly generated a storm that has never abated. Darwin's views were to have extraordinary consequences on scien- tific and religious beliefs and remain among the greatest intellectual achievements ofall time, “Whenever | have found that | have blundered, or that my work has been imperfect, and when I have been contemptuously criticized, and even when Ihave been overpraised, 0 that have felt mortified, ithas been my greatest comfort to say hundreds cof times to myself that have worked as hard and as wells could, and ne man can do more than this.” Chorls Darwin, in his cutbiorphy, 1876 (Once Darwin's caution had been swept away by the publication (of On the Origin of Species, he entered an incredibly productive period of evolutionary thinking for the next 23 years, producing five revisions of On the Origin of Species and a dozen new books. He continued his scientific interactions with Wallace, who carefully documented the geographic distributions of animal and plant spe- cies and thereby founded the field of historical biogeography (p. 798). Darwin died on April 19, 1882, and was buried in Westminster Abbey. The litle Beagle had already disappeared, having been retired in 1870 and sold for scrap. 6.2 DARWINIAN EVOLUTIONARY THEORY: THE EVIDENCE Perpetual Change ‘The main premise underlying Darwinian evolution is that te living worlds neither constant nor perpetually ycling, but alvays chang- ing and with hereditary continuity from past to presen life. Perpetial change inthe form and diversity of animal lie troughout its 600-0 700-milion-year history is seen most directly inthe fossil record. A fossils remnant of past life uncovered from the erst ofthe earth (Figure 68), Some fosils constitute complete remains (insets in amber and mammoth), actual hard parts (eth and bones). and pet sified skeletal parts infiltrated with silica o other minerals (ostaco- derms and molluscs). Other fossils include footprints or other impressions, burrows of marine worms in sediment on the seafloor, and fossil excrement (coprolites). In addition to documenting ogan- ismal evolution, fosils revel profound changes in the earth's envi- ronment, including major changes in the distributions of lands and seas, Foils formed on the floors of ancient seas can be quaried high atop current mountains (p. 105), Fessi remains may on rare occasions include soft tissues preserved so well that recognizable cellular organelles are revealed by electron microscopy! Insects are frequently found entombed in amber, the forsized resin of trees. One study of 2 fy entombed in 40-millin-year-ld amber revealed structures corresponding to muscle fibers, nck ribosomes, pid droplets, endoplasmic reticulum, and mitochondria (Figure 6.8). This, extreme case of mummifiation probably occurred because chemicals in the plant sap dilused into and embalmed the inseet’s tissues. 108 PARTTWO Continuity and Evlution of Anal Life Interpreting the Fossil Record ‘The fossil record is biased because preservation is selective, ‘Vertebrate skeletal parts and invertebrates with shell and other hard structures left the best records (Figure 6.8). Sofi-bodied animals, including jelyfishes and most worms, are fossilized only under very ‘unusual circumstances, such as those that formed the Burgess Shale ‘of British Columbia (Figure 6.9). Exceptionally favorable conditions {or fossilization produced the Precambrian fossil bed of South Australia, the tar pits of Rancho La Brea (Hancock Park, Los Angeles), the great dinosaur beds (Alberta, Canada, and Jensen, ‘Utah; Figure 6.10), and fossil beds of the Yunnan and Lianoning provinces of China. Fossils form in stratified layers with new deposits lying on top ‘of older ones. I left undisturbed, whichis rare, the ages of fossils in preserved sequence are directly proportional to depth in the strat fed layers. Characteristic fosils often serve to identify particular layers, Certain widespread marine invertebrate fossils, including various foraminferans (p. 238) and echinoderms (p. 474), are such ‘good indicators of specific geological periods that they are called index." oF “guide,” fossils. Unfortunately, the layers are usually tilted or show faults (racks). Old deposits exposed by erosion may bbe covered with new deposits in a different plane. When exposed to tremendous pressures or heat, staified sedimentary rock metamor- phoses into crystalline quarizite, slate, or marble, which destroys fossils. © Stratigraphy of fossils for two major groups of Affican ante- lopes is shown on Figure 6.11, Species in this group have different ‘characteristic sizes and shapes of horns, which form much of the fossil record ofthis group. Solid vertical lines in Figure 6.11 show the temporal distributions of species determined by presence oftheir ‘characteristic hors in rock strata of various ages. Red lines denote the fossil records of living species, and gray lines denote the fossil records of extnet species. The dotted gray lines show the inferred relationships among living and fossil species based on their sharing ‘of homologous structural features Geological Time Long before the earth's age was known, geologists divided its history into a table of succeeding events based on the ordered layers of sed= mentary rock. The “law of stratigraphy” produced a relative dating with the oldest layers at the bottom and the youngest at the top ofthe Sequence. Time was divided into eons, eras, periods, and epochs as shown inthe Timeline of Major Biological Events om the endshect inside the back cover of this book. Time during the last eon (Phanerozoic) is expressed in eras (for example, Cenozoic), periods (For example, Tertiary), epochs (for example, Paleocene), and some times smaller divisions of an epoch. In the late 1940s, radiometric dating methods were developed for determining the absolute age in years of rock formations. Several fei etGeieey Figure 6.8 Four examples of fossil material, Fish osi om rocks ofthe Green Rivet Formation, Wyoming. Such fsh swam here during the Eocene ‘epoch ofthe Tertiary period, aprenimataly 50 min years age. B, Stalked crinids (ass Crnaides, p. 489) rom Devonian rocks. The oss record af ‘hese echinoderms shows that they reached tex peak millons of years ear ad began a slow decine tothe present C, An insect fess that got sic in the resin a te approximately 25 mon years ago and thet has sine hardened into ember D Electron micrograph of Ussue rom e ly fesiized as shown in ithe nucloo of celle marked nro. wow. mbhe.com/hickmanipzt 7 (CHAPTER. Organic Evolution 105 Figure 6.9. Fosstuiobitevsble atthe Burgers Shale Quay. the Rocky Mountains of Brlsh Columbia, nado 8, Animal of te Cmts prio, epproxmately 505 milion years {ag0, 8 econsroted from fsa preserved in the Burgess Shale. The main now body plans hat appeered rather abruptly at thistime established the body plans of enimas fami to us ode. evn SchaloiAmy Sock Pho | ae yt Some 106, PART TWO Continuity and Evolution of Anima Life independent methods are now used, all based on the radioactive decay of naturally occurring elements into other elements “These “radioactive clocks” are independent of pressure and temperature changes and therefore are not affected by often violent cearth-building activities, (One method, potassium-angon dating, uses the decay of potas sium-40 (®K) to argon-40 (Ar) (12%) and caleium-40 (Ca) (88%). Argon is noble gas that evaporates from liquid media, It ‘accumulates inthe crystal structure of rock only after the rock has solidified andthe nuclear decay of potssium-40 produces a trapped ‘stom of argon. The half-life of potassium-40 is 1.3 billion years: half ‘ofthe orginal atoms will decay in 1.3 billion years, and balf of the remaining atoms wll be gone atthe end ofthe next 1.3 billion years. ‘This decay continues until all radioactive potassium-40 atoms are ‘gone. To measure the age of the rock, one caleulates the ratio of remaining potassium-40 atoms to the amount of potasium-40 orii- Figure 6.10 A hotoursaeton prtety excreted tom rocket nally there (the remaining potassium-40 atoms plus the argon-40 and Dinosour Province Park, Abert tion of many new structures, some of which will persist and give rise tonew homologies. Multiplication of Species ‘A branch point on the evolutionary tree means that an ancestral spe- ies has spit into two different ones. Darwin's theory postulates that genetic variation present within a species, especially variation that ‘occurs between geographically separated populations, provides the material from which new species arise. Because evolution is a branching process the total number of species produced by evolu tion increases through time, although most ofthese species event ally become extinet without leaving descendant species. A major challenge for evolutionists is to discover the process by which an ancestral species “branches” to form two or more descendant species ‘This theory adds a spatial dimension to evolutionary processes. ‘When populations ofa species become isolated from each other by ‘geographic barriers, the isolated populations undergo separate evo- Itionary change and diverge from each other. For example, when sea level was higher in the past than itis now, low areas of Cuba were ‘inundated, dividing its land area into multiple isolates. Lizard popu: lations that were formerly parts of a single species evolved species-level differences in isolation before another lowering of sea level reconsolidated Cuba as we know it today. Before we explore multiplication of species, we must decide ‘what we mean by “species.” As explained in Chapter 10, no consen- sus exists regarding definition of species. Most biologists agree, however, that important criteria for recognizing species include (1) descent of all members from a common ancestral population forming a lineage of ancestor-descendant populations, (2) reproduc tive compatibility (ability to interbreed) within and reproductive incompatibility between species for sexually reproducing animals, tnd (3) maintenance within species of genotypic and phenotypic cohesion (lack of abrupt differences among populations in allelic frequencies and organismal characteristics). The criterion of repro- ‘ductive compatibility has received the greatest attention in studies of species formation, also called speciation. Biological features that prevent different species from inter- breeding are called reproductive barriers. The primary prob- Tem of speciation is to discover how two initially compatible populations evolve reproductive barriers that cause them to become distinct, separately evolving lineages. How do popula- tions diverge from each other in their reproductive properties while maintaining complete reproductive compatibility within ‘each population’? Reproductive barriers btwoen populations usually evolve grad- ually. Evolution of reproductive barriers requires that diverging pop- ‘ulations must be kept physically separate for long periods of time. If diverging populations reunite before reproductive barriers have ‘evolved, interbreeding occurs between the populations and they merge. Speciation by gradual divergence in animals may require ‘extraordinarily long. periods of time, perhaps 10,000 to 100,000 us PARTTWO Continuity and Evolution of Animal Lite years or more. Geographical isolation followed by gradual diver- [gence is the most effective way for reproductive barriers to evolve, and many evolutionists consider geographical separation a prerequi- site for branching speciation. Geographical barriers between populations are nat the same thing 35 reproductive barriers. Geographical barriers refer to spatial separation of two populations. They prevent gone exchange and are usually a precondition for speciation Reproductive barriers result from evolution and refer te various behavioral, physical, physiological and ecological factors that prevent interbreeding between diferent species. Behavioral barriers often evalve faster then other kinds of reproductive barriers. Geographical barriers do not guarantee that reproductive barriers wll evolve, Reproductive barriers are most ely to evolve under conditions that inclide a generation of small population size, a favorable combination of selective factors, and long periods of geographical isolation. One or both of pair of geographically isolated populations may become extinct prior to evolution of reproductive harriers between them. Over the vast span of geolosical time, however, conditions| sulficient for speciation have occurred milions of times. Allopatrie Speciation Allopatic (“in another land”) populations of a species are those that occupy separate geographical areas. Because of their geo- ‘Eraphical separation, they cannot interbreed, but would be expected to do so ifthe geographic barriers between them were removed. If populations are allopatric immediately preceding and ‘during evolution of reproductive barriers between them, theres ing speciation is called allopattie speciation or geographic spe- ciation. The separated populations evolve independently and adapt to their respective environments, generating reproductive barriers hetween them as a result of their separate evolutionary paths. Because their genetic variation arises and changes independently, physically separated populations will diverge genetically even if their environments remain very similar: Environmental change between populations also can promote genetic divergence between them by favoring different phenotypes in the separated popula- tions. Ernst Mayr (Figure 6.19) has contributed greatly to our knowledge of allopatric speciation through his studies of speci tion in birds. Allopatri speciation begins when a species splits into two or more geographically separated populations, This spliting can hap- pen in either of two ways: by vieariant speciation or by a founder ‘event. Vicarant speciation is initiated when climatic or geological changes fragment a species’ habitat producing impenetrable barriers that Separate different populations geographically. For example, ‘mammalian species inhabiting a lowland forest could be divided by uplifting ofa mountain barrier, sinking and flooding ofa geological, fault, or climatic changes that eause prairie or desert conditions to ‘encroach on the forest. Formation ofthe isthmus of Panama sepa rated populations of the sea urchin genus Eucidaris into separate ‘Caribbean and Pacific Ocean isolates, leading to formation of the pair of species shown in Figure 1.50 (p. 6) ‘Vicariant speciation has two important consequences. Although the ancestral population is fragmented, individual fragments are Figure 6.19 erst Mayr 1904 t 2005}. major contrbuterto our knowledge of speciation and of evolution In general usually left fel intact. The vicariant process itself does not induce genetic change by reducing populations to a small size or by tans- porting them to unfamiliar enviconments, Another important conse (quence is that the same vicariant events may fragment several diferent species simultaneously. For example, fragmentation of a lowland forest most ikely would disrupt species from numerous and diverse taxonomic groups. including salamander, frogs, snails, and ‘many other forest dwellers. Indeed, the same geographic patterns are observed among closely elated species indifferent groups of organ- isms whose habitats are similar, Such patterns provide strong evi- dence for vicariant speciation ‘An alternative means of initiating allopatric speciation is for 1 small number of individuals to disperse to a distant place where 1no other members oftheir species occur. The dispersing individ uals may establish a new population in what is called a founder event. Allopatric speciation caused by founder events has been observed, for example, in the native fruit fies of Hawaii. Hawaii contains numerous patches of forest separated by voleanie lava flows. On rare occasions, strong winds ean transport a few Flies from one forest to another, geographically isolated forest where the flies are able to start a new population. Sometimes, a single fertilized female may found a new population. Unlike what hap- pens in vicariant speciation, the new population initially has a very small size, which ean cause its genetic structure to change quickly and dramatically from that of its ancestral population (see p. 123). When this event happens, phenotypic characteristics that were stable inthe ancestral population often reveal unprece- dented variation in the new population. Ax the newly expressed variation is sorted by natural selection, large changes in pheno- type and reproductive properties occur, hastening evolution of reproductive barriers between the ancestral and newly founded populations. “The term founder event its most general usage denotes a ispersal of organisms from an ancestral population across a ‘geographic barrier to start anew allopatric population. Origins ‘of Galtpagos finches by South American emigrants provide ‘good examples (p. 117). founder event doesnot aways directly cause important changes inthe genetic constitution of the new population relative to the old one although such changes are expected when the number of founding individuals isvery smal (les than Sto 10individuas for example) and the {ancestral population has alarge amount of genetic variation ‘Achange nthe genetic constitution ofa newly founded population because of small number of founders is termed a Jounder fect, which includes a population bottleneck (p. 125) Ifa founder effects so profound that selection ats new ways ‘on reproductvelyimportant characters, the founder event can induce speciation. Furderindvced speciation describes the subset ‘of founder eventsin which a founder effect hastens specieslevel clvergence ofthe newly founded population. Speciation in Hawaiian Orosphil fut flies as escried inthe text illustrates ‘ounder.indced speciation, Excluded from founder-nduced speciation are founder events whose ral in speciation i strictly to establish anew allopatric population capable of independent ‘evolutionary change ‘Surprisingly, we often learn most about the genetics of allopat- ric speciation from cases in which formerly separated populations regain geographic contact following evolution of incipient reproduc tive barriers that are not absolute. The occurrence of mating between pl — Disruptive selection Figure 6.33 Rosponsos selection ona continuous polygenie cheracte coloration in ane A, The equency dstbuton of eolraton Defoe selection. 8, Stabzing selection cls extreme vrents rom the Population, inthis ease eiminating mens that re unusaly ight (or dak, thereby sting the meon,C, Dectona selection shits the population mean, inthis case by favoring dey colored variants 1, Daruptive select overs both extremes but nat the meer the meen 's unchanged but the population nolonger hs ebellshaped dsbuton of phenctypes. 6.5 MACROEVOLUTION: MAJOR EVOLUTIONARY EVENTS Macroevolution describes large-scale events in organic evolu- tion. Speciation links macroevolution and mieroevolution. Major trends in the fossil record (see Figures 6.11, 6.12, and 6.13) are clearly within the realm of macroevolution. Patterns and pro- cesses of macroevolutionary change emerge from those of micro- evolution, but they acquire some degree of autonomy in doing so. ‘The emergence of new adaptations and species, and the varying rates of speciation and extinction observed in the fossil record, g0 beyond the fluctuations of allelic frequencies within populations. Stephen Jay Gould recognized three different “ters” of time at Which we observe distinct evolutionary processes. The fist ier con- stitutes the timescale of population genetic processes, from tens to thousands of years. The second tier covers millions of years, the scale on which rates of speciation and extinction are measured and ‘compared among different groups of organisms, Punctuated equilib: rium is a theory of the second tier, explaining the occurrence of speciation and morphological change and their association over mil- lions of years. The third tier covers tens to hundreds of millions of ‘yeas, and is marked by occurrence of episodic mass extinctions. In ‘the fossil record of marine organisms, mass extinctions recur at inter vals of approximately 26 million years. Five ofthese mass extnc tions have been particularly disastrous (Figure 6.34). The study of long-term changes in animal diversity focuses onthe third-tie times cale (see Figures 6.13 and 6.34). Speciation and Extinction Through Geological Time Broltioamy change althe second te proven new panpectve ca, Darwin's theory of natural selection. Although a species may persist, formany millions of years i ulimately has two posable elation ay fates: It may give ie to new species or become extnet without Ieavng descendants, Rates of speciation and extinction vary among lineages and incages that have the highest speciation rats and lo est extinction ates produce the greatest number of living species ‘The characteristics ofa species may make it more or les likely than citer to undenge speciation or extinction events, Beatme way characteristics re passed from ancestral to descendant species (anal ‘ogous to here tthe organismal level), ineages whose character istics increase the probably of speciation and confer resistance to extinction should dominate the living wor This species vel ror ess that prodoces differential rates of speciation and extinction mong lineages is analogous in many ways to natural selection. I ‘epresents an expansion of Darwin's theory of natural election This expansion i particularly important for macroevolution if one acepts thethoory of punctate equilib, which tats that the evolton arly important variation ocurs primarily among rather tan within species. Species selection encompasses the differential survival and imitation of species through gcologial ime based on variation among lineages, especialy in emergent, species-level properis “These specietevel properties inclode mating rituals soil struc ing. migration pattems, gographicdsribation, and all othr proper fe tht anergy a the species level (oe p= 5). Dement eprion sally resemble ther ancestors in these properties. Fr example, & “harem” prem of mating in whicha single male and several females compose 8 breeding unit characterizes some mammalian lineages tut no thers: We expect speciation mts to be enhanced by soa systems thal promote founding of new population by smal numbers of individuals. Cerin social systems may increase the likelihood thata species wil survive environmental challenges though cooper ative action. Such properties woud be favored by species selection over geological time. Differential speciation and extinction amon lineages also can tbe caused by variation in oganimal-evel properties (suchas spe- ciolied vers generalized feeding) rather tan spcies-level proper tis (ep. 5) Organisms that specialize in cating a restricted range of foods, for example, may be subjected more readily than generale ined feeders to geographic isolation among populations, because aces where ther preferred food is scarce o absent wil fanetion as scographic barriers to dispersal. Such geographic isolation could {generate more frequen opportunites for speciation to cur hrogh- ft pological time. The fossil records of two major groupe of www.mhhe.com/hickmanipet7e 3 8 [Number of famiies of marine animals oo Geologie time (108 yours b Figure 6.34 chengesin numbers oftaxonomicfamiles (9108) of marine animals trough ‘ime from the Cambrian peroat the present Sharp drops represent five mojo extinctions of the extinctions the vera numberof meine skeletonzed marine animes, Note thet des {anes hea incensed tthe present. African antelopes suggest tis result (see Figure 6.11). A group of specialized grazers that contains blesboks, hartebeest, and wilde bocsts shows high speciation and extinction rates. Since the late “Miocene, 33 extinct and 7 living species are known, representing at least 18 events of branching speciation and 12 terminal extinctions. In contrast, a group of generalist grazers and browsers that contains impatas shows little branching speciation or extinction during this ‘same interval of time. Interestingly, although these two groups differ {really in speciation rates, extinction rates, and species diversity, they do not difer significantly in total number of individual animals alive today. CHAPTER 6 Organic Evolution 129 Paleontologist Elisabeth Veba, whose research produced the results in Figure 6.11, uses the term ‘effect macroevolution to describe differential spe-

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