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Williams, George C. “Pleiotropy, natural selection and the evolution of senescence.” Evolution 11 (1957): 398–411. ———. Sex and Evolution. Princeton, N.J.: Princeton University Press, 1975. Linnaean system The Linnaean system is used by biologists to classify species of organisms into nested categories. The system was invented by the Swedish botanist Karl Linné (see Linnaeus, Carolus) in the 18th century. Prior to Linnaeus, the classification of organisms was in disarray. First, each species was usually identified by long Latin descriptions. For example, the common ground cherry was called Physalis amno ramosissime ramis angulosis glabris foliis dentoserratis, which means, in part, that it had angled stems and the leaves were hairless and had toothed margins. Linnaeus shortened these long names to two-word names (in this case, Physalis angulata, a name it still uses). Thus Linnaeus established the practice of binomial nomenclature (“naming system that uses two names”). Second, the same species could be given different descriptive names by different botanists. Linnaeus standardized the names. Was Rosa sylvestris alba cum rubore, folio glabro the same as Rosa sylvestris inodora seu canina? Linnaeus standardized it to Rosa canina. Mistakes still happen, in which the same species is accidentally given two different names, but it is now much less common. These innovations were accepted by the biological community with a sense of relief, but this was not the most important innovation of the Linnaean system. Before Linnaeus, there was no recognizable basis for classification. Some zoologists classified animals on the basis of terrestrial vs. aquatic, others on the basis of big vs. small. French paleontologist Georges Buffon (see Buffon, Georges) organized animals according to their usefulness in the human economy. Linnaeus brought order into this chaos. In classifying plants, Linnaeus recognized that reproductive characteristics were less variable than the characteristics of leaves and stems. He therefore classified plants on the basis of the numbers and condition of the stamens (male parts) and pistils (female parts) of their flowers. Consistent with the male bias of science and society, Linnaeus’s major axis of classification was the number of stamens: 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 12, and many individual stamens; the grouping of the stamens (tetradidymous, dididymous, monadelphous, diadelphous, polyadelphous); the fusion of stamens to one another or to pistils; plants with separate male and female flowers on the same tree (monoecious) or different trees (dioecious) or a mixture of arrangements. His last class, the cryptogams (crypto- means hidden), consisted of plants that had no obvious flowers (see seedless plants, evolution of). Within the male axis, he divided plants on the basis of their female parts. He applied his system in a rigid fashion, resulting in classifications that are surprising to modern botanists. For example, he placed oaks, beeches, hazelnuts, and cattails into the same category; modern botanists recognize cattails as very different from the other three (see angiosperms, evolution of). The central feature of the Linnaean system is that it is a nested hierarchy. The specific epithet (the second of the Linnaean system two words in the name) referred to the species, and the genus name (the first word) was given to a set of closely related species. As the system was modified by later scientists, closely related genera were clustered into families; closely related families into orders; closely related orders into classes; closely related classes into phyla; phyla into the major kingdoms, of which there were originally only two: plants and animals. Linnaeus inspired the scientific world because he brought, or revealed, order within the diversity of organisms on the Earth which was just then being thoroughly explored. The Linnaean system seemed to correspond closely to the concept of the Great Chain of Being (see scala naturae). All things, from rocks to plants to animals to humans to angels to God, were believed to be connected like a tapestry of fibers. Scientists were not surprised to find intermediate entities—for example, a sponge as intermediate between animal and rock, or beings halfway between humans and apes—because it was all part of the created tapestry. Linnaeus had no thought of evolutionary origins of organisms; this is revealed by the fact that he applied his system to rocks as well as to organisms. Linnaeus tried to make the system as natural as possible by using characteristics that were a realistic reflection of the Chain of Being. While Linnaeus gave no thought to evolution, the fact that the natural world could be so readily classified into nested categories struck Charles Darwin as being a major piece of evidence favoring evolution (see origin of species [book]). Despite naturalists’ belief in the Scala Naturae, there actually were not many intermediates between modern entities. Sponges are not really half-rock, half-animal. Dogs, wolves, coyotes, and foxes form a group of similar canines; cats, lions, tigers, leopards, and cheetahs form a group of similar felines; and there are no cat-dogs. Organisms, as Linnaeus classified them, display a pattern like branches of an evolutionary tree. Today, biologists attempt to classify organisms using the Linnaean system but try to make the system as natural as possible by defining genera, families, orders, classes, and phyla on the basis of evolutionary divergence (see cladistics). The species themselves are recognized when possible on the basis of reproductive isolation (see speciation). When a group of species share many features with one another, scientists conclude that they evolved from a common ancestor more recently than species that share few features. The Latin (or Greek) generic and specific names may describe the characteristics that make the genus or species different from other species or genera; they may describe visible features of the organism, or the location in which it lives; they may be the ancient Greek or Latin names for the organisms; or they may be a tribute to a scientist who studied the organisms. For example, dogs are Canis familiaris—genus Canis, species familiaris. Canis is a Latin word for dog. Other members of the genus Canis include Canis lupus, the wolf (lupus is Latin for wolf), and Canis latrans, the coyote (latrans comes from the Latin for barking). This system of groupings also explains the higher levels of the Linnaean system (see figure on page 248): • Genera that evolved from a common ancestor most recently, and are therefore most similar to one another, are
Linnaean system The tree of life used in this figure brings the Linnaean system together with modern evolutionary science, with a focus on animals. The last universal common ancestor (LUCA) evolved into bacteria, archaebacteria, and eukaryotes. Most biologists classify the bacteria and archaebacteria into Kingdom Monera. The eukaryotes diversified into many lineages, most of which are classified into Kingdom Protista. One eukaryotic lineage evolved into the modern plant kingdom, and another into the modern animal and fungus kingdoms. Dashed lines represent symbiogenesis. grouped together into the same family. The canid family, Canidae, contains all the doglike animals, such as foxes, along with the true dogs. • Families that evolved most recently from a common ancestor, and are therefore most similar to one another, are members of the same order. Canids, along with cats, bears, raccoons, weasels, badgers, skunks, and otters, are members of the Order Carnivora, the carnivores. The families in the carnivore order share common features such as the kinds of teeth they possess.
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ENCYCLOPEDIA OF Evolution
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Encyclopedia of Evolution Copyright
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Contents Foreword ix Acknowledgment
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Foreword The theory and facts of ev
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IntroduCtIon What you do not know a
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other issues coming along with it.
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adaptation Adaptation is the fit of
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from the allometric patterns of gro
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considered as an example. (The taxo
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English) were a resounding success,
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Some Centers of the Evolution of Ag
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helps the DNA insert into the host
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Even within a single patient, HIV e
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then into the insect body; carbon d
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chromosomes (they are diploid) whil
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genetic basis. About 35,000 years a
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emained dormant and sprouted after
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Further Reading Wise, Steven M. Rat
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Because the DNA of many archaebacte
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Koi and goldfish have been bred tha
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Barringer Crater, Arizona, was form
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y a silent wall of darkness advanci
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Skeleton of “Lucy,” the Austral
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Further Reading Alemseged, Zerxenay
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acteria, evolution of Examples of t
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0 Bates, Henry Walter advantage. Th
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ehavior, evolution of In another sp
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ig bang make nests with horizontal
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iodiversity How Much Do Genes Contr
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iodiversity Biodiversity (as indica
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0 biogeography Another problem with
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iogeography Biogeography of Selecte
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iogeography their species through d
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iology design) or adaptation to the
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iology D. Response to environment.
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0 birds, evolution of • The foot.
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irds, evolution of • doves and pi
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Burgess shale early career. By heat
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Burgess shale jointed legs, Anomalo
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Cambrian period occurred when anima
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0 Cenozoic era opens the way to an
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Chicxulub Pritchard, J., and Dolph
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cladistics The above examples used
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coevolution descent, and that scien
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coevolution Coevolution of Organism
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0 coevolution What Are the “Ghost
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coevolution What Are the “Ghosts
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commensalism Wilson, Michael. Micro
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continental drift was an animal tha
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continental drift During geological
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0 convergence bee-pollinated flower
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convergence Both birds and bats hav
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creationism explained these observa
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creationism used to justify militar
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creationism • In the early 21st c
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00 Cretaceous period however, other
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0 Cro-Magnon evolution had occurred
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0 Cro-Magnon areas, perforated shel
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0 Cuvier, Georges Cuvier held stron
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0 Darwin Awards Darwin Awards “Th
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0 Darwin, Charles forever. It would
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Darwin, Charles and animals did hav
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Darwin’s finches has its own uniq
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Dawkins, Richard ate the pulp. More
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developmental evolution Part II. In
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0 Devonian period gene from an inve
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dinosaurs what DeVries thought were
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diseases, evolution of • Thyreoph
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disjunct species have happened? Som
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DNA (evidence for evolution) genes,
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0 DNA (evidence for evolution) seco
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DNA (raw material of evolution) DNA
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DNA (raw material of evolution) The
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Dobzhansky, Theodosius Mice have tw
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duplication, gene prominent brow ri
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0 ecology Second, organisms can con
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Eldredge, Niles they possess no str
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eugenics grieving, or experiencing
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eugenics because government officia
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eukaryotes, evolution of of the sor
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0 Eve, mitochondrial within cells,
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evolutionary ethics the ability to
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evolutionary ethics Can an Evolutio
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evolutionary medicine Can an Evolut
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evolutionary medicine variable in t
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0 extinction past (a genetic bottle
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fishes, evolution of his obvious vi
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Flores Island people Christopher St
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fossils and fossilization The grain
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founder effect Further Reading Fort
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0 Gaia hypothesis nitrogen oxide (N
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Galton, Francis constituted a premi
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gene pool Cocos Island is too small
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Gould, Stephen Jay animals, and the
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Gray, Asa he had long accepted Lyel
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0 group selection used the carbon t
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gymnosperms, evolution of began per
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Haeckel, Ernst (1834-1919) German E
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The initial divergence between homi
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Although Homo ergaster, the presume
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tions were spreading through Africa
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Selected Examples of Homo heidelber
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sunlight of tropical regions, and t
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Page 218 and 219: gene transfer from a bacterium, per
Page 220 and 221: to evolutionary science. He also ap
Page 222 and 223: Examples of Intergeneric Crosses Re
Page 224 and 225: ice ages The term ice ages usually
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Page 230 and 231: ence their intelligence. There is a
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Page 234 and 235: e exactly the right ones. Bovine in
Page 236 and 237: Van Till, Howard J. “E. coli at t
Page 238 and 239: food particles with whiplike struct
Page 240 and 241: would readily interbreed and produc
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Page 244 and 245: isotopes When evaporation from the
Page 246: Java man See Homo erectus. Johanson
Page 249 and 250: 0 Kenyanthropus chemist Henri Becqu
Page 251 and 252: language, evolution of is not consi
Page 253 and 254: language, evolution of Italian, Por
Page 255 and 256: Lascaux caves Some of the original
Page 257 and 258: Leakey, Richard Richard Leakey, in
Page 259 and 260: 0 life history, evolution of it inv
Page 261 and 262: life history, evolution of is: sele
Page 263 and 264: life history, evolution of Why Do H
Page 265: life history, evolution of Why Do H
Page 269 and 270: 0 living fossils admired. The genus
Page 271 and 272: Lysenkoism about evolution. When Da
Page 273 and 274: Malthus, Thomas intelligent design)
Page 275 and 276: mammals, evolution of the reptilian
Page 277 and 278: markers another precocious and crea
Page 279 and 280: 0 Mars, life on • The spacecraft
Page 281 and 282: Maynard Smith, John producing a 200
Page 283 and 284: meiosis Zoology. He became director
Page 285 and 286: Mendel, Gregor of the fertilized eg
Page 287 and 288: Mendelian genetics an American gene
Page 289 and 290: 0 Mendelian genetics the environmen
Page 291 and 292: microevolution Stanley Miller did o
Page 293 and 294: missing links Batesian mimicry. Nam
Page 295 and 296: mitochondrial DNA Hominin skulls ha
Page 297 and 298: molecular clock these scientists di
Page 299 and 300: 0 mutualism function of the protein
Page 301 and 302: natural selection Fact 1. Populatio
Page 303 and 304: natural selection Three types of na
Page 305 and 306: natural theology different kinds of
Page 307 and 308: Neandertals different species of hu
Page 309 and 310: 0 Neolithic Neandertals lived short
Page 311 and 312: new synthesis In contrast to progen
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origin of life Are Humans Alone in
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00 origin of life It is often said
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0 Origin of Species (book) ribose);
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0 Orrorin ———. On the Origin
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0 Paley, William in the Chordata) e
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0 peppered moths There remained som
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0 Permian extinction produced in la
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Permian period upon the exploitatio
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Piltdown man found in slightly diff
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plate tectonics The Earth’s surfa
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population Percent of Mammal Genera
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0 population genetics Vandermeer, J
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population genetics If allele A is
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Precambrian time at least some gene
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progress, concept of arboreal prima
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promoter Each chromosome contains t
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0 punctuated equilibria and persist
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punctuated equilibria Gradualism wa
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Quaternary period epoch of the Quat
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ecapitulation would cause some of t
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eligion, evolution of pathogens nev
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0 reproductive systems Catkins of m
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eproductive systems Scobell has inv
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eproductive systems fact, there is
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eptiles, evolution of Blanckenhorn,
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esistance, evolution of ineffective
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0 respiration, evolution of There i
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Sagan, Carl (1934-1996) American As
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tain features are universally recog
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The testing of hypotheses accumulat
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the results are sufficiently intere
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een wrong: His hypothesis of the co
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Comparison of Inherit the Wind with
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and others form more complex struct
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endonuclease gene is then used as a
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e able to do the same thing. Second
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mosomes come in groups of five rath
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eliminated—that is, if sexual rec
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one, free of parasites, is likely t
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nesses of resource acquisition and
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Zahavi, Amotz. The Handicap Princip
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and it is not happening extensively
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this, too, is a genetically based u
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monkeyflowers of the genus Mimulus)
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Further Reading Abrahamson, Warren
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comes from the unfertilized egg, ra
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of the bacteriophages. Therefore, h
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technology Technological and Cultur
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thermodynamics • Plants. The flow
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thermodynamics return to their orig
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00 Triassic period more resulting l
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unconformity An unconformity is a g
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sissippi River is getting shorter e
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ago. This was the birth of the Sun.
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0 vestigial characteristics algae.
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Wallace, Alfred Russel (1823-1913)
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genetic inferiority of the lower cl
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that point onward he questioned the
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Carl R. Woese pioneered the use of
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Darwin’s “One LOng argument”:
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the young seedlings must compete wi
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Traits can reappear after even hund
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ated: It has been an advantage in o
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chapter 10. On the imperfection of
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out in competition with the species
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languages. Linguists classify all R
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My theory has been much maligned. T
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Index Note: Boldface page numbers i
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asexual propagation 370-371 Ashfall
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Bryan, William Jennings creationism
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ird evolution 62-63 cladistics 72-7
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divergence molecular clock 278-279
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extinction 159-160. See also mass e
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Gondwana (Gondwanaland) 68, 84, 86,
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Huxley, Julian S. 200 eugenics 146
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Mendelian genetics and 268 Spencer,
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Origin of Species (Darwin) 433-434
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ocean currents 179, 207 Oceanian re
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polar cells 368 Polkinghorne, John
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Sanger, Frederick 55 Sanger, Margar
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spores 264, 370 spotted hyena (Croc
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useless characteristics 409 Ussher,
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