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618 PART 5 / Macroevolution Figure 22.3 Phylogenies of North American Tetraopes (beetles) and their food plants, milkweeds (Asclepias). The phylogenies are mainly mirror images, or cophylogenies. The two or three exceptions may be due to errors in phylogenetic inference or to host shifts (as discussed in Section 22.3.3). These beetles exploit their food plants both as larvae (which bore into the roots) and as adults (which eat the flowers and leaves). Redrawn, by permission of the publisher, from Farrell & Mitter (1994). Cophylogenies can arise because of... . . . coevolution ... Tetropes (beetles) Phaea grp 1 Phaea grp 2 T. comes T. ineditus T. elegans T. discoideus (MX) T. discoideus (US) T. umbonatus T. melanurus T. S-maculatus T. annulatus T. pilosus T. mandibularis T. tetropthalmus T. varicornis T. femoratus T. sublaevis T. basalis Convolvulaceae Apocynaceae Matelea Marsdenia A. subulata A. curassavica A. subverticillata A. glaucescens A. tuberosa A. amplexicaulis A. sullivanti A. arenaria A. latifolia A. syriaca A. notha A. speciosa A. erosa A. eriocarpa Asclepias (milkweeds) Simple cardenolides More complex, toxic cardenolides Most toxic cardenolides concentrated in latex 22.3.2 Two taxa may show mirror-image phylogenies, but coevolution is only one of several explanations for this pattern Figure 22.3 shows on the left the phylogeny of 15 out of the 25 beetles in the genus Tetraopes that live in North America, together with two relatives. On the right is the phylogeny of their food plants, the milkweeds (Asclepias). Milkweeds contain poisons (cardenolides), but Tetraopes are not harmed by them. The beetles store the toxins in their own bodies, making them unattractive to birds and mammals. Tetraopes are brightly colored in orange and black. The striking feature of the two phylogenies in Figure 22.3 is that they are near mirror images. With two or three exceptions, the phylogeny of the beetles matches the phylogeny of the plants they eat. In technical language, the phylogenies of the two taxa are cophylogenies. The phylogenies of two taxa are cophylogenies if they have the same (or much the same) branching pattern. Statistical tests exist to determine whether two phylogenies are more similar than would be expected by chance. Cophylogenies can arise for at least three reasons. One is coevolution in the full sense of the word. The two taxa have exerted evolutionary influences on each other, and evolution leading to speciation in one taxa tends to cause speciation in the other taxon too. For instance, two subpopulations of one ancestral milkweed species might have ..
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.. EVOLUTION
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© 2004 by Blackwell Science Ltd a
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vi Brief Contents PART 5. MACROEVOL
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viii Full Contents 3.5 Ring “spec
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x Full Contents 7.2 Rates of molecu
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xii Full Contents 10.5 Genetics of
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xiv Full Contents PART 4. EVOLUTION
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xvi Full Contents 14.12 Identificat
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xviii Full Contents 17.7 Geographic
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xx Full Contents 21.4.3 Ordovician
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Preface The theory of evolution is
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xxiv Preface evolve resistance to d
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.. Introduction Part one When Darwi
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4 PART 1 / Introduction Evolution i
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6 PART 1 / Introduction Examples ex
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8 PART 1 / Introduction Time (a) (b
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10 PART 1 / Introduction . . . look
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12 PART 1 / Introduction Time (a) (
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14 PART 1 / Introduction Figure 1.7
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18 PART 1 / Introduction . . . and
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20 PART 1 / Introduction Further re
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22 PART 1 / Introduction DNA is car
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24 PART 1 / Introduction Most genes
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26 PART 1 / Introduction Table 2.1
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28 PART 1 / Introduction Original D
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30 PART 1 / Introduction (a) Transp
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32 PART 1 / Introduction . . . per
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34 PART 1 / Introduction Dominance
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Figure 2.11 Two populations with 10
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42 PART 1 / Introduction Study and
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44 PART 1 / Introduction Life could
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48 PART 1 / Introduction Tooth qual
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50 PART 1 / Introduction Artificial
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52 PART 1 / Introduction Ring speci
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54 PART 1 / Introduction Human obse
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58 PART 1 / Introduction 5' pG Anti
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62 PART 1 / Introduction (a) α-hem
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64 PART 1 / Introduction Age (Myr a
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66 PART 1 / Introduction The groups
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68 PART 1 / Introduction The scient
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70 PART 1 / Introduction Study and
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Plate 3 Here in the lower row are s
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Plate 6 Scrub jays (Aphelocoma coer
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Plate 8 Chromosomal races of the ho
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Plate 10 Geological map of North Am
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72 PART 1 / Introduction Cod produc
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74 PART 1 / Introduction The strugg
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76 PART 1 / Introduction Many chara
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82 PART 1 / Introduction Chromosome
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86 PART 1 / Introduction The condit
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90 PART 1 / Introduction Summary 1
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.. Evolutionary Genetics Part two T
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.. 5 The Theory of Natural Selectio
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.. Figure 5.1 The general model of
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.. . . . and use Mendel’s rules t
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.. A simpler proof of the Hardy- We
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.. The MN human blood group system
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.. CHAPTER 5 / The Theory of Natura
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.. We need to know more to understa
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.. Figure 5.4 Peppered moths natura
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.. Mark-recapture experiments sugge
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.. Manchester The fitness estimates
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.. Pests, such as mosquitoes, evolv
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.. Figure 5.8 The mortality of mosq
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.. Box 5.2 Resistance Management Th
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.. Other factors will be at work in
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.. . . . which can sometimes be use
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.. Figure 5.9 The global incidence
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.. In host-parasite relations, the
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.. Populations may be subdivided CH
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.. We construct a model of gene fre
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.. We construct a model of selectio
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.. Further reading CHAPTER 5 / The
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.. 6 Random Events in Population Ge
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.. Box 6.1 Random Sampling in Genet
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.. Probability of polymorphism 1.0
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.. Evolution can occur by random dr
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.. Random drift has consequences fo
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.. Generation 1 Generation2 Adults
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.. CHAPTER 6 / Random Events in Pop
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.. Heterozygosity ( H ) 1.0 0.9 0.8
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.. Summary 1 In a small population,
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.. 7 Natural Selection and Random D
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.. Two extreme views b selectionist
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.. The original neutral theory has
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.. . . . and levels of polymorphism
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.. homozygotes by normal Mendelian
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.. Amino acid differences 1.0 0.9 0
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.. Globin evolution in “living fo
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.. Long-generation species, e.g., w
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.. Observed heterozygosity 1.0 0.8
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.. The nearly neutral theory can ex
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.. Insulin illustrates the effect o
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.. CHAPTER 7 / Natural Selection an
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.. Biologists came to accept, throu
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.. Elevated dN/dS ratios are observ
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.. Box 7.4 Model Organisms for Biom
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.. But the test has proved illumina
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.. Evidence shows that synonymous c
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.. The genomic era is allowing new
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.. Summary 1 The neutral theory of
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.. Study and review questions 1 Dra
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.. The tiger swallowtail butterfly
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.. Mimicry requires the correct com
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.. A haplotype is a haploid combina
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.. Linkage disequilibrium ( D) r =
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.. The more complex multilocus mode
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.. Expected frequency ( piqi ) (a)
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.. A model of haplotype frequency c
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.. Evidence for linkage disequilibr
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.. (proportion of polymorphic sites
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.. CHAPTER 8 / Two-locus and Multil
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.. Adaptive topographies can be use
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.. Mean population fitness ( w ) -
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.. p' 0 1 0 Frequency of A allele S
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.. Study and review questions 1 Her
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.. Beak size influences feeding eff
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.. Population size Seed √depth ×
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.. Continuous characters are influe
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.. Genetic and environmental influe
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.. Several non-additive genetic fac
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.. Box 9.1 Some Statistical Terms U
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.. Parent-offspring similarity depe
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.. Frequency Frequency Generation 1
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.. New variation can be introduced
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.. (a) 1976-1977 (b) 1981-1982 (c)
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.. Figure 9.11 (a) The main veins i
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.. Certain non-linear relations can
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.. Behavior Physiology Life history
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.. Some heritable characters do not
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.. Further reading CHAPTER 9 / Quan
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.. Adaptation and Natural Selection
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.. 10Adaptive Explanation T his cha
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.. . . . factually, ... . . . or th
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.. Not all evolution is adaptive CH
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.. Figure 10.2 Stages in the evolut
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.. Some critics suggest that the in
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.. Box 10.1 Molecular Cooption The
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.. Fitness (a) (b) Small mutation L
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.. Figure 10.5 A cross between the
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.. . . . by experiment, ... . . . a
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.. Figure 10.6 The fruits of (a) Cr
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.. Heterozygous advantage may lead
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.. Figure 10.7 Developmental asymme
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.. . . . that can be tested between
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.. Fitness (a) Time 1 (b) Time 2 (c
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.. Fitness Fitness (a) Time 1 (c) T
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.. Imperfect adaptations may or may
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.. The “design” of an eye for s
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.. hypothesis of adaptation or cons
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.. Study and review questions 1 Wha
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.. An adaptation may benefit one le
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.. The sd gene gains an advantage f
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.. CHAPTER 11 / The Units of Select
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.. We define Hamilton’s rule Hami
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.. Kin selection is at work in this
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.. Selfish The theories of group se
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.. Box 11.1 Group Selection for Egg
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.. The unit of selection, in a seco
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.. * * The word “gene” is being
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.. Summary 1 Adaptations evolve by
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.. in Sexual Reproduction 12Adaptat
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.. Figure 12.2 Non-reproductive sex
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.. Figure 12.3 Evolution in (a) ase
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.. Asexual reproduction has a spind
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.. . . . as illustrated by a motor
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.. Log fitness of organisms Number
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.. Figure 12.7 Frequency changes of
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.. Genetic cycles in snails are con
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.. Darwin provided comparative evid
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.. . . . which may explain extreme
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.. Theory requires open-ended choic
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.. But selection may have removed h
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.. Courtship bouts Net reproductive
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.. Theory predicts deviations from
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.. CHAPTER 12 / Adaptations in Sexu
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.. Study and review questions 1 Wha
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346 PART 4 / Evolution and Diversit
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17Evolutionary Biogeography B iogeo
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522 PART 5 / Macroevolution looked
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524 PART 5 / Macroevolution Fossili
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526 PART 5 / Macroevolution Era Per
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528 PART 5 / Macroevolution N = N 0
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530 PART 5 / Macroevolution Early l
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532 PART 5 / Macroevolution Cells h
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.. Multicellular life originated ov
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.. Present ~540 Myr ~1,200 Myr Echi
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.. Figure 18.8 The rise of the angi
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.. Figure 18.9 The origin of tetrap
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.. Probainognathus Thrinaxodon Proc
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.. . . . and cynodonts but their te
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.. . . . and sexual dimorphism all
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.. Homo sapiens c. 100,000 years ag
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.. Macroevolution may be due to ext
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.. Summary 1 Fossils are formed whe
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.. Study and review questions 1 Rev
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.. Evolutionary genomics aims to an
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.. Figure 19.1 The history of the h
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.. (a) A (b) 2 1 B C Vertebrate 1 2
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.. Following symbiosis ... . . . ge
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.. X and Y chromosomes evolve apart
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.. Figure 19.5 (a) A chromosomal in
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.. Mice do not show the same patter
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.. Study and review questions 1 (a)
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.. Morphological evolution is drive
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.. Figure 20.2 (a) Evolution by ter
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.. Developmental change can be by h
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.. . . . than might be expected fro
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.. A similar gene works in eye deve
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.. Figure 20.6 History of the Hox g
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.. Changes in arthropod limbs are a
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.. Switching systems may have made
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.. Study and review questions 1 If
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.. Horse teeth are an example of ho
Page 1254: .. Horse teeth show a representativ
Page 1258: .. Figure 21.2 Possible phylogeny o
Page 1262: .. Figure 21.3 The relation between
Page 1266: .. . . . disappear after the measur
Page 1270: .. Box 21.1 Two Meanings of Gradual
Page 1274: .. Time Punctuated equilibrium extr
Page 1278: .. Time General younging direction
Page 1282: .. Score Change in score per millio
Page 1286: .. Time (Myr) Two species/million y
Page 1290: .. Summary CHAPTER 21 / Rates of Ev
Page 1294: .. 22Coevolution C oevolution happe
Page 1298: .. Figure 22.2 Coevolution means th
Page 1302: .. Pollinator relationships have le
Page 1308: 620 PART 5 / Macroevolution . . . o
Page 1312: 622 PART 5 / Macroevolution One stu
Page 1316: 624 PART 5 / Macroevolution Box 22.
Page 1320: 626 PART 5 / Macroevolution Table 2
Page 1324: 628 PART 5 / Macroevolution Nematod
Page 1328: 630 PART 5 / Macroevolution The cop
Page 1332: 632 PART 5 / Macroevolution Figure
Page 1336: 634 PART 5 / Macroevolution Carnivo
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Page 1344: 638 PART 5 / Macroevolution Number
Page 1348: 640 PART 5 / Macroevolution Coevolu
Page 1352: 642 PART 5 / Macroevolution Study a
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644 PART 5 / Macroevolution Adaptiv
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646 PART 5 / Macroevolution Extinct
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648 PART 5 / Macroevolution . . . b
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Tertiary Lower Danian Cretaceous Up
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652 PART 5 / Macroevolution The ast
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654 PART 5 / Macroevolution 10 6 ye
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656 PART 5 / Macroevolution Extinct
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658 PART 5 / Macroevolution Darwin
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660 PART 5 / Macroevolution (a) Ext
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662 PART 5 / Macroevolution Figure
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664 PART 5 / Macroevolution Hansen
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666 PART 5 / Macroevolution Taxa wi
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668 PART 5 / Macroevolution Fossil
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670 PART 5 / Macroevolution One gro
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672 PART 5 / Macroevolution Molecul
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674 PART 5 / Macroevolution Changes
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676 PART 5 / Macroevolution . . . o
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678 PART 5 / Macroevolution Summary
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680 PART 5 / Macroevolution (2002),
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Glossary Words in italics cross-ref
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684 Glossary individuals, such as p
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686 Glossary isolating mechanism An
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688 Glossary random drift Synonym o
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Answers to Study and Review Questio
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692 Answers to Study and Review Que
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700 References Arnold, M.L. (1997).
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702 References Briggs, D.E.G. & Cro
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704 References Clarke, C.A., Sheppa
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706 References Dietl, G.P., Alexand
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708 References Flynn, J.J. & Wyss,
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710 References Grant, P.R. & Grant,
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712 References Hey, J. (2001). Gene
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714 References Jeffreys, A.J., Royl
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716 References Kruuk, L.E.B., Meril
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718 References MacFadden, B.J. (199
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720 References Milinkovitch, M.C.,
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722 References Orr, H.A. (1998). Th
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724 References Rice, W.R. (2002). E
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726 References Sequeira, A.S., Lant
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728 References Surlyk, F. & Johanse
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730 References Vrba, E.S. (1993). T
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732 References Wright, S. (1977). E
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734 Index angiosperms (cont.): inse
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736 Index coloration (cont.): prezy
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738 Index evolution (cont.): long-t
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740 Index genetics (cont.): charact
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742 Index immunoglobulins 24 immuno
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744 Index molecular distance 440-2
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746 Index parasites (cont.): genome
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748 Index reproductive development
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750 Index synonymous changes (cont.
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