Index

ABCDE model of flower development, 86 age structure of populations, 64–5 abd-B , 264 Agrostis tenuis (grass), 58 abdomen, 125–8 alleles, 71 Acanthostega, 171–2 allometric growth, 88, 113, 158, 336 acoel flatworms, 170, 240 allopatric speciation, 274–5, 278 Acrothoracica, 125, 126 allopolyploidy, 275 actinomorphic flowers, 244–5 alpha chains (haemoglobin), 136 Actinopterygii (ray-finned fishes), 171, 300 altered developmental gene products, 137–9 Acyrthosiphon pisum (pea aphid), 256 altered morphology, 140–44 adaptation altered pigmentation classic concept, 320–21 Biston betularia (peppered moth), 57, coadaptation distinction, 203 68–71, 73, 186, 188, 191, 197 connection to and gene Drosophila melanogaster, 264–5 co-option, 311–26 heterotypy, 139–40 direction of , 185–99 Manduca sexta , 268 adaptive hypothesis, 127 altered sequence mutations, 77–80 adaptive landscape model, 190–91, 209 Amborellaceae (angiosperms), 176, 178, 336 adaptive scenarios Ambystoma mexicanum (axolotl), 65 appropriateness, 98–9 amino acid substitutions, 136, 139 definition, 336 Amniota, 154, 173, 336 developmental plasticity, 253 amphibians, 9, 154, 173, 216, 249, 336 eye positions in flatfish, 108 Anura, 304 fins-to-limbs, 173 COPYRIGHTEDfrogs, 35, MATERIAL36, 304–5 rodent teeth formation, 117 Xenopus laevis, 37 sinistral/dextral snail shells, 111 toads, 304 snake fang position, 112 axolotl (Ambystoma mexicanum), 65 adhesion, cells, 246–7 paedomorphic salamanders, 94, 95 adult complexity increases, 299–302 amphioxus (cephalochordates), 163 adultocentric view (Minelli), 8 amphipods, 89 African brown butterfly (Bicyclus anynana), anagenesis, 274 128–32, 212, 243, 254 ancestrula, zooid modules, 9

Evolution: A Developmental Approach, by Wallace Arthur © 2011 Wallace Arthur

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Anchiornis, 197 terminology, 241, 242 angiosperms trunk outgrowths, 236 definition, 336 Arabidopsis thaliana Hydatella, heterochrony, 103–5 altered products of developmental , leaf arrangements, 214–15 137–9 origins, 176–9 close relative of, 85 plants, lineage, 302 gene naming conventions, 359–61 Solanaceae, heterotopy, 116–17 tcp1 gene co-option, 244 taxon sampling, 328–9 arachnids, 168 animals; see also individual species/types Archaeopteryx, 197 body plans, direction of evolution, arctic fox, 192 284–90 arthropods complexity, lineage, 301–2 heterochrony, 95–9 embryonic cleavage patterns, 18–19 inversion of bilaterian dorso-ventral axis, genetic toolkit concept, 245–9 114–16 sequence discovery, 25 kingdom composition/complexity, gene co-option, 235 328–9 origins, 165, 167, 169 model organisms, 35–6 repatterning, 82–5, 87 origins of multicellularity, 11–14 artificial selection segmentation, mapping to trees, 163–9 butterfly eyespots, 130–31, 211–13 taxon sampling, 328–30 Drosophila melanogaster, genetic terminology/taxonomy, 297 assimilation, 264–7 Annelida (segmented worms), 163–7 polyphenism from stable , Antennapedia complex, 223, 226 267, 269 antennapedia gene, 26, 83–4 asexual reproduction, 58–60 antero-posterior axis aspm (abnormal spindle-like microcephaly- bilaterian animals, 248 associated) gene, 122 Drosophila, 49–50 assimilation, genetic, 264–7, 342 hedgehog gene, 48 Astyanax mexicanus (blind cave-fish), heterotopy, 112–16 132, 133 Hox genes, 27 atomised cluster, 228, 229 polarity determination, 35 Australopithecus spp., 123 segmentation origins, 163–9 Aves see birds (Aves) Anura, 304, 305, 337 axolotl (Ambystoma mexicanum), 65 aphicarus gene, 256 aphids, wing polyphenism, 254–6 balancing selection, 72, 186 Aphis fabae (bean aphid), 256 barnacles (Cirripedia), 125–8 apomorph, 337 basal (primitive) angiosperms, 103–5 apoptosis, 38–40, 337 Bateson, William, 225 appendages beak shape/size, Darwin’s finches (Geospiza), centipedes 6, 7, 186–90 leg-pairs, 83, 213–14, 257 bean aphid (Aphis fabae), 256 poison claw segment, 141–3 bears, 191–2 definition, 241, 242 beauty of universal laws, 335 gene co-option, 237–43 beetles, horns, 133 limb origins, 169–175 beta chains (haemoglobin), 135, 136

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bi-directional heterometry, 128–32 Fisherian vs. Goldschmidtian views, bias see developmental bias 90–91 bicoid gene see maternal (bicoid) gene legless/shelled tetrapods, 325 Bicyclus anynana (African brown butterfly), body scale/fur colour coadaptation, 191–4 128–32, 212, 243, 254 body size, 124–8, 253 bilateral symmetry, gene co-option, 248 bone morphogenetic (BMPs), bilaterian animals 114, 189 definition, 337 bones, mammalian ear, 151–5 developmental gene examples, 223–5 box jellies (Cubozoa), 240 dorso-ventral axis inversion in vertebrates/ brain size, humans/chimps, 122–4, 128 arthropods, 114–16 Brassicaceae, 84, 85, 337 gene co-option, 238 breeding patterns, semelparous/ Hox gene clusters (Duboule), 228, 229 iteroparous, 64 left–right axis, 249 brown algae, 301, 334 micro-RNA families mapping, 307 Bryozoa (moss animals), 9, 337 protostomes, 167 Burgess Shale, 116, 337 biogenetic law (Haeckel), 16, 17 Butterflies; see also individual species birds (Aves), 151–4, 197–8 Colias butterflies/caterpillars, 53 Darwin’s finches (Geospiza), 56, 58, eyespots, 128–32, 211–13, 243 191, 196 Inachis io (peacock butterfly), 128 beak size/shape, 6, 7, 186–90 Maniola jurtina, 56 feathers, 197, 251 neck structure, 201, 202 cacti, appendages, 242 Pinaroloxias inornata (Cocos finch), cactus gene, 224 186–8 cadherins, 39, 246, 247 sex chromosomes, 58 Caenorhabditis elegans, 35, 105, theropod dinosaurs, relationship, 197 113–14, 347 Biston betularia (peppered moth), 57, 68–71, calmodulin, 190 73, 186, 188, 191, 197 Cambrian explosion, 284, 286–9, Bithorax complex, 226 337, 363 bithorax Drosophila mutant, 21–3, 91 , epigenetic landscapes, 23, 337 , 266, 267 Caniformia, 192 blastoderm, 50, 99, 337, 352 canonical signalling pathways, 46–7, 338 blastula, 37, 38, 337 Capsella bursa-pastoris (shepherd’s purse), blind cave-fish (Astyanax mexicanus), 85, 86 132–3 carapacial ridge, turtle, 282, 283 Blind Watchmaker, The (Dawkins), 293 Carboniferous Period, 173, 176, 364 blood vessel linings, 145, 146 Carnivora, dogs/bears, 192 BMP4 (bone morphogenetic 4), cascades of development, 49–51 189, 190 cascades of interacting genes, 137, 138 body form, coordinated change, 20 case study, definition, 338 body plans caste formation, social insects, 261 Cambrian explosion, 285, 287–290 caudal gene, 32, 50, 214, 359 chelonians, 215 Caudal protein, 50 definition, 337 cave-fish (Astyanax mexicanus), 132–3 direction of evolution, 284–9 cell differentiation, 37–40, 338 Evo–Devo explanations, 334 cell signalling see signalling pathways

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cell-level developmental processes, 34–53 chelonians (turtles/tortoises), 215, 216, 282–4 basics, 37–40 Cheverud, Jim, 332–3 death/apoptosis, 38–40, 337 chick (Gallus), 35, 149, 167, 174, 190 differentiation, 37–40, 338 Chilopoda see centipedes gene structure/expression, 40–45 chimpanzee (Pan troglodytes), 122–4 migration, 38, 39 Chinese lantern-type flowers, 116–17 proliferation, 37–8 chirality, 109–11, 159, 267, 338, 366–8 signalling pathways, 45–8 Choanoflagellata, 12, 13, 247, 338 long-range, 51–3 Choloepus didactylus (two-toed sloth), 63 variety of cell types, 39, 295–6, 306 Chordata, 115–16, 164, 166, 338 heterotypy chordin gene, 114 centipede poison claw, 144 chromatin, 138, 338 new cell types, 144–7 chromosomes, 41, 79–80, 349 cell–cell adhesion, 246, 247 Cichlidae, opsin genes, 101–2 centipedes Cidaroids, 8 , distal-less gene, 11 Cirripedia, vestigial abdomen, 125–8 engrailed gene expression, 44, 45 cis-regulatory regions, 137, 156, 338 fossil evidence, 141 clades, 12, 338 gene expression stripes, 85, 127, cladistics school, 161, 163, 338 166, 214 cladogenesis, 54, 274, 338 heterochrony, 95, 98, 99 cladograms, 301, 302, 338 leg-bearing segments, 83, 84, 213, classic evolutionary concepts, 319–20, 325 214, 257 classification meristic variation, 257–9, 262 developmental genes, 222–3 segmentation, 169 Evo-Devo information, 30, 32 developmental bias, 213, 214 naming genes, 359–62 heterochrony, 95–9 signalling proteins, 222 meristic variation, 257–9, 262 cleavage, 18–19, 110, 338 poison claw modification, 141–4 clock and wave-front model, 167, 168 Strigamia maritima, 11, 169, 214, clusters, Hox genes, 228–9 257–9, 262 Cnidaria centromeres, 67, 80, 338 Cubozoa (box jellies), 240 Cepaea nemoralis (land snail) engrailed gene homologue, 238 altered pigmentation, 139–40 new cell types, 145 life cycle complexity, 304 coadaptation population studies, 55–60 adaptation, distinction, 203 reproductive isolating mechanism, 276–8 definition, 338 speciation, 278–9, 281 developmental/functional, two characters, Cephalochordata, 115, 163, 338 191–4 Cerion (West Indian snail), 267 embryogenesis, 193 cervical vertebrae, number/size, 201–3 gene-level forms, 230–31 characters mammalian vertebrae, 203 dependence/independence (Darwin/ modifier genes, genetic Wallace), 208–9 accommodation, 267 value change during development, 7 Whyte’s ideas, 23 weighting in evolutionary trees, 368 coastal centipede (Strigamia maritima), Chelicerata (arachnids), 168 257–9, 262

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Cocos Island, 186, 187 antennapedia gene, 84, 87 codevelopment, gut bacteria, 269–271 barnacles, vestigial abdomen, 125–8 coding DNA, 41, 42, 81 crabs, 89, 111, 125, 241 coding genes, 136, 138–9 embryos, distal-less gene, 11 coding regions, 81, 231–3 engrailed gene expression, 44, 45 cohorts, 64, 339 morphological disparity, 125 cold-acclimation, 138–9 Pancrustacea, 87, 304, 348 Colias butterflies/caterpillars, 53 Thompsonian transformations, 89 colinearity phenomenon, 27, 339 Ctenophora (comb jellies), 13, 339 collar cell of sponge, 13–14 Cubitus interruptus (Ci) protein, 48 colonial groups, tunicates, 59 Cubozoa (box jellies), 240 colouration, body fur, 191, 192 cusp patterns, rodent teeth, 117 comb jellies (Ctenophora), 13, 339 cyc gene, 244, 245 comparative embryology, nineteenth century, 16–19, 313 Darwin, Charles, 18, 312, 313, 319 compensation, heterometric, 132–4 correlation of growth, 20–21 complexity dependence vs. independence of characters, evolution of, 291–308 208–9 adults, 299–302 finches (Geospiza), 56, 58, 186–91, 196 decreases/increases, 297–302 genetic theory, 4, 6 definitions, 291–3 phylogeny illustration, 161, 164 life-cyle complexity change, 302–5 Darwin’s Finches (Lack), 188 measures of complexity, 295 daughter species, 7, 274, 281, 320, 339, 351 molecular level, 306–8 Dawkins, Richard, 293, 358 spurious Law of Increasing Complexity, de Beer, Gavin, 16, 21, 22, 24, 109 293–9, 302 deletions, 78, 80, 81 repatterning, 148–9 delta gene, co-option, 237 component parts, numbers/types, 292 Democricetodon (LCA of mice/voles), 118 compound nature of evolutionary dental epithelium, 113; see also teeth change, 179 denticles, Drosophila spp, 155–7 conformation, chromatin, 138 dependence/independence of characters conspecifics, 12, 339 (Darwin/Wallace), 208–9 constraint see developmental constraint desert locust (Schistocerca gregaria), 254 continuous developmental plasticity, 252 deuterostomes, 18, 19, 115, 288, 339, 368–9 continuous variation, 86–7, 185–90, Chordata, 115–16, 164, 165, 338 259–60, 355, 356 Hemichordata, 115, 343 convergent evolution, 204, 339 Development and Evolution (de Beer), 21 co-option see gene co-option developmental bias coordinated changes, Thompson’s theory, definition, 339 20–21 direction of evolution, 200–17 Copepoda, 89, 126 routine evolution, 211–15 courtship behaviour, 278, 279 gene co-option, 249–51 covariance matrix, 196 gene level, 230–31 covariation, 206–10, 212–13 origin of novelties, 216–17 crabs, 89, 111, 125, 241 origin of term, 205 cross-vein, Drosophila, 265, 277 meets Evo–Devo, crustaceans 209–11

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developmental bias (cont’d) developmental repatterning, 10, spandrels, 205, 332, 351, 358 121, 340 terminology, 207, 208 between levels of organisation, developmental constraint 155–7 definition, 339 categories/subcategories, 157–9 direction of evolution, 200–17 causes, 159–60 neck structure in mammals, 202–3 chapters overview, 75–6 quantitative genetic theory, 332–3 complex process, 148–9 role in evolution, 131 definition, 82 spandrels paper, 205, 332, 351, 358 different kinds producing similar results, terminology, limitations/ 149–51 misunderstandings, 204–8 directionality, 87 developmental drive, 206–7, 339 fitness relationship, 89–92 developmental genes four component types, 11 altered products, heterotypy, 137–9 integrative nature of, 148–61 bias/coadaptation at gene level, 230–31 key relationships, 323 BMP4/calmodulin, 190 mapping to phylogenetic trees, classification, 222 161–80 definition, 339 molecular level examples, 82–8 developmental plasticity, 261–4 mutations, 77–92 developmental/genetic level, 218–19 single level of organisation, 151–5 examples, 223–5 standing variation, 72–3 Hox genes, 225–30 terminological suitability, 10 interactions/regulation, 137 types, 32 naming, 359–62 whole organism level, 88–9 neutral evolution (Kimura), 102 developmental selection (Whyte), 193, 203; overview, 219–23 see also internal selection regulatory vs. coding change, 231–3 developmental trajectory, 6, 9–11 taxon sampling, 330 definition/usefulness, 52, 340 transposition mutations, 79 developmental repatterning, 82, 87–8 developmental level, 21 8–9 Drosophila, 23 developmental patterning, 340 human brain size increase, 124 developmental plasticity Waddington’s views, 21–4 co-development, evolution, 269 Devonian Period, 170–71, 173 developmental genes, 261–4 dextral/sinistral snail shells, 109–10 direction of evolution, 270–71 Diapsida (snakes), 112–13, 154, 165 discrete variants, winged/wingless insect dichotoma gene, 244 forms, 255–7 Dicyemida, 297–8 genetic accommodation, 267 differentiation of cells, 37–41, 112, 338 genetic assimilation, 264–6 Dimidiochromis compressiceps, 103 meristic variation, 253, 257–9, 262 Diploidy, 18, 55, 57, 60, 340 patterns, development, 261–71 Diptera (flies), 57, 58, 227, 340; see also plant growth, continuous variation, Drosophila spp. 259–60 direct development polyphenism, experimental evolution, 269 age structure of populations, 64 reaction norms, evolution, 266–7 body size, developmental plasticity, 254 types, 252–4 complexity change, 302

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definition, 9 antennapedia gene, 26, 83–4, 87 phylogenetic positions, 3 bithorax mutant, 21–3, 91, 266, 267 directional selection, 71–2, 186 caudal gene, 32, 50, 214, 359 discrete developmental plasticity, 252 chromosomal inversions, 80 discrete variants, 254–6, 355, 356 D. mauritiana, 155, 157 Dishevelled protein, 46, 47 D. melanogaster, 22, 95, 99–101 disorganized Hox gene cluster, 228, 229 abdominal pigmentation, 263–4 disruptive selection, 186 antennapedia gene, 26 distal-less (dll) gene, 11, 32, 131, 242, bithorax mutant, 21–3, 266, 267 243, 361 developmental plasticity, 253 distributions egg, 35 geographical, 54–5, 60–63, 275, 276, genetic assimilation, 264–6 278, 343 hedgehog mutant, 27 statistically normal, 86, 87, 185–6 , classification, 32 divergence svb gene, 156, 157 duplication and divergence, 244, 248, Wingless signalling, 48–9, 50 307–8, 340 D. pseudoobscura, 99–101, 275–6, 278 evolutionary trees, 6 population studies, 56, 57 von Baerian, 16–19, 317, 318, 353 D. sechellia, 155–7 dll (distal-less) gene, 11, 32, 131, 242, 243, 361 D. simulans, 99–101, 155, 157 dlx family of genes, 243 denticles, 155–7 DNA developmental genes, classification, 223 altered sequence mutations, 77–80 distal-less expression, 243 binding motifs, 220, 221 dorso–ventral axis establishment, 223–5 DNA–histone complex, 41 eye development, 225, 226 eukaryotic, human, 66–7 gene nomenclature, 359–61 fingerprinting, 79 homeotic transformations, 225–6 homeobox sequence, 25, 26, 30, 32, 220, Hox gene complexes, 227 221, 344 pair-rule genes, 99–101 methylation pattern, inheritance, 78 segmentation molecular clock hypothesis, 287–9 heterochrony, 97 repatterning, mutations, 77–92 origins, 167, 169 Dobzhansky, Theodosius, 55, 56, 356 genes, 221 domains, 340 Duboule, Denis, 228, 229 homeodomains, 25, 26, 220, 221, 344 dung flies/beetles, 204 , 71, 340 duplication and divergence, 244, 248, dorsal gene, 224, 248 307–8, 340 dorso–ventral axis bilaterian animals, 249 ear bones, mammalian, 151–5 Drosophila, 223–5 early developmental stage, evolution, 8 heterotopy, 112–16 Earth’s history, 363–9 inversion, vertebrates/arthropods, 114–16 ecdysone, metamorphosis, 52 double helix DNA structure, 41 Ecdysozoa, phylogeny, 166, 167 downstream regulatory regions, 137, 340 echinoderms dpp (decapentaplegic) gene, 114, 224, 248 dorso–ventral axis inversion, 115 dromeosaurs, 197 engrailed gene co-option, 238 Drosophila spp. larvae

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echinoderms (cont’d) Eumetazoa, terminology use, 296 phylogenetic tree, 8 Eusthenopteron, 172 yolk/plankton feeders, 8, 334–5 even-skipped (pair-rule) gene, 50, 99–101, ecological genetics approach, 10 221, 223 eco–devo school of thought, 91–2 Evo-Devo ectopic expression, 83, 225, 226, 340 advent in 1980s, 25–7 egg-laying, nematode, 113–14 broad and narrow views, 27–8 egg-timer concept, 317–19, 330 foundations, 313 Elapidae, evolution, 112–13 history/overview, 15–32 Eldredge, Niles, 281 integrative, 332–4 embryogenesis; see also larvae key concepts and connections, 311–26 centipedes, 95 molecular level, 327–31 poison claw segment, 142–3 tenets, 163, 164 coadaptation, 193 Evolution of Man, The (Haeckel), 16 definition, 340 evolutionary trees, 6, 366–9; see also distal-less gene expression, 11 phylogenetic trees ear bones, formation, 151 concept (Gerhart/Kirschner), hominid brain size, 124 219, 323–4, 341 nauplius larva, 125–7, 347 exaptation nineteenth century comparative definition, 196, 341 embryology, 16–19, 314 direction, 196–9 recapitulation, 15–18, 21, 22, 24, 109, flowers/limbs, 179 350 gene co-option, 249–51 Sonic Hedgehog signalling, 149–51 origin of term, 127 Embryos and Ancestors (de Beer), 21 origins of multicellularity, 12 endo-symbionts, 41, 269, 270, 341 exons, 42, 137, 341 endosperm, 105, 341, 349 expression patterns, 341 endothelial cells, 145–7 stripes, 84, 85, 127, 214 engrailed (segment polarity) gene, 35 extant, definition, 341 centipede, 262 extremist views on evolution, 162 co-option, 235, 237–9 exon, 251 expression, 44, 45 eyeless gene, 225, 226 stripes, 84, 85, 126, 127, 214 eyes, 107–9, 125, 132–3, 225, 237–40 heterochrony, 97 eyespots, butterfly, 128–32, 211–13, 243 enhancers, 137, 232, 350 environmental factors, 6, 7, 192–3, 252–71 FA (fluctuating asymmetry), 6, 259, 342 enzyme-producing genes, 356 feathers, 197, 250 epigenesis, 341 Feliformia, 192 epigenetic landscape, 21, 23, 341 FGF (fibroblast growth factor), 168 , 78, 160, 341 FGFR (fibroblast growth factor ), 87 Epimorpha, heterochrony, 96–7 fingerprinting, DNA, 79 epithelium, 40, 113, 118, 341 fish Erwin, Douglas, 272 Astyanax mexicanus (blind cave-fish), 132 ether vapour, 23, 264 Cichlidae, opsin gene heterochrony, 101–2 Eucalyptus trees, 261 fossil jawless, 286 euechinoids, 8 lobe-finned, phylogeny, 172 Eukaryota, 40–45, 66, 67, 341 Pleuronectiformes (flatfish), 107–9

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recapitulation, 22 mammalian ear bones, evolution, teleosts, 107, 132–3 151, 152 transformations (Thompson), 20 rodent teeth heterotopy, 117 Fisher, Ronald, 90, 91, 356 tetrapods, 171–4 fitness; see also natural selection turtle (Odontochelys), 284 accelerated segment formation in foxes, 63, 191, 192 centipedes, 97–9 frame-shift mutations, 81 continuously variable characters, 186 Frizzled receptor, 46, 47, 49 definition, 342 frog, 35–7, 304, 305 developmental repatterning, 89–92 front-fanged snakes, 112–13 developmental trajectory, 6, 8 fruit fly see Drosophila spp. direction of evolution, 200–201 ftz (fushi tarazu) gene, 223 heterochrony, 102–5 fungi, 12, 40, 301, 334 mammalian neck structure, 203 fur colour, bears/foxes, 192 peaks in an adaptive landscape/seascape, 190–91 G matrix, 210, 211 polymorphisms, 71 Galapagos Islands, 186, 187, 196 fixation, alleles, 72 Gallus (chick), 35, 149, 166, 167, 174, 190 fixed-phenotype evolutionary theory, 263 gametophytes, 60 flamingo neck structure, 201, 202 Gastropoda, 109–11, 139–40, 304, 305, flatfish (Pleuronectiformes), 107–9 342; see also Cepaea nemoralis; snails flies (Diptera), 57, 58, 227, 228, 340; see also gastrulation, 37, 38, 342 Drosophila spp. gel electrophoresis, 66, 79, 356 flounder, 107 general evolutionary theory, 332–4 flowers genes; see also mutations ABCDE model, 86 co-option, 234–51 diversity, 177 appendage evolution, 241–3 flowering time, heterochrony, 105 definition, 234–7 origins, 176–9 exaptation/developmental bias, 249–51 typical parts 176, 178 genetic toolkit concept, 245–49 zygomorphic, gene co-option, 244–5 molecular taxon sampling, 330 fluctuating asymmetry (FA), 6, 259, 342 segments/eyes evolution, 237–40 food supply, larval body size, 253 zygomorphic flowers evolution, 244–5 forelimb/hindlimb length co-variation, deletions, 78, 80, 81 206, 207 duplication, 149, 223, 342; see also fossil evidence/record duplication and divergence angiosperms, 176 expression barnacles, 127 cell differentiation, 38 Cambrian explosion, 284, 286–9, developmental processes, 40–45 337, 363 ectopic, 83, 225, 226, 340 centipedes, 142 embryos, distal-less gene, 11 geological time, 363–5 engrailed stripes, 84, 85, 127, 166 sinistral/dextral snail shells, 111 heterochrony, 93–105 trilobites, 332, 333, 365 heterotopy vertebrates, 286, 287 dorso-ventral axis inversion, 114 fish, 108, 171, 172, 174 Physalis, 116–7 human brain size increase, 123, 124 location of genes, 40–42

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genes; see also mutations (cont’d) greenflies 254, 255 regulation, complexity measures, 307 groups of genes, naming, 361 segmental stripes, 84, 85, 127, 166, 214 gut bacteria, symbiotic, 269, 270 switching on genes, 42, 43, 45 gymnosperms, 105, 343 insertions, 78, 81 naming conventions, 359–62 Haeckel, Ernst, 121, 294, 313 regulatory, 41, 81, 137, 231–3, 350 law, 16, 18, 19 three classes, 220 haemoglobin, 135–6 genet individuals, 58, 342 hairy gene expression, 99, 100 genetic accommodation, 267 Haldane, J.B.S., 19, 24–5, 356–8 genetic assimilation, 264–6, 342 Haldane’s rule, 276 genetic code, 81 Hanski, Ilkka, 61 genetic drift, 66, 80, 102, 103, 320, 342 haplodiploidy, 60, 61 genetic imperialism problem, 357–8 haploidy, 60, 343 genetic structure of natural populations, Harpinia plumosa, 89 65–7 head size, humans, 88 genetic toolkit concept, 245–49 hearts, vertebrates, 145, 146 genic DNA, 41, 66 heat shock, 264, 265, 268, 269 , 219–20, 306, 342 heavy metal tolerance, 58 genotype–developmental pattern map, 90 hedgehog gene, 27, 47–9, 84, 149, genotype–fitness map, 90 150, 361 genotype–phenotype map, 89, 342 Hedgehog pathway, 47–8 Geoffroy Saint-Hilaire, Etienne, 114 height distribution, humans, 86–7 geographical range, 54, 60–61, 63, 274–7, Helicidae (gastropods), 139–40 343 Heliocidaris spp., 8 geological time, 363–5 helix-loop-helix, 220, 221 Geophilomorpha, see centipedes helix-turn-helix, 26, 220, 221 Geospiza spp. (Darwin’s finches), 56, 58, Hemichordata, 115, 343 186–91, 196 hemimetabolous development, 52, 343 Geranium carolinianum, 259, 260 Hemiptera, aphids, 255–7 Gerhart, John, 219 Hennig, Willi, 161 germ band, 18, 343 herding behaviour, 61 germ layer, 94, 342, 343, 346 , 4, 77–8, 343, 355, 356 germ-line mutations, 78 hermaphrodites, 58 giant (gap) gene, 50 heterochrony, 93–105 Gilbert, Scott, 10 Arabidopsis/Linaria, 245 giraffe, 201 categorisation, 32 glyptodonts, 216, 343 classification, 158 Goldschmidt, Richard, 21, 22, 75, 83, 90–91, de Beer’s theory, 21 334, 357 definition, 93–4, 343 Goodwin, Brian, 214 fitness, 102–5 Gould, Stephen Jay, 23–4, 109, 197, 203, Gould’s ideas, 24 205, 207, 211, 267, 280, 332, heterochronic genes/mutations 333, 358 terminology, 105 gradual evolution hypothesis, 108 molecular level, 99–102 gradualist views (Fisher), 90, 343 off-putting terminology, 94 grass (Agrostis tenuis), 58 organismic level, 95–9

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over-emphasis on, 9, 10 geological time, 363–5 repatterning component, 11 nineteenth-century comparative types/levels, 94–5 embryology, 16–19 Heterokontophyta, 334 holometabolous insects, 52, 131, 303, 304, heterometry, 121–34 344; see also Diptera; Lepidoptera bi-directional, 128–32 HOM complexes, 226 categorisation, 32 homeobox classification, 159 definition, 344 compensation, 132–4 discovery/characteristics, 25, 26 decrease in relative size, 124–8 Drosophila, 30, 32 definition, 121–2, 343 transcription factor genes, 220, 221 increase in relative size, 122–4 homeodomain, 25, 26, 220, 221, 344 repatterning component, 11 heterotopy 106–20; see also homeosis arthropod segment identity genes, 83 Arabidopsis/Linaria, 244, 245 bithorax Drosophila, 91 A–P/D–V axes, 112–16 brassicacean flowers, 84, 85, 87 categorisation, 32 definition, 344 classification, 159 transformations, Drosophila spp., 91, definition, 106–7, 343 225–6 left-right asymmetry, 107–11 Homo spp.; see also humans repatterning component, 11 H. floriensis, 128 Thompsonian transformations, 88–9 H. habilis, 123, 124 heterotypy, 135– 47 H. heidelbergensis, 123 altered morphology/novelty origins, 140, H. neanderthalensis, 123, 128 142, 144 H. rudolfensis, 123 altered pigmentation, 139–40 H. sapiens, brain size, 122–4, 128 altered products of developmental genes, homology of genes 137–9 classic concept, 320 categorisation, 32 co-option, 237 classification, 159 definition, 344 definition, 135–6, 343–4 eye development, 225 Drosophila sechellia, svb (shavenbaby) gene homoplasy, distinction, 163 regulation, 156–7 human/chimp brain size, 122–3 new cell types, 144–7 mapping repatterning to trees, 162, 163 repatterning component, 11 orthology/paralogy, 250 heterozygous advantage, 72, 136, 344 homonomous segmentation, 292 hierarchies of development, 51 homoplasy, homology distinction, 163, 344 hindlimb/forelimb length co-variation, 206, 207 homozygous state, 18, 55, 66, 111, 136, 344 histones, 41, 344 hormones, 89, 344 history horns, beetles, 133 conclusions based on, 24–5 hos2 gene, 139 Darwin’s evolutionary theory, 4, 6 hour-glass model, 317, 318 Evo-Devo housekeeping genes, 41, 66, 102, 344 advent in 1980s, 25–7 Hox genes fore-runners 1900–1980, 19–24 abdominal pigmentation, 263–4 Fisherian vs. Goldschmidtian views, altered/new role, 235 90–91 antennapedia gene, 83, 84

99781444337204_6_index.indd781444337204_6_index.indd 339393 111/5/20101/5/2010 11:41:3811:41:38 AMAM 394 Index

Hox genes (cont’d) engrailed expression, 44, 45, 84, 85 antero–posterior body expression, 27 germ band, 18 centipede poison claw segment, 142 metamorphosis, 52 definition, 344 pattern–fitness, 91 Drosophila, 32, 227, 263–4 phylogeny, 303 Duboule cluster types, 228, 229 segmentation origins, 166, 167, 169 fins-to-limbs phylogeny, 174, 175 social caste formation, 261, 262 genetic toolkit concept, 246 syncytial cleavage, 19 mammalian vertebrae, 203 winged/wingless forms, developmental overview, 221, 225–30 plasticity, 254–7 humans zygomorphic flower pollination, 244 insertions, point mutations, 78, 81 recapitulation of evolution, 17, 18, 22 instars, 64, 268, 344; see also larvae webbed hands/feet, 40 integrative nature of repatterning, gene nomenclature, 360, 361 148–60 head size, 88 integrative Evo-Devo, 332–4 height distribution, 86–7 integrins, 39, 345 populations interactions, component parts of organism, DNA types, 66–7 292 five characteristics, 55, 57 interbreeding, species definition, 273 sickle cell anaemia, 136 internal selection, 191, 193, 194, Tree of Life (Haeckel), 294, 295 203, 204 Huxley, Julian, 88 introns Huxley, T. H., 3 definition, 345 hybrids, 277–279 developmental genes, 137, 138 Hydatellaceae, 103–5, 344 eukaryotic, 42, 43 Hymenoptera, 60, 61, 261 mutations, 81 Hyperia galba, 89 shuffling, 250 invected gene, 239 imaginal discs, 83, 131, 344 inversions, chromosomal, 80 imperfect recapitulation, 22 islands, population studies, 61, 63 Inachis io (peacock butterfly), 128 isolating mechanisms, 276, 277, 279 incus bone, 151, 152, 154 isometric changes, 88, 345 indirect development; see also larvae italic script, gene names, 359–61 age structure of populations, 64 iteroparous breeding patterns, 64 complexity definition, 292, 302 definition, 9 jaw joint, mammalian ear bones, developmental plasticity, 253 151–5 genes, 220 jaws, mouse, 194–6 phylogenetic position, 303 Jeffreys, Alec, 79 Indohyus (extinct mammal), 6 jellyfish, poison cells, 145 Indri indri (indri lemur), 63 journal publications, Evo-Devo, 28, 31 induction, nematode vulval position, 113 industrial melanism, 10, 68–71, 73, 186, karyotype, 345, 356 188, 191, 197 key concepts insects; see also Drosophila spp.; individual species Evo-Devo/classical, 312–26 antp expression, 83, 84 list, 314–16

99781444337204_6_index.indd781444337204_6_index.indd 339494 111/5/20101/5/2010 11:41:3811:41:38 AMAM Index 395

Keywords and Concepts in Evolutionary population studies, 55, 58 (Hall/Olson), Leroi, Armand, 272 317 leucine zipper, 220, 221 Kimura, Motoo, 102 levels of organisation, repatterning between, kinorhynch, 164, 165 155–7 Kirschner, Marc, 219 Lewis, Ed, 226 Klingenberg, Chris, 194, 196 Lewontin, Richard, 205, 211, 332, 333, 356, 358 Lack, David, 188 life cycles Lamarckian theory, 23, 201, 313 advantageous forms, 7, 8 land snail see Cepaea nemoralis Colias butterflies/caterpillars, 53 landmark features, 194–6 complexity of organism larvae; see also indirect development change, 302–5 butterflies, development, 131 definition, 291–3 complexity, definition, 292 haplodiploid, 60, 61 definition, 345 Schistosoma, 293 Drosophila spp., 155–7, 160, 253 Xenopus, 37 echinoid, feeding/non-feeding, 8, 334–5 /receptor groups, 45–7, 345 Manduca sexta, polyphenism evolution, light stimulus, plants, 260–62 267, 268 limbs; see also appendages phylogenetic position of insects, 303 leg-bearing segments, 83, 84, 213, 214, 257 population studies, 57 legless tetrapods, 325 laws length co-variation, 206, 207 biogenetic law, 16 origins, 171–7 characteristics/requirements, 18, 19 terminology, 241, 243 patterns rather than, 24 trunk outgrowths, 236 of recapitulation (Haeckel), 16 Linaria, 244, 245 spurious universal laws, 293–9, 302, 335 lineages, 293–5, 299–302, 330, 345 von Baer’s laws, 16 linear reaction norms, 266 LCA (last common ancestor), 163, 170 Lithobiomorpha (centipedes), 96, 98, 213 leaf arrangements, angiosperms, 214–15 Lithobius spp., 96 leafy gene, 177 Littorina (sea snail), 304, 305 left-right axis, 107–111, 160, 249, 267, 338, lobe-finned fishes, 171 366–8 locus (gene), 345 lefty gene, 111, 248 locust (Schistocerca gregaria), 255 leg-pairs, centipedes, 213–14, 257 long-germ development, 168, 345 legless tetrapods, 325 long-range signalling, 51, 53 Lepidoptera (butterflies/moths) long-term evolution, direction, 196–9 bi-directional heterometry, 128–32 Lophotrochozoa, 166, 167 Bicyclus anynana (African brown butterfly), loss of structures, 125 128–32, 212, 243, 254 Lymnaea (water snail), 64, 110, 304, 305 Biston betularia (peppered moth), 57, 68–71, 73, 186, 188, 197 MacArthur, Robert, 55 Colias butterflies/caterpillars, 53 macro-evolution, 273, 282, 345 eyespots, 128–32, 211–13, 243 macromutation, 334 Inachis io (peacock butterfly), 128 definition, 346 Maniola jurtina (butterfly), 56 direction of evolution, 280

99781444337204_6_index.indd781444337204_6_index.indd 339595 111/5/20101/5/2010 11:41:3811:41:38 AMAM 396 Index

macromutation (cont’d) mesoderm, 48, 167, 169, 282, 346 Evo-Devo explanations, 334 mesomutations, 87, 280, 346 flatfish, left-right asymmetry, 108 Mesorhabditis, vulval position, 113 role in novelty, 216 Mesozoa, use of term, 296, 297 MADS-box genes, 25, 116, 117, 176, 346 Mesozoic Era, 365 mads16 gene, 116, 117 messenger RNA, splicing, 43 malaria, sickle cell anaemia, 136 metamorphosis, 52, 346 Malawi, Lake, 101 metapopulations (Hanski), 61, 63, 346 male genetic polymorphism, pea aphid, 256, Metazoa, use of term, 296 257 methylation, DNA, 78 malleus bone, 151, 152, 154 Metriaclima zebra, 103 mammals; see also individual mammals Mexican blind cave-fish (Astyanax mexicanus), ear bones, 151–5 132, 133 neck structure, 201–3 micro-evolution, 282, 334, 335, 346 manatee, vertebrae number, 201, 202 micro-RNAs (miRNAs), families, 307 mandible, mouse, 194–6 microcephalin gene, 122 Manduca sexta (tobacco hornworm), 267, 268 micromutation, 87, 346 Maniola jurtina (butterfly), 56 Microtus rossiaemeridionalis (vole), 117, 118 mapping repatterning to trees, 161–80 midge larvae, 57 flower origins, 176–9 migration of cells, 38, 39 pattern/process/homology, 161–3 millipede segmentation origins, 164 segmentation of animals, 163–9 mimicry, 57, 346 vertebrate fin-to-limb transition, 169–75 Minelli, Alessandro, 8 marsh grass (Spartina townsendii), 275 model organisms, 35, 36, 56–8, 346, 359–61 mass extinction events, 365 modern synthesis school, 161, 162, 346 maternal genes, 50 modifier genes, 267 bicoid, Drosophila egg, 35 , 9, 51, 317–19, 332, 347 Bicoid proteins, 50 molecular clock hypothesis, 287, 288 Drosophila spp. segmentation origins, molecular level 167, 168 complexity evolution, 306–8 naming, 359 developmental processes, 34–53 sinistral/dextral snail shells, 110 Evo-Devo, 313, 327–31 mating disadvantage, snail shell chirality, heterochrony, 99–102 110, 111 heterotopy, 108, 117 Maynard Smith, John, 205 heterotypy, 135–6 Mazoscolopendra, 97 repatterning Medawars, Peter and Jean, 162 examples, 82–8 mega-evolution concept (Simpson), 196, integrative nature, 149–51 273, 346 Mollusca; see also snails melanism engrailed gene co-option, 238, 239 definition, 346 left-right asymmetry, 109–11, 338, 366–8 industrial, 10, 68–71, 73, 186, 188, 191, partial segmentation, 164, 165 197 monophyletic groups, 141, 186, 347 Mendel, Gregor, 18, 86, 231, 312, 313, monoplacophoran molluscs, 164, 165 355, 356 Monosiga brevicollis (unicellular meristic variation; see also segmentation choanoflagellate), 246, 247 developmental plasticity, 252, 257–9, 261 Morganucodon, 152, 153

99781444337204_6_index.indd781444337204_6_index.indd 339696 111/5/20101/5/2010 11:41:3811:41:38 AMAM Index 397

, 89, 347 natural populations, 54–73 morphological disparity, 125 natural selection morphological geometry, 194–6 butterfly eyespot size/colour, 211–13 morphospace, 12, 347 centipede leg-bearing segments, 214 mosquito larvae, 57 continuously variable characters, 185–90 moths Evo-Devo, 163 Biston betularia (peppered moth), 10, 57, flowers/limbs, 179 68–71, 73, 186, 188, 197 industrial melanism, 10, 68–71, 73, 186, motifs, DNA binding, 220, 221 188, 191, 197 mouse see Mus musculus interplay with heritable variation, 4 movement long-term evolution, 196–9 cells, 38–9 morphological geometry, 194–6 heterotopy, 158 population studies, 67–73 mpf2 gene, 116 two characters, adaptive landscape, mRNA, 42, 43, 81 190–91 mule, 277 nauplius larva, 125–7, 347 multicellularity, 11–14, 248 neck structure, birds/mammals, 201–3 Muridae, 195 negative co-variation of two characters, Mus musculus (mouse) 206, 207 gene nomenclature, 360 negative heterometry, 155 Hox genes, 226–7 Nematoda, 35, 105, 113–14, 238, 299, 347 flies comparison, 227 neo-sympatry, 278 life cycle, 293 , 21, 32, 94, 347 morphological geometry, 194–6 nervous system, 122–4, 128, 149–50, 238, teeth heterotopy, 117–19 243, 347 mutations networks of development, 51 apparently without consequence, 82 neural plate, 243 classic concept, 319–20, 325 neural tube, 149, 150, 347 coding/regulatory regions, 232 neutral evolution (Kimura), 102 definition, 347 neutral mutations, 66, 72, 102, 103, 133, deletions, 78, 80, 81 267, 288, 347, 356 developmental repatterning, 77–92, 160 new types see novelty developmental trajectory, 6, 10 Nijhout, Fred, 267 Drosophila, 21–3, 27 Nile tilapia (Oreochromis niloticus), 101–3 Evo-Devo, 163 nineteenth-century comparative embryology, four types, 78–80 16–19 frame-shift, 81 Nkx2.2 transcription factor, 149, 150 germ-line, 78 nodal gene, 111, 248 hundreds/thousands of bases, 79 noncoding DNA, 41 neutral, 66, 72, 102, 103, 133, 267, 288, normal distributions, 86, 87, 185 347, 356 notch gene, 237 proximate functional consequences, Notch-Delta signalling, 97, 98, 167–9 80–82 notochord, 149, 285, 286, 347 Myriapoda; see also centipedes novelty relationships, 141, 142 cell types, 144–7 developmental bias, 216–17 Nanos protein, 50 Evo-Devo explanations, 334, 335

99781444337204_6_index.indd781444337204_6_index.indd 339797 111/5/20101/5/2010 11:41:3811:41:38 AMAM 398 Index

novelty (cont’d) Paranthropus spp., 123 heterotopy, 120 parapatric speciation, 274, 275 origins, 140–44, 281–4 paraphyletic groups, 128, 154, 348 role definition, gene co-option, 235–7 parasitic organisms, 296, 297, 299 nucleotide base triplets, 81 Parhyale (crustacean), 11 nucleus, 41, 43, 45 parthenogenetic reproduction, 255 nudibranch (sea slug), 109 particulate theory (Mendel), 18, 86, 231, 312, 313, 355, 356 octopus endothelial cell linings, 145 parts, number/types, 292 Odontochelys (fossil turtle), 283 Partula (snail), 63, 110 Oikopleura, 228 pathways of development, 51 Oithona nana, 89 , 340; see also OLIG2 transcription factor, 149, 150 developmental repatterning On Growth and Form (Thompson), 20, 89 definition, 348 oncogenes, 45, 48–9, 84, 348 developmental plasticity, 265–71 ; see also developmental embryonic cleavage, 18, 19 definition, 348 search for, 55 recapitulates phylogeny, 15–18, 21, 22, 24, von Baerian divergence, 16–19, 317, 318, 109, 350 353 Ontogeny and Phylogeny (Gould), 23, 24 pattern–fitness map, 90 opsin genes, 101–3 , 246 Orchidaceae, 244, 245 gene, 133, 225, 239–40 Oreochromis niloticus (Nile tilapia), 101, 103 pea aphid (Acyrthosiphon pisum), 256 organisational levels, 155, 157 pea plants (Pisum), 86, 87, 348 organised Hox gene cluster, 228, 229 peacock butterfly (Inachis io), 128 organismic level, heterochrony, 95, 99 peacock’s tail, 191 Origin of Species, The (Darwin), 4, 161, pectoral girdle, turtle, 281–3 162, 208 pentadactyl limb, 172 orthogenetics, history, 91 peppered moth (Biston betularia), 10, 57, orthology, homology of genes, 250 68–71, 73, 186, 188, 191, 197 Orthonectida, 296, 297 perfect/imperfect recapitulation, 22 Orthoptera, 254 peripatric speciation, 275 ossicles, mammalian ear, 151–5 Phanerozoic Era, 363, 364 outgroups, 101, 141, 167, 187, 188, 246, phenotype, 4, 7, 189, 211, 348 329, 348 see developmental Owen, Richard, 162 plasticity phyletic constraint, 204, 205, 332 paddlefish (Polyodon), 174, 175 phylogenetic trees paedomorphic salamanders, 94, 95 Amniota, 154 paedomorphosis, 65, 94, 95, 348 angiosperms, 179 pair-rule genes, 50, 99–101, 221, 223 Cepaea (land snail), 278 Pan troglodytes (chimp), 122–4 echinoderms, 8 pan-externalist view, 203, 207 Geospiza (Darwin’s finches), 186, 188 Pancrustacea hypothesis, 87, 304, 348 human/chimp, 123 Panderichthys, 171, 172 insects, 304 pangenesis theory of inheritance, 4 lineages, complexity change, 293–5, paralogy, homology of genes, 249 299–302

99781444337204_6_index.indd781444337204_6_index.indd 339898 111/5/20101/5/2010 11:41:3811:41:38 AMAM Index 399

mapping repatterning to, 161–80 poison cells, Cnidaria, 145 flower origins, 176–9 poison claw segment, centipedes, 141–3 pattern/process/homology, 161–3 Polistes canadensis (social wasp), 261, 262 segmentation of animals, 163–9 polychaete worm, 164 vertebrate fin-to-limb transition, 169–75 polygeny, 189, 280, 349 origins of multicellularity, 12, 13 polymorphism snakes, fang positions, 112 Acyrthosiphon pisum (pea aphid), 256 Tree of Life (Haeckel), 294 Biston betularia (peppered moth), 57, phylogeny, 126, 348 68–71, 73, 186, 188, 191, 197 phylotypic stage, 317, 318, 348–9 Cepaea nemoralis shell colour, 139, 140 phylum, definition, 349 definition, 349 Physalis pubescens, 116, 117 heterozygous advantage, 136 pigmentation history, 356 abdominal, Drosophila melanogaster, population studies, 65–67, 71, 72 263, 264 snail shell chirality, 109–111, 159, 267, altered, heterotypy, 139–140 338, 366–8 Biston betularia (peppered moth), 57, VNTRs, 79 68–71, 73, 186, 188, 191, 197 Polyodon (paddlefish), 174, 175 Manduca sexta polyphenism, 267, 268 polyphenism Pinaroloxias inornata (Cocos finch), 186–8 aphids, 254–6 Pisum (pea plants), 86, 87, 348 butterfly eyespots, 129 pitx gene, 111 definition, 349 Placozoa, 239, 240, 349 experimental evolution, 269, 271 plaice, 107 Manduca sexta, 267, 268 plankton-feeding echinoderm larvae, 8, 334, polymorphism contrast, 254 335 polyphyletic groups, 297, 349 plants; see also flowers; individual species/types polyplacophoran molluscs, 164, 165 appendages, cacti, 241, 242 polyploidy, 80, 349 body plans, use of term, 284 polytene chromosomes, 79, 349 complexity, lineage, 301, 302 pond snails, 64, 110 developmental plasticity, 259–60 populations Hydatellaceae, heterochrony, 103–5 natural, 54–73 kingdom composition/complexity, 328, 329 age structure, 64–5 MADS-box genes, 25 genetic structure, 65–7 model organisms, 35, 36, 85, 138, 139, human, 55, 57, 66, 67 254, 359–61 natural selection, 67–73 origins of multicellularity, 11–14 spatial structure, 60–63 population studies, 60 Porifera, 13 separation of individuals, 58 positional information, 48, 349 Plasmodium, 136 positive co-variation, 206–8 plasticity, 7, 77, 349; see also developmental potato (Solanum tuberosum), 116, 117 plasticity predation Platyhelminthes, 292, 293 Biston betularia (peppered moth), 57, , 189, 349 68–71, 73, 191, 197 plesiomorphy, 349 butterfly eyespots, 130, 131 Pleuronectiformes (flatfish), 107–9 primary structure, 220, 221 point mutations, 78 primordium, 318, 330, 349

99781444337204_6_index.indd781444337204_6_index.indd 339999 111/5/20101/5/2010 11:41:3911:41:39 AMAM 400 Index

progenesis, 32, 95, 349 reproductive character displacement, 278 programmed cell death, 38–40, 337 reproductive darts (land snail), 278–9, 281 prokaryotes, 40, 306, 343, 349 reproductive heterochrony, 94–5, 157, 158 proliferation of cells, 37–8 reproductive isolating mechanism, 276, 277 promoter regions, 42, 117, 137, 350 reproductive openings, 113, 141, 142 proteins reptiles classification, signalling pathways, 222 mammalian ear bones, evolution, 151–5 gene transcription/translation, 45 scales/feathers, 197, 198 haemoglobin, heterotypy, 135, 136 snakes, 112–13, 154, 164 homeodomain, 25, 26 Rhamphoverritor reduncus (fossil barnacle), naming conventions, 359–62 127 protoconch, 140 rhea, neck structure, 201, 202 protostomes, 165, 167, 288, 350, 368–9 Rhizocephala, 125, 126 Protozoa, terminology, 296, 297 ribosomes, translation, 43 pseudogenic DNA, 66–7 ribs, turtles, 282 Pteridium aquilinum (fern), 63 RNA, 42, 43, 81, 306, 307 punctuated equilibrium (Eldredge/Gould), RNA polymerase, 42, 43, 138, 350 128, 280, 350 quantification, 331 rodents; see also Mus musculus (mouse) QTL (quantitative trait loci), 87, 189, 281, teeth, 117–19 350 Rosen, Donn, 162 quantitative genetics, 209–11, 332, 333 Rotifera, 59, 351 queens, Polistes canadensis, 261, 262 routine evolution, 211–15 rudimentary limbs, 236 radial cleavage, 18, 19 ramet individuals, 58, 350 Sacculina carcini (barnacle), 125, 126 ray-finned fishes, 171 salamanders, paedomorphic, 94, 95 reaction norms saltationism (Goldschmidt), 21, 90, 91, 351, definition, 350 357 developmental, 211 Sander, Klaus, 317 three types, 259 Sapphirina, 89 developmental plasticity, 266–7 Sarcopterygii (lobe-finned fishes), 172 direction of evolution, 270 Satyridae, Bicyclus anynana (African brown environmental variation, response, 7 butterfly), 128–32, 211–13, 243, Manduca sexta, 268 254 recapitulation, 15–18, 21, 22, 24, scala naturae approach, 291, 293 109, 350 scale-dependence of evolution, 272–3 recruitment see gene co-option scapula, turtles, 282, 283 red blood cells, 136 Schistocerca gregaria (desert locust), 254 red fox (Vulpes vulpes), 63 Schistosoma life cycle, 293 reductive heterotopy, 159 Scinkidae, 216 regulatory genes, 41, 81, 137, 231–3, 350 Scolopendromorpha, 96, 97, 99 reinforcement see Wallace effect (reproductive Scutigeromorpha, 96 character displacement) sea slug, 109 relative size see heterometry sea squirts (urochordates), 115, 164, 228, 353 repatterning see developmental repatterning seasonal polyphenism, 129, 254 repetitive DNA, 67 seaweed, 334

99781444337204_6_index.indd781444337204_6_index.indd 440000 111/5/20101/5/2010 11:41:3911:41:39 AMAM Index 401

secondary loss of structures, 125 annelids/arthropods, false idea, 167 secondary structure, 220, 221 centipedes, 141 segmentation compound repatterning, 151 animals, origins, 163–9 definition, 351 centipedes deuterostomes, 115 developmental bias, 213–14 divergence, 6 heterochrony, 95–8 Mesozoa, 296 poison claw modification, 141–3 Monosiga brevicollis, 246 Drosophila spp., 49, 50, 97, 221, 331 multicellularity, origins, 11, 12 gene co-option, 237–40 novelties, origin, 216 segment identity, 82–5, 351 Pancrustacea, 348 segment polarity tetrapods, 172 definition, 351 size change see heterometry genes sloth, 201–3 arthropods, 45, 49, 50, 235 slugs, 109 Drosophila, 97, 331 sna gene, 224 striped expression, 84, 85, 127, 168, snails; see also Cepaea nemoralis (land snail) 214 altered pigmentation, 139–40 selective coefficient (s), 72 Cerion (West Indian snail), 267 semelparous breeding patterns, 64 left-right asymmetry, 109–11, 160, 267, sex chromosomes, 55, 58 338, 366–8 sexual dimorphism, 257 life cycle complexity, 304, 305 sexual females, aphids, 255 Littorina (sea snail), 304, 305 sexual selection, peacock’s tail, 191 Lymnaea (pond snail), 64, 110, 304, 305 shark, 174, 175 Partula (Pacific island snail), 63 shell reproductive isolating mechanism, 277 chelonians, 281, 282 snakes, 112–13, 154, 164 snails, chirality, 109–11, 159, 267, 338, SNPs (single nucleotide polymorphisms), 67 366, 368 social insects, 262 tetrapods, 325 sog (short gastrulation) gene, 114, 224, 248 shepherd’s purse (Capsella bursa-pastoris), 85–7 Solanaceae, 116 short-germ development, 168, 351 Solanum tuberosum (potato), 116, 117 short-term evolution, 185–90, 191–4, 196 soma, 351 sickle cell anaemia, 136 somatic development, 94, 95 signalling pathways somatic heterochrony, 158 basics, 45–8 somitogenesis, 167, 168, 351 classification, 32 Sommer, Ralf, 113 definition, 351 sonic hedgehog (shh) gene genetic toolkit concept, 246 rodent teeth heterotopy, 118, 119 long-range signalling, 51–3 snake fang positions, 113 Notch-Delta signalling, 97, 98, 167–9 vertebrate eye development, 133 proteins, classification, 222 Sonic Hedgehog signalling, 149, 150 Sonic Hedgehog, 149–52 spandrels paper (Gould/Lewontin 1979), 205, Simpson, G.G., 196, 273 332, 351, 358 sinistral/dextral snail shells, 109, 110, 338, Spartina townsendii (marsh grass), 275 366, 368 spatial development changes see heterotopy sister-groups spatial distributions, populations, 62

99781444337204_6_index.indd781444337204_6_index.indd 440101 111/5/20101/5/2010 11:41:3911:41:39 AMAM 402 Index

spe gene, 85 syncytial blastoderm, 50, 99, 296, 352 speciation, 72, 186–90, 320, 351 systems of development, 52 species; see also evolutionary trees; lineages; phylogenetic trees Tabin, Clifford, 189, 190 cladogenesis, 54 tandem repeats, 78–9 distributions, red fox/indri lemur/two-toed taste buds, blind cave-fish, 132, 133 sloth, 63 taxon/taxa, 328–30, 352 origins, 273–81 TCP family of transcription factors, 244, 245 speciose, definition, 351 tcp1 gene, co-option, 244 spiders, 44, 50, 168, 169 teeth spiral cleavage, 18, 19, 110 rodents, 117–19, 194–6 splicing process, 42, 43 snake fangs, 112–13 split Hox gene cluster, 228, 229 teleosts, 107, 132–3, 352 sponges, 13 telomere, 67, 352 sporophytes, 60 temperature squid, 145 Drosophila melanogaster abdominal stabilising selection, 186 pigmentation, 264–5 stage structure, populations, 64 Manduca sexta polyphenism evolution, stamenoid petals gene, 85, 361 267–9 standing variation, 72–3, 216, 219, 351 meristic variation, 257–9, 261, 262 Stanley, Stephen, 280 temporal expression differences see stapes bone, 151, 153 heterochrony statistical distributions, 185, 186 Teratorhabditis, 113 Stegocephalus inflatus, 89 terminal target genes, 41, 352 stem cells, 38, 352 terminology sterile offspring, 277, 278 appendages/limbs, 240, 242 Sternberg, Paul, 113 change in developmental trajectory, 10–11 stinging cells, Cnidaria, 145 developmental, 51–3 stmads16 gene, 116 developmental constraint, 204–8 stochastic processes, 61, 324, 352; see also Eumetazoa, 296 genetic drift gene nomenclature, 359–62 Strepsiptera, 91, 303, 352 heterochrony, 94, 95, 105 Strigamia maritima (coastal centipede) internal selection/coadaptation, 194 embryo, distal-less gene, 11 Mesozoa, 296, 297 meristic variation, 257–9, 262 Metazoa, 296 segmentation, 169, 214 Protozoa, 296, 297 striped expression, 84, 85, 127, 166, 214 repatterning of development, 158, 159 substitutions, point mutations, 78 upstream/downstream, 137 svb (shavenbaby) gene, 156, 157, 159 tetrapod vertebrates swan neck structure, 201, 202 chelonians, 215, 216, 281–4 switching on genes, 42–6 definition, 352 symbiotic gut bacteria, 269, 270 hindlimb/forelimb length co-variation, symmetry/asymmetry 206–7 chirality, 109–11, 159, 267, 338, 366–8 legless/shelled, 325 plants, 260–61 phylogeny, 170 sympatric speciation, 274, 275 transition from fishes, mapping to trees, synapsid skulls, 154 169–75

99781444337204_6_index.indd781444337204_6_index.indd 440202 111/5/20101/5/2010 11:41:3911:41:39 AMAM Index 403

theropod dinosaurs, 197 tunicates, 59, 353 Thompson, D’Arcy, 19, 20–21 turbot, 107 Thompsonian transformations, 88–9 turtle, 215, 216, 281–4 Thoracica, 125–7 two-toed sloth (Choloepus didactylus), species thyroid hormone, 108 distribution, 63 Tiktaalik, 171, 172, 174 Tyrannosaurus rex, 207 tilapia, 101–2, 103 timing change see heterochrony unicellular organisms tinman gene, 224 choanoflagellate (Monosiga brevicollis), toad, 305 246, 247 tobacco hornworm (Manduca sexta), 267–9 complexity questions, 293, 302, 303 tomato plants, 189 multicellular origins, 12 tortoises, 215, 216 types, 248 trajectory see developmental trajectory upper/lower case for gene names, trans-environment fitness profiles, 360–61 193, 352 upstream regulatory regions, 137, trans-regulation, 138, 156, 352, 353 138–9, 353 transcription urbilaterian eukaryotic, 42 bilateral symmetry, 249 factors definition, 353 binding, regulatory region mutation segmentation questions, 170–71 effects, 81 segments/eyes, gene co-option, 237, 238, definition, 352 240, 242 developmental genes, 137, 138 Urochordata (sea squirts), 115, 164, 228, 353 gene switching, 42, 43 genes, overview, 220–21 variance-covariance matrix, 210–11 Sonic Hedgehog signalling in vertebrate variation, 210–11; see also mutations embryos, 149–52 continuous variation, 86–7, 185–90, upregulation, hos2 gene single amino 260–62, 355, 356 acid mutant, 139 covariation, 206–7, 209–10, 211–13 without translation, 67 discrete variants, 254–6, 355, 356 transformations theory (Thompson), 20–21, neutral, 66, 72, 102–3, 133, 268, 289, 88–9, 353 347, 356 translocations, chromosomal, 80 standing variation vs. waiting for a transmembrane proteins, 39, 47, 48, mutation, 73 222, 225 variety of life forms, 5 transposons, 67, 79, 353 Vendobionta, 289–90, 353, 363, 364 Tree of Life (Haeckel), 294 ventralisation, 114, 224 trees see evolutionary trees; phylogenetic Veronicaceae, 244, 245 trees vertebrae, neck, 201–3 trees (plants), 9, 10 vertebrates; see also individual vertebrates trichomes, 155–8, 160 appendages, gene co-option, trilobites, 332, 333, 365 243, 244 triplet code, 81 embryonic recapitulation, 17–19, 21–2, trochophore larva, 304, 353 24, 109, 350 trunk outgrowths, 236, 242–3 endothelial cell linings, blood vessels, trunk segment modification, 141–4 145–7

99781444337204_6_index.indd781444337204_6_index.indd 440303 111/5/20101/5/2010 11:41:3911:41:39 AMAM 404 Index

vertebrates; see also individual vertebrates Waddington, C.H., 21, 23, 91, 266, 267 (cont’d) Wallace, Alfred Russel, 208–9 eye development genes, 133 Wallace effect (reproductive character fin-to-limb transition, mapping to trees, displacement), 279, 280 171–7 wasp, social, Polistes canadensis, 261–2 flatfish, left-right asymmetry, 107–9 webbed feet, 40, 173 hearts, 145–7 West Indian snail (Cerion), 268 hindlimb/forelimb length co-variation, 206–7 West-Eberhard, Mary Jane, 268 inversion of bilaterian dorso-ventral axis, Westlothiana (fossil tetrapod), 174 114–16 whales, Indohyus divergence, 6 model organisms, 35, 36 Whitehead, Alfred North, 311 origin of , 286–8 Whyte, Lancelot Law, 23, 193 proportion of animal kingdom, 329 wild-type alleles, 65, 354 rodents, teeth, 117–19 winged/wingless insect forms, 254–6 snakes, fang position, 112–13 wingless gene, 48–9, 84; see also Wnt somitogenesis, 167–9 pathway Sonic Hedgehog signalling, 48, 149–52 Wnt pathway, 45–7, 168 turtle shell, 281–4 workers, Polistes canadensis, 261–2 vestigial structures, 124–8, 353 Wright, Sewall, 190, 356 Victoria, Lake, 102 Viperidae, 112–13 Xenopus laevis, 37, 38 VNTRs (variable number of tandem repeats), 67, 79 yolk-feeding echinoderm larvae, 8, 334–5 vole (Microtus rossiaemeridionalis), 117–19 von Baerian divergence, 16–19, 313, 314, zebrafish, 171, 174, 175, 243, 360, 361 317, 318, 353 zen gene, 223, 250 Vrba, Elizabeth, 197 zinc finger, 220, 221 Vulpes vulpes (red fox), 63 zooid modules, 9, 354 vulval position in nematodes, 113–14 zygomorphic flowers, 244–5

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