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Appendix: Geological Time Scales

Appendix: Geological Time Scales

Appendix: Geological Time Scales

Three recent geological time scales for the Phanerozoic Eon. The time for beginning of each interval of geological time is given in millions of before present. Time scale divisions after Harland et al. [1], except , which follows Snelling [3].

Interval Time of Beginning

(1) (2) (3)

Cenozoic Era Quaternary Sub-era 0.01 0.01 0.01 Epoch 2.0 2.0 1.6 Tertiary Sub-era Neogene Period Epoch Piacenzian 3.4 Zanclian Stage 5.1 5.5 5.3 Epoch Messinian Stage 6.5 Tortonian Stage 11.3 10.5 Serravallian Stage 15.1 Langhian Stage 16.3 Burdigalian Stage 21.8 Aquitanian Stage 24.6 23.0 23.7 Paleogene Period Epoch Chattian Stage 32.8 27.0 30.0 Rupelian Stage 38.0 34.0 36.6 Epoch Priabonian Stage 42.0 37.0 40.0 Bartonian Stage 39.0 43.6 Lutetian Stage 50.5 45.0 52.0 Ypresian Stage 54.9 53.0 57.8 Epoch Thanetian Stage 60.2 59.0 62.3 Danian Stage 65.0 65.0 66.4 Mesozoic Era Period Maastrichtian Stage 73 72 72 Campanian Stage 83 83 83 Santonian Stage 87.5 86 86 Coniacian Stage 88.5 88 88 430 Appendix

Interval Time of Beginning

(1) (2) (3)

Turonian Stage 91 91 91 Cenomanian Stage 97.5 95 95 Albian Stage 113 107 107 Aptian Stage 119 112 114 Barremian Stage 125 114 116· Hauterivian Stage 131 119 120 Valanginian Stage 138 126 128 Berriasian Stage 144 130 135 Period Tithonian Stage 150 135 139 Kimmeridgian Stage 156 140 144 Oxfordian Stage 163 150 152 Callovian Stage 169 158 159 Bathonian Stage 175 170 170 Bajocian Stage 181 178 176 Aalenian Stage 188 181 180 Toarcian Stage 194 189 188 Pliensbachian Stage 200 195 195 Sinemurian Stage 201 201 201 Hettangian Stage 213 204 205 Period Rhaetian Stage 219 210 210 Norian Stage 225 220 220 Carnian Stage 231 229 230 Ladinian Stage 238 233 235 Anisian Stage 243 239 240 Scythian Epoch (includes Spathian, 248 245 250 Smithian, Dienerian, and Griesbachian Stages) Paleozoic Era Period Tatarian ( = Dzhulfian) Stage 253 250 255 Kazanian Stage 260 Ufimian Stage 258 258 Kungurian Stage 263 265 270 Stage 268 273 Stage 280 280 Stage 286 290 290 Carboniferous Period Sub-period Stephanian Epoch 296 300 Appendix 431

Interval Time of Beginning

(1 ) (2) (3)

Westphalian Epoch 315 310 Namurian Epoch (B+C) 320 Sub-period Namurian Epoch (A) 333 320 325 ( = Serpukhovian Epoch) Visean Epoch 352 355 Tournaisian Epoch 360 360 355 Period Famennian Stage 367 Frasnian Stage 374 375 375 Givetian Stage 380 Eifelian Stage 387 385 390 Emsian Stage 394 Siegenian Stage 401 Gedinnian Stage 408 400 405 Period Pridoli Epoch 414 Ludlow Epoch 421 420 Wenlock Epoch 428 425 Llandovery Epoch 438 418 435 Period Ashgill Epoch 448 425 440 Caradoc Epoch 458 438 455 Llandeilo Epoch 468 455 460 Llanvirn Epoch 478 470 Arenig Epoch 488 475 490 Tremadoc Epoch 505 495 510 Period Late Cambrian 525 Middle Cambrian 540 Lenian Stage Atdabanian Stage 550 Tommotian Stage 590 530 570

References (1) Harland, W. B.; Cox, A. V.; Llewellyn, P. G.; Pickton, C. A. G.; Smith, A. G.; and Walters, R. 1982. . Cambridge: Cambridge University Press. (2) Odin, G. S., ed. 1982. Numerical Dating in Stratigraphy (2 Vols.). Chichester: Wiley. (3) Snelling, N. J., ed. 1985. The Chronology of the Geological Record. Geol. Soc. Lond. Mem. 10. Oxford: Blackwell. List of Participants with Fields of Research

BABIN, C. CHARLESWORTH, B. Laboratoire de Paleontologie Dept. of Biology et Stratigraphie du Paleozoique University of Chicago Universite de Bretagne Occidentale 1103 E. 57th St. U.E.R. Sciences Chicago, IL 60637 Avenue Ie Gorgeu USA 29283 Brest Cedex Population genetics and evolu• France tionary theory Paleozoic Mollusca Bivalvia: evolution, biostratigraphy CONNOR, E. F. Dept. of Environmental Sciences BAMBACH, R.K. Clark Hall Dept. of Geological Sciences University of Virginia Virginia Polytechnic Institute Charlottesville, VA 22903 and State University USA Blacksburg, VA 24061 Population and community USA ecology, biogeography, and insect• Community paleoecology, ecospace plant interactions utilization and patterns of diversity change through time DZIK, J. Zaklad Paleobiologii PAN BANDEL, K. Aleja Zwirki i Wigury 93 Institut fUr Paliiontologie 02-089 Warsaw Universitat Erlangen-Niirnberg Poland Loewenichstrasse 28 Evolutionary paleontology 8520 Erlangen Federal Republic of Germany Paleontology, paleobiology, evolution of molluscs List of Participants with Fields of Research 433

ERBEN, H.K. FURSICH, F. T. Institut fUr PaUiontologie Institut fUr Palaontologie Universitat Bonn und Historische Geologie Nussallee 8 Universitat Munchen 5300 Bonn 1 Richard-Wagner-Strasse 10 Federal Republic of Germany 8000 M unchen 2 Phyletic extinctions (esp. "mass Federal Republic of Germany extinctions" ), general patterns of Paleoecology of mesozoic benthic organismic evolution in Phanero• marine systems zoic time FUTUYMA, D.J. FISHER, D. C. Dept. of Ecology and Evolution Museum of Paleontology State University of New York University of Michigan Stony Brook, NY 11794 Ann Arbor, MI 48109 USA USA Evolution of associations among Paleontology - functional plants and herbivorous insects, morphology, phylogenetic from genetic, ecological, and inference, taphonomy phylogenetic perspectives; coevolution FLESSA, K. W. Dept. of Geosciences GOULD, S.J. University of Arizona Museum of Comparative Zoology Tucson, AZ 85721 Harvard University USA Cambridge, MA 02138 Paleobiology, biogeography USA Evolutionary biology FLU GEL, E. Institut fUr Palaontologie HALLAM, A. Universitat Erlangen - Nurnberg Dept. of Geological Sciences Loewenichstrasse 28 University of Birmingham 8520 Erlangen P.O. Box 363 Federal Republic of Germany Birmingham B15 2TT Evolution offossil reefs England Evolutionary paleobiology, paleobiogeography 434 List of Participants with Fields of Research

HSU, K.J. LaBARBERA, M. Geologisches Institut Dept. of Anatomy ETH-Zentrum University of Chicago 8092 Zurich 1025 East 57th Street Switzerland Chicago, IL 60637 Sedimentology, geochemistry, USA paleoenvironmental analyses Functional morphology and biomechanics of living and fossil HUSSNER, H. M. marine invertebrates Institut fUr PaHiontologie LEVINTON, J. S. Universitat Erlangen - Nurnberg Dept. of Ecology and Evolution Loewenichstrasse 28 State University of New York 8520 Erlangen Stony Brook, NY 11794 Federal Republic of Germany USA Carbonate sedimentology, Marine ecology, evolutionary evolution of reefs, mass extinctions biology

JABLONSKI, D. MEETER, D. A. Dept. of Geophysical Sciences Dept. of Statistics University of Chicago Florida State University 5734 S. Ellis Avenue Tallahassee, FL 32306 Chicago, IL 60637 USA USA Ecological statistics: time analysis, design of experiments and Paleobiology and macroevolution tests of hypotheses JARVINEN, O. MOSBRUGGER, V. Dept. of Zoology Institut fUr Palaontologie University of Helsinki Universitat Bonn P. Rautatiekatu 13 Nussallee 8 00100 Helsinki 10 5300 Bonn 1 Finland Federal Republic of Germany Community ecology of birds; Paleobotany, evolutionary biology ecological zoogeography; population biology of endangered populations List of Participants with Fields of Research 435

MULLER, G. RAUP, D.M. Institut fUr Anatomie Dept. of Geophysical Sciences Wahringerstrasse 13 University of Chicago 1090 Vienna 5734 S. Ellis Avenue Austria Chicago, IL 60637 Experimental embryology, USA vertebrate limb morphogenesis, Theoretical paleobiology and evolutionary biology evolutionary biology REIF, W.-E. NAGL, W. Institut und Museum fUr Geologie FB Biologie und Palaontologie Universitat Kaiserslautern Universitat Tiibingen Postfach 3049 Sigwartstrasse 10 6750 Kaiserslautern 7400 Tiibingen Federal Republic of Germany Federal Republic of Germany Genome organization and Evolutionary biology, functional evolution ofspecies, DNA evolution morphology in cell cultures, cell differentiation and malignant transformation de RICQLES, A. J. Laboratoire d' Anatomie NIKLAS, K. J. Comparee Section of Plant Biology Universite Paris VII Cornell University 2, Place Jussieu Ithaca, NY 14850 75005 Paris USA France Plant development and evolution Evolutionary biology, paleohistology PANCHEN, A.L. Dept. of Zoology RIEGER, R. M. The University Institut fUr Zoologie Newcastle upon Tyne NE1 7RU Universitat Innsbruck England Universitatsstrasse 4 6020 Innsbruck Vertebrate paleontology: origin Austria of land vertebrates and early The origin and radiation of the tetrapods; taxonomic theory bilateria as deduced from neonological information (with special reference to the new ultrastructural information) 436 List of Participants with Fields of Research

RUNNEGAR, B. SOULE, M.E. Dept. of Geology and Geophysics 118 Little Oaks Rd. University of New England Encinitas, CA 92024 Armidale, NSW 2351 USA Australia Evolutionary genetics and Evolution of the mollusca, phenetics; conservation biology; crystallography of carbonate environmental ethics biominerals, protein evolution SOUSA, W.P. SEILACHER, A. Dept. of Zoology Institut und Museum fUr University of California Geologie und Paliiontologie Berkeley, CA 94720 Universitat Tiibingen USA Sigwartstrasse 10 7400 Tiibingen 1 Disturbance and successional Federal Republic of Germany dynamics in marine intertidal Paleobiology communities; ecology of host• parasite interactions SELANDER, R. K. Dept. of Biology STEARNS, S. C. University of Rochester Zoologisches Institut River Campus Rheinsprung 9 Rochester, NY 14627 4051 Basel USA Switzerland Molecular population genetics Evolutionary theory, evolutionary SEPKOSKI, Jr., J. J. ecology, life-history evolution Dept. of Geophysical Sciences University of Chicago STINNESBECK, W. 5734 S. Ellis Avenue Institut fUr Palaontologie Chicago, IL 60637 Universitat Bonn USA Nussallee 8 Paleontology, paleobiology 5300 Bonn 1 Federal Republic of Germany SIMBERLOFF, D. Cretaceous-tertiary boundary Dept. of Biological Science events, upper cretaceous mol/uses, Florida State University paleoenvironmental analyses Tallahassee, FL 32306 USA Ecology, evolution, biogeography List of Participants with Fields of Research 437

TURNER, J.R.G. VERMEIJ, G.J. Dept. of Genetics Dept. of Zoology University of Leeds University of Maryland Leeds LS2 9JT College Park, MD 20742 England USA Evolutionary genetics Paleobiology, malacology

UNDERWOOD, A. J. WAGNER, G.P. Dept. of Zoology Institut fUr Zoologie School of Biological Sciences der Universitat Wien University of Sydney Althanstrasse 14 Sydney NSW 2006 1090 Vienna Australia Austria Experimental intertidal ecology; Quantitative genetic theory of epistemology of ecological phenotypic evolution, evolution of experiments development WAKE, D.B. URBANEK, A. J. Museum of Vertebrate Zoology Zaklad Paleobiologii PAN 2593 Life Sciences Building Zwirki i Wigury 93 University of California 02-089 Warsaw Berkeley, CA 94720 Poland USA Paleontology and evolutionary Evolutionary biology, with biology, morphogenesis and evolu• emphasis on patterns and processes tion of colonial organisms of evolution in urodeles; functional (graptolites, pterobranchs) and evolutionary morphology, genetics in relation to speciation VALENTINE, J. W. and geographical ecology Dept. of Geological Sciences University of California WEIDICH, K.F. Santa Barbara, CA 93106 Institut fUr Palaontologie USA und Historische Geologie Origin of phyla Universitat Miinchen Richard-Wagner-Str. 10/11 8000 M iinchen 2 Federal Republic of Germany Mesozoic microfaunas; cretaceous foraminifera Author Index

Babin, C. 331-350 Milller, G. 167-182 Bambach, R. K. 331-350, 407-428 Nagl, W. 167-182, 223-232 Bandel, K. 167-182 Niklas, K. J. 331-350, 383-405 Charlesworth, B. 167-182 Panchen, A. L. 331-350 Connor, E. F. 47-67, 119-147 Raup, D. M. 1-5, 7-22, 235-257 de Ricqles, A. J. 47-67 Reif, W.-E. 47-67 Dzik, J. 47-67 Rieger, R. M. 47-67 Erben, H. K. 235-257 Runnegar, B. 167-182 Fisher, D. C. 47-67,99-117 Seilacher, A. 47-67 Flessa, K. W. 235-257 Selander, R. K. 167-182 Flugel, E. 331-350 Sepkoski, J. J., Jr. 235-257, 277-295 Fursich, F. T. 331-350 Simberloff, D. 259-276,331-350 Futuyma, D.J. 331-350,369-381 Soule, M. E. 235-257 Gould, S. J. 7-22, 47-67 Sousa, W. 235-257 Hallam, A. 235-257 Stearns, S. C. 23-44, 167-182 Hsu, K. J. 235-257, 297-312 Stinnesbeck, W. 235-257 Hussner, H. M. 235-257 Turner, J. R. G. 167-182, 183-207 Jablonski, D. 1-5, 7-22, 235-257, Underwood, A. J. 331-350, 351-367 313-329 Urbanek, A.J. 167-182 Jarvinen, O. 331-350 Valentine, J. W. 167-182, 209-222 LaBarbera, M. 47-67,69-98 Vermeij, G. J. 235-257 Levinton, J. S. 167-182 Wagner, G. P. 47-67, 149-165 Meeter, D. A. 47-67 Wake, D. B. 47-67 Mosbrugger, V. 47-67 Weidich, K. F. 331-350 Subject Index

Aclonal growth form, clonal vs. 344 Astogeny 49 Actuotaphonomy 347 Atavisms, experimentally induced 156 Adaptation(s) 11,23-44,108,112, 113 Autocorrelation function 121 Adaptive landscape 216, 325 Autocovariance function 121 - peak 216 Autonomization 49 - radiation(s) 87, 183-206, 314, 326, 339, 345, 348 Background and mass extinction - replacement 99, 101, 104 processes 324 - selection 223 - and mass extinction regimes 246, Allometric coefficient 72 323 - equation 71 - conditions 318 - models 73-78 - extinction(s) 235-239, 246, 248, 278, Allometry 70-79,87-89 314, 322, 372 -, evolutionary 70, 87-89 -- rates 247 -, interspecific 70, 79 - levels of extinction 16 -, intraspecific 70, 79 Backward smearing 241,242 -, ontogenetic 70, 88 Baupliine 33, 167-169, 209-221 Alpha diversity 360 Bellerophontid mollusks 324 Ammonite(s) 10,321,323 Benthic communities 214,372 Ammonoids 285 Beta diversity 360 Amniotes 394, 400 Biogeography 19, 20, 319, 332, 335, Amniotic egg 400 336, 338, 339, 341 Amphibians 394 - of extinction 319 Analogies 151 Biological hierarchy 28, 29 Angiosperms 339, 345, 401 Biology, comparative 39 Animal and plant communities - of constraints 156-161 383-403 Biomass 388, 398 Annelids 168 Biomere 279 Anoxia 252, 254 Biotic change 409 -, oceanic 252, 254 Bioturbation 325,419 Anoxic events 252 Birds 383, 394, 398 Antelopes 14 Bivalve mollusks 56,319,323,325,326 Antipredatory morphology 324 Body plan 33 Aptian 286 - size 16, 214, 246-248 Archaeopteryx 398 Bolide, extraterrestrial 249 Arthropods 394 Bootstrap techniques 138 Articulate brachiopods 323, 325, 326 Bose-Einstein condensation 227-228 Assemblage 332-334, 346, 352 Boundary, Cretaceous/Tertiary 243, -, community vs. 332-334, 346 249-253, 304, 307, 309 Assemblages, changes in species -, Permo/Triassic 304, 310 341-348 -, -Cambrian 209, 210, -, species 333, 338, 341-348 304, 308, 310 440 Subject Index

Brachiopods 11,319,323,325,326 -, Benthic 214, 372 -, articulate 323, 325, 326 -, climax 353 Brain size 89 -, coevolution in 369-379 - -, evolution of 14 - convergence 343-344 Branching process 100, 105 -, displacement 414-416 Bryozoans 320, 323 - evolution 17, 18, 331-350, 384, Burgess shale 10, 211 409-411,421, 422 Butterflies 178 -, plant 383-403 -, reef 319, 320, 325 Calcite dissolution 310 - structure 411,419, 420, 422, 423 Cambrian 3,288 -, tropical reef 320 -, Metazoan radiation at the beginning - vs. assemblage 332-334, 346 of the 3 Comparative biology 39 Carbon isotope fractionation 300 - method 162 Carnivora, order 324 Competition 17,101,104,111, Carnivores 393, 396 332-334, 345 Causality 102, 103, 105, 110-112 Competitive exclusion 325 Causes of extinction 259-274 - interactions 2, 377 Cenomanian 252, 287 Competitors, predators, prey 18 - /Turonian extinction(s) 252 Completeness of record 215 Change in sea level 250-254 Complexity 52, 54 Changes in species assemblages Conformation, DNA 225, 226 341-348 Congruence 52 Character displacement 81 Constraint(s) 23-44, 55, 107, 112, 114, Chronostratigraphic resolution 241 153-162, 225 Clade(s) 54, 101, 106-108, 114, 247, -, archaic 157 248 -, biology of 156-161 -, species-richness within 247, 248 -, demographic 38 Cladistics 189 -, developmental 37, 154, 156, 157, Cladogenesis 53, 194 159,170,172 Cladograms 191 -, functional 156 Classification of organisms 225 -, genetic 36 Climatic change 252 -, physiological 37 Climatic cooling 252, 254 -, selective 36 Climax communities 353 -, systemic 36 Clonal groups 320 Constructional morphology 11 - reproduction 10 Continental drift 336, 340 - vs. aclonal, growth form 344 Continuity 314 Close mimicry 191 Convergence 49 Cnidarians 168 -, community 343-344 Coevolution 338-339,347,348, Cooling, climatic 252, 254 369-379,420,422 Cope's Rule 53, 83, 84, 89, 107, 246 - in communities 369-379 Corals 320, 323 -, plant/insect 377 -, rugose and tabulate 323 Collective properties, emergent vs. 337, Corridor model 158 338 Covariance matrices 37 Community 17, 18, 214, 319, 325, Cretaceous, end of the 326 351-365, 369-379, 383-403, 407-423 - extinctions, terminal 249, 253 -, animal and plant 383-403 - mass extinctions, terminal 277 Subject Index 441

- jTertiary 243, 24~248 Drosophila 171, 173 -j- boundary 243, 249-253, 304, 307, Dwarfing 84,85 309 -j- extinctions 255 Crinoids 323, 326 E. coli 172 -, stalked 326 Early Metazoan speciation 168, 169 Criterion of coincidence 152, 154 Ecological succession 336 -- structure and position 151 Ecomorphology 343,347-348 Cryptic species 15 Ecophenotypic 12 Ecospace 409,417,423 Ecosystem 354 Demographic constraints 38 Ecotones 354 Determinism 223, 224 Ectotherms 397 -, physical 223 fauna 211 Development and evolution 167, 170, Effect hypothesis 14 172 Effects of mass extinction 254, 255 Developmental constraints 37, 154, Elaboration 57 156, 157, 159, 170, 172 Elastic similarity 75-79 Devonian mass extinctions 290 Emergence 103, 104, 113 Diagenetic changes 333, 335, 347 Emergent properties 361-363,422 Differential reproductive success 25 - vs. collective properties 337, 338 Differentiation 49 Endothermy 383 Diffusion 100, 107, 108, 112, 114 Eocene 287 Dinosaurs 18, 324 Environmental gradients 354,410--412, Directional selection 13, 171 422 - speciation 14 Epeiric seas 251 Directionality 48, 100, 101, 105, 108, Epifaunallife 214 161 Epiphenomenon 421 Directions 52 Equation, allometric 71 Dispersal 16, 31, 246 Equilibrium, punctuated 175-179, 192, - ability 16, 246 200,377 Displacement 422, 423 Escalation 61 Dissolution, calcite 310 Eukaryotes 218 Distribution(s), geographic 248 Evolution, community 17,18,331-350, -, species-abundance 334, 335, 347 384 Diversification 314, 324-326 -, development and 167,170,172 -, post-event 324-326 -, hierarchical view of 319 Diversity 49, 277, 284, 340--341, 347, -, iterative 10 360,372,379,388,409,417,420,423 -, modifier 160 -, alpha 360 - of brain size 14 -, beta 360 -- reptiles 395 -, species 83, 214, 360, 372 -- sex 31 DNA conformation 225, 226 -, parallel 49 -, excited states of 227 -, phyletic 175-177, 191, 202 Drift 9, 178, 179, 216, 263, 336, 340 -, physical aspects of 224 -, continental 336, 340 -, punctuated phyletic 191 -, genetic 216, 263 -, punctuational 183-206 -, species 178, 179 -, rate of 157 Driving forces 227 -, sequence 168, 169, 174 442 Subject Index

Evolutionary allometry 70, 87-89 - rate 247, 346 - dynamics underlying extinction events - rates, background 247 322 - regime, background and mass 323 - novelties 64 -, selectivity of 245-248 - significance of mass extinctions 3 -, Toarcian 252 - trends 12, 14, 199 Extraterrestrial bolide 249 - -, large-scale 12 - objects, impact of 249, 250 - window(s) 158, 159 Excimers 227 Fauna, Ediacaran 211 Excited states of DNA 227 -, insect 383, 393 Exclusion, competitive 325 Faunal replacement 325 Expansion of oxygen-minimum zone Feedback selection 160, 161 304, 305 Fertility perturbation 300 Experimentally induced atavisms 156 Fitness 23-44, toO, 109-112,264 Experimentation 49 Fixation of traits 28 External vs. internal factors 224 Food chain 370 Extinction(s) 3,15,16,86,87,127,185, - webs 363 197, 235-241, 245-253, 255, 259-274, Foraging 377 278,285,288,314,316,318-325,336, Foraminifera 10, 335 337, 342, 369, 372, 379 Fossil assemblages 417 Extinction(s), background 235-239, - record 3, 5, 63, 209-221, 277, 369, 246, 248, 278, 314, 322, 372 383-389, 393-398 -, biogeography of 319 -- completeness 213 - cascading extinction waves 336, 337 - -, hypothesis testing 3 -, causes of 259-274 -- of terrestrial plants 383, 385-389 -, Cenomanian-Turonian 252 --- terrestrial plants and animals -, contemporary 259 383 -, Cretaceous-Tertiary 236,249,253, --- vertebrates 393-398 255, 321, 323 Fossilization 19 - during historic time 239-241 Fossils, reworking of 241 -, Ordovician 236, 318, 325 Fourier analysis 131 -, Permian-Triassic 3,216,252,253, Fractionation, carbon isotope 300 284, 316, 318 Frasnian-Famennian extinction 252 -, - /Triassic-Jurassic 285,318 Function 52, 121 - event(s) 235-239, 253, 322 -, autocorrelation 121 - -, evolutionary dynamics underlying -, autocovariance 121 322 Functional constraints 156 - -, mass 322 - morphology 333, 335, 347 -, Frasnian-Famennian 252 Functionalism 224 - intensities 235, 243, 248 -, late Devonian 236, 252, 318 Gastropods 14, 178, 319, 323 -, levels of background 16 -, volutid 14 -, mass (see Mass extinction) Gene transfer, lateral 172 -, mechanisms of 249 Generalists 403 - of species 369 Generalized species 340, 348 -, Paleozoic 288 Genes, regulatory 40, 169, 173, 174 -, probability of 239, 245, 246, 248 -, structural 173 - processes, background and mass Genetic constraints 36 extinction 324 - deterioration 263 Subject Index 443

- drift 216, 263 Impact hypothesis 322 - mechanisms 9 - of extraterrestrial objects 249, 250 - population structures 9 Improvement 99-103, 108, 109, - recombination 10 111-115 - systems 9, 13 Inbreeding 263, 264 - theory, population 2 Increase of information and entropy Geoflora 336, 346 224 Geographic distribution 248 Individualistic vs. holistic view 336, - range 245-247, 319 337, 346 Geographical isolation 203 Industrial melanism 189 Geologic time scale, quality of 242 Innovation 57 Geometric similarity 74, 76, 77 Insect(s) 18, 375-378, 383, 389, 392, Goldschmidtian macromutations 203 393 Gradualistic change 13, 229 - fauna 383, 393 Graptolites 177, 320 - jplant coevolution 377 Great American Interchange 345 -, pollinating 378 Growth form, clonal vs. aclonal 344 Intensities, extinction 235, 243, 248 Gymnosperms 401 Interactions, competitive 2, 377 Guild structure 335, 340-343, 409 Intermediate structure 33, 38-40 Guinea pig 171 Internal vs. external factors 224 Interspecific allometry 70, 79 Habitat choice and stasis 172 - variation 152 Hard substrates 412,416 Intraspecific allometry 70, 79 Heliconius 186, 192 Intrinsic rate of increase (rm) 80, 89 Herbivores 394, 396 Iridium 249,254 Herbivorous reptiles 396 Irreversibility 49 Herbivory 375, 392 Isotope fractionation, carbon 300 Hereditary information, horizontal Iterative evolution 10 transfer of 10 Iteroparity 31 Heritability 25, 73, 89 Heterozygosity 86 Keystone species 336 Hierarchical levels 246, 247 - view of evolution 319 Lagerstatte(n) 10, 11, 18, 335, 347 Hierarchy 28,29,48 Large-scale evolutionary trends 12 -, biological 28, 29 Larval development, planktotrophic Higher taxon 154 325, 326 History 33 Lateral gene transfer 172 Hitchhiking 178, 179 Lazarus effect 242, 316 Holistic vs. individualistic view 336, Legumes 173 337, 346 Linkage 37 Home range 82 Homologue 151, 153 Macroevolution 24,32,41, 154, 155, Homology 150-154 162, 204, 378 Homo sapiens 239,240 Macroevolutionary role of speciation 2 Horizontal transfer of hereditary Macromutations, Goldschmidtian 203 information 10 Maestrichian 287 Hybridization 194, 271 Magnetic reversals 253 Hypermorphosis 14 Major adaptations 19 Hypothesis testing - fossil record 3 Major axis 73 444 Subject Index

Major innovations 18 Method, comparative 162 Mammal-like reptiles () 18, Microevolution 24, 32,41, 55 324 Microplankton 12 Mammals 319,324,326,378,383,394, Miocene, middle 287 397 Model(s), allometric 73-78 -, placental 319, 326 -, corridor 158 Marine communities 407-423 -, Markovian 99, 101-105, 112-114 - -, structure of 416-420 -, probabilistic 51 Marine regressions 16 - I regressions 72 - reptiles 396 - II regressions 73 - revolution 323, 324 Mimicry 191, 196, 197 Markov processes 51 -, close 191 Markovian models 99, 101-105, Minimum viable populations 263 112-114 Modem Synthesis 185,200 Marsupial 319 Modes of life 409, 419, 420 Mass extinction(s) 3, 16, 111,235-239, Modifier evolution 160 243, 246-255, 277-292, 315, 318, 319, Molecular drive 229 322-324, 344, 345, 348 Mollusks 11, 56, 168, 324 -- are a potent macroevolutionary -, bellerophontid 324 force 318 -, bivalve 56 - -, Cretaceous-Tertiary (end-Cretaceous) Monophyletic groups 54 277, 319 Morphological integration 156 - -, definition of 315 - stasis 2 - -, Devonian 290 Morphology 11,150,324,333,335,347 - -, effects of 254, 255 -, antipredatory 324 -- events 322 -, constructional 11 - -, evolutionary significance of 3 -, functional 333, 335, 347 - -, Ordovician 289,319 Morphometrics 344,347,348 - -, patterns of differential survivorship Morphospace 10 associated with 319 Morphospecies 15 -- processes, background and 324 Mutation, random 223 -- regimes, background and 323 -, somatic 9 - -, survivorship in 319, 344, 348 - -, timing of 243 Natural experiment 50 Matrices, covariance 37 - selection 23-44, 99, 108-114 Mechanical similarity 77, 78 - system of organisms 150 Mechanisms of extinction 249 Nematoda 173 Mendelian 9 Neo-Darwinian paradigm 101, 108, Mesozoic marine revolution, precursor 114 to 324 -jDarwinism 199,202 - revolution 323, 324 Neotaphonomy 347 - time scale 242 Neoteny 14 Metabolic rate 78 New synthesis 228 Metamerism 54 Niche partitioning 403,416 Metazoa 210 - structure 417 Metazoan speciation, early 168, 169 NO. 308 - radiation(s) 3, 209, 219 Nonadaptive processes 205 -- at the beginning ofthe Cambrian 3 Non-amniote tetrapods 395 Meteorite impact(s) 239, 311 Nonrandomness 224 Subject Index 445

Nuclear winter 253, 310 Phyletic evolution 175-177,191,202 -- scenarios 250 - -, punctuated 191 Nutrients 421 - transformation 12, 217 Phylogenetic relationships 345 Ocean pollution 302 Physical aspects of evolution 224 Oceanic anoxia 252, 254 - determinism 223 Ontogenetic allometry 70, 88 Physiological constraints 37 Ontogeny 49 Placental mammals 319, 326 Opportunistic species 254 Planktonic foraminifera 10 Optimality theory 35 Planktotrophic larval development Ordovician, lower 288 325,326 - mass extinction 289 Plant adaptations 375 Organismal integration 39 - communities 383--403 Organisms, classification of 225 - grades 335, 341 Organization 332-335 - jinsect coevolution 377 Organizational complexity 49 -, vascular 320 Origin of a homologue 155 Platyhelminths 168 Originations 127 Pleistocene 252 Orthogenesis 8 -, late 321 Oxygen concentration 420 Pliensbachian 286 - crisis 308 Pliocene 287 - deficiency 305-308 Pollinating insects 378 Oxygen-minimum zone, expansion of Polycohort survivorship 237 304, 305 Population density 82 - genetic theory 2 - structures, genetic 9 Paleocommunities 333, 408, 417 Positional information 161 Paleontologic resolution 241 Post-event diversification 324--326 Paleontological samples 9 Precambrian 370 Paleozoic extinction 288 -jCambrian boundary 209,210,304, Parallel evolution 49 308, 310 Parasites 378 Precursor to the Mesozoic marine Parasponia 173 revolution 324 Patterns 47 Predation 325, 378,418, 422 - of differential survivorship associated Predators, prey, competitors 18 with mass extinction 319 Primary producers 370 Periodicity 119,120,131,135,138,145, Principal components analysis 71 243-245 Probabilistic models 51 Permian extinction, late 236, 253, 284, Probability of extinction 239, 245, 246, 316 248 - mass extinction, late 284 Process(es) 51, 55, 100, 105, 205, 324 Permo-Carboniferous 252 -, background and mass extinction 324 - jTriassic 252, 304, 308, 310 -, branching 100, 105 -j- boundary 304,310 -, Markov 51 -j - extinction 252 -, nonadaptive 205 Persistence of trends 323, 324 -, stochastic 51 Perturbation, fertility 300 Productivity 247 Phanerozoic terrestrial biota 372 Progenesis 14 Photon emission 227 Progress 99, 100, 102-104, 111 446 Subject Index

Prokaryotes 218 Reworking of fossils 241 Provinciality 20, 214, 252-254 Risk minimization 29 Pseudo-genes 174 - /minimizing traits 42 Pteridophytes 346 Rugose and tabulate corals 323 Pull of the recent 15,128 (rm) 86,89 Punctuated equilibrium 175-179,192, 200, 229, 377 Salmonella 172 - phyletic evolution 191 Scale 53 Punctuational evolution 183-206 Scaling coefficient 74 - pattern 13, 217 - exponent 74 Sea level 250-254 Quality of the geologic time scale 242 - -,change in 250-254 Quantitative genetics 9 Sedimentary completeness 213 Seed 400 Racheting 57 Selection 2, 9, 13, 14, 23--44, 55, 99, Radiation(s), adaptive 87, 183-206, 108-114, 160, 161, 171, 178, 179, 204, 314, 326, 339, 345, 348 217, 223 - at the beginning of the Cambrian, -, adaptive 223 Metazoan 3 -, directional 13, 171 -, Cambrian 288 -, feedback 160, 161 -, Metazoan 3, 209, 219 -, natural 23--44, 99, 108-114 Random changes 49,50 -, species 2, 14, 27, 55, 178, 179, 204, - mutation 223 217 - walk 100, 105, 106 -, stabilizing 13, 160, 171 Randomness 50 Selective constraints 36 Rate of evolution 157 Selectivity 245-248, 254, 314, 317-322 -- extinction 346 - of extinction 245-248 Red Queen Hypothesis 16,269 Self-organization 224 Reduced major axis regressions 73 Sequence evolution 168, 169, 174 Reducibility 104 Sex, evolution of 31 Reductionist explanation 34 - ratio 263, 268 Reef(s) 3, 337, 343, 344, 412--414, 416, Sibling species 15, 200, 202 419 Simulation 51, 52 - communities 319, 320, 325 Size 49, 70, 321 - -, tropical 320 -, body 16, 214, 246-248 Refugia 255 Smearing, backward 241,242 Regulatory genes 40,169,173,174 Soft substrates 410--412 - mechanisms 226 Somatic mutations 9 Repetition 52 Spatial structure 335, 346 Replacement 49, 255 Specialization 49 -, adaptive 99, 101, 104 Specialized species 340, 348 -, faunal 325 Speciation 2, 11-14, 19, 168, 169, Reproductive success, differential 25 176-180,185,217,322 Reptiles 18, 324, 394-396 -, directi onal 14 -, evolution of 395 -, early Metazoan 168, 169 -, herbivorous 396 -, macroevolutionary role of 2 -, mammal-like 18, 324 - rates 322 -, marine 396 Species-abundance distributions 334, -, 18, 324 335, 347 Subject Index 447

- assemblages 333, 338, 341-348 Tabulate and rugose corals 323 - -,changes in 341-348 Taphocoenosis 333, 347 -, cryptic 15 Taphonomy 8, 12, 417 - diversity 83, 214, 360, 372 - feedback 417 - drift 178, 179 Taxocenes 333, 352 -, extinction of 369 Temporal heterogeneity 334, 346 -- family ratios 316 - scales 334 -, generalized 340, 348 Terrestrial arthropods 389-393 - interactions 332-339, 347, 348 - plants, fossil record of 383, 385-389 -, keystone 336 -- and animals, fossil record of 383 -, opportunistic 254 Tetrapods 383, 394, 395 - packing 3 -, non-amniote 395 - richness 247, 248, 388 Thanatocoenosis 333, 341 -- within clades 247, 248 Therapsid reptiles 18, 324 - selection 2, 14,27, 55, 178, 179,204, Thermodynamic code 226 217 Thermodynamics 230 Species, sibling 15, 200, 202 Tiering 419 -, specialized 340, 348 Time-averaging 9, 12, 17, 19, 334, 346, Stabilizing selection 13, 160, 171 408, 410, 416, 417 Standing crop 408 - resolution 12, 19 Stasis 2, 13, 18, 28, 159, 168, 169, 171, - series 119-145 172, 175, 196, 339 -- analysis 119-145 -, habitat choice and 172 Timing of mass extinctions 243 Static stress similarity 75 Tithonian, upper 286 Statistics 49 Toarcian extinction 252 Sticklebacks 176 Transitivity 103, 104, 113 Stochastic processes 51 Transposable elements 173 Strangelove Ocean 298-304, 306 Trends 14, 17, 48, 99-104, 107, 108, - perturbation 300, 304, 310 112-115,199 Stratigraphic ranges 242 -, evolutionary 14, 199 - resolution 241,315 Triassic extinction(s), late 236,252,285 Structural genes 173 Trilobites 10, 325 Structuralism 224 Trophic levels 370 Structure 332-335 - structure(s) 335, 363, 417, 418 -, guild 335, 340--343 Tropical floras 319 - of marine communities 416-420 - forest 3 Succession 17, 336, 416, 417 - reef community 320 -, ecological 336 - setting 247 Superorganism concept 336-338 Unterbauplane 209, 212, 219 Survivorship in mass extinction 319, 344,348 Variability 25 -, polycohort 237 Vascular plants 320 Symmigy 333 Vectors 52 Synergetics 225 Vertebrates 168 Synorganization 49 -, fossil record of 393-398 Systemic constraints 36 Vicariance biogeography 338 Systems approach 149 Volatility 321 - conditions 225 Volutid gastropods 14

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