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Ecology, genetics and evolution of metapopulations i hanski, o gaggiotti (elsevier, 2004)

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CONTRIBUTORS Numbers in parentheses indicate the pages on which the authors" contributions begin Jon Aars (515) NERC Molecular Genetics in Ecology Initiative, Aberdeen Population Ecology Research Unit, School of Biological Sciences, University of Aberdeen, Aberdeen AB24 2TZ, Scotland; current address: Norwegian Polar Institute, Polarmiljosenteret, N-9296 Tromso, Norway Janis Antonovics (471) Biology Department, University of Virginia, Charlottesville, Virginia 22904 Ottar N Bjornstad (415) Departments of Entomology and Biology, Penn State University, University Park, Pennsylvania 16802 Benjamin M Bolker (45) Zoology Department, University of Florida, Gainesville, Florida 32611 Cajo J.F ter Braak (105) Biometrics, Wageningen University and Research Centre, Box 100, NL-6700 AC Wageningen, The Netherlands Mar Cabeza (541) Metapopulation Research Group, Department of Ecology and Systematics, University of Helsinki, FIN-00014 Helsinki, Finland Jean Clobert (307) Laboratoire d'Ecologie, Universit~ Pierre et Marie Curie, B~timent A, 75252 Parix cedex 05, France O Eriksson (447) Department of Botany, Stockholm University, SE-106 91 Stockholm, Sweden Rampal S Etienne (105) Community and Conservation Ecology Group, University of Groningen, Box 14, NL-9750 AA Haren, The Netherlands Oscar E Gaggiotti (3, 337, 367) G~nomique de Populations et Biodiversit~, LECA-CNRS UMR 5553 Universit~ Joseph Fourier, F-38041 Grenoble Cedex 9, France Sergey Gavrilets (275) Department of Ecology and Evolutionary Biology, Department of Mathematics, University of Tennessee, Knoxville, Tennessee 37996 ix 682 REFERENCES Wu, C.-I (2001) The genic view of the process of speciation J Evol Biol 14, 851-865 Wu, C.-I., and Palopoli, M E (1994) Genetics of postmating reproductive isolation in animals Annu Rev Genet 27, 283-308 Yodzis, P (1978) Competition for space and the structure of ecological communities In "Lecture Notes in Biomathematics," Vol 25 Springer-Verlag, Berlin Young, A., Boyle, T., and Brown, T (1996) The population genetic consequences of habitat fragmentation for plants Trends Ecol Evol 11, 413-418 Yu, D.W., Terborgh, J.W., and Potts, M.D (1998) Can high tree species richness be explained by Hubbell's model? Ecol Lett 1, 193-199 Yu, D.W., and Wilson, H.B (2001) The competition-colonization trade-off is dead; long live the competition-colonization trade-off Am Nat 158, 49-63 Yu D.W, Wilson, H.B., and Pierce, N.E (2001) An empirical model of species coexistence in a spatially structured environment Ecology 82, 1761-1771 Zabel, J., and Tscharntke, T (1998) Does fragmentation of Urtica habitats affect phytophagous and predatory insects differentially? Oecologia 116, 419-425 Zera, A.J., and Denno, R.E (1997) Physiology and ecology of dispersal polymorphism in insects Annu Rev Ecol Syst 42, 207-231 Zeyl, C., Mizesko, M., and de Visser, J (2001) Mutational meltdown in laboratory yeast populations Evolution 55, 909-917 Zhang D.Y., and Lin, K (1997) The effects of competitive asymmetry on the rate of competitive displacement: How robust is Hubbell's community drift model? J Theor Biol 188, 361-367 INDEX effects, 200 variance, 160 alternative stable equilibria, 12 strategies, 141 American mink (Mustela vison), 520, 522 population size and extinction risk, 527 American pika (Ochotona princeps), 100, 132, 503, 517 ancestral process, 175 ancient lineages in a metapopulation, 188 androdioecy, 245 anther smut, 473-4 anthropogenic threats, 566 assignment test, 371 assortative mating coefficient, 377, 382 A abundance distribution, 136 adaptation, 18, 223 in subdivided populations, 165 to local conditions, 295 to marginal habitats, 409 to sink habitats, 406, 409-10 adaptive landscapes, 18, 280 plasticity, 263 additive genetic effects, 264 genetic variance, 203, 205-6, 208, 213, 264, 348-9 age at maturity, 246 at reproduction, 246 age-structure, 252, 399 age-structured population, 102 aggregation, 62, 64 Aland Islands, 74, 492, 500-1,506 Allee effect, 13, 87, 90, 346, 458 alleles average effects of, 208 allelic diversity, 349 B badger, 28 Bateson-Dobzhansky-Muller model, 279, 296 Bay checkerspot butterfly, see Euphydryas editha Bayesian approach, 184 approach, hierarchical, 378, 383, 386 approach, potential problems, 384 683 684 Bayesian (Continued) inference, 374 model choice, 380 BDM model, 279, 296 multilocus generalization of, 281 bet-hedging strategy, 243 Bicyclus anynana, 359 biodiversity, 541 black hole sink habitat, 158 adaptation to, 405 black-tailed prairie dog (Cynomys ludovicianus), 517, 519 boundary length minimization problem, 556-7 breeder's equation, 203 breeding dispersal, 528 phenology, 411 value, 201,208 bubonic plague, 435, 438-40 Burramys parvus, 532 C Carlina vulgaris, 246 carrying capacity, 389 Centaurea jacea, 459 chaotic fluctuations, 346 childhood diseases, 428 chinook salmon metapopulation, 567 clade disparity, 289 Clarkia concinna, 458 classic metapopulation dynamics, 506 model, theory, 12, 83 small mammal, 517 climate change, 507-9 coalescence approach, 17, 368 in metapopulations, 186 in panmictic populations, 175 in two-sex diploid populations, 179 times, 176 coalescent inference analytical methods, 182 computational method, 183 in metapopulations, 196 interval, 176 standard model, 177 coalescent predictions in metapopulations, 194 coalescent process, 175, 178 at a multisite genetic locus, 178 INDEX making inferences with the, 182 simulations, 193 standard, 175-6, 192 theory, 174 coevolution in a metacommunity, 270 of dispersal and habitat specialization, 326 of phenotype and context, 259-60 coevolving species, 270 coexistence, 146-7, 149 of multiple species, 134 collecting phase, 189-93 colonization, 85, 87, 247, 367, 370, 385, 505 dataset, 116 density-dependent effects on, 376 distance effects on, 464 events, 382 evolutionary effects of, 275 rate, 92, 130 related to connectivity, 115 colonization and extinction processes, 82 rates, 78 colonization-competition trade-off, 146, 149 colonizer syndrome, 238, 251 colonizing groups, composition of, 378 common lizard, 313, 316, 318, 323-5 community heritability, 271 estimate of, 272 competition, 251 competitive ability, 148, 272 interactions as forms of IGEs, 269 complementarity, 543 connectivity, 25, 100, 102, 108, 114, 527, 550, 552, 563 asymmetrical, 28 in a metacommunity, 138 measure of, 111,498 of networks, 501 conservation, 8, 511 biology, 20, 338, 541 conspecific attraction, 111,315, 519, 529, 531,534 context evolving, 263 kinds of, 262 variance in, 262 continuum models, 50 core-satellite distributions, 90 correlated landscape, 424 random walk, 13 INDEX corridors, 144 coupled-map lattice, 48, 100 critical community size, 423, 429 CSI method, 385 cyclic populations, 530 D 101,360 decision theory, 127 decline long-term geometric rate of, 577 rate of, 596 declining-population paradigm, 339, 365 deleterious alleles, 162 fixation of, 353 deleterious mutations, 327, 350, 353-4, 363 accumulation of, 356 demographic rescue, 358 stochasticity, 84, 339-40, 344, 346 variance, 340 density dependence, 267, 338 density-dependent dispersal, 251,314, 329 immigration, 533 DIC, 381 diffusion coefficient, 86 modeling, 461 approximation approach, 572 approach, practical implications, 567 direct genetic effects, 272 disease colonization event, 480 colonization rate of, 482 decline, 486 dynamics, global coupling, 419 effect on population growth, 475 eradication, 442 extinction, 482 impact on population extinction, 475 incidence, 476 local abundance, 476 persistence, 423 prevalence, 476, 486-7 regional abundance, 476 resistance, 350, 473, 487 spread of, 488 transmission, 356, 473, 483, 487 dispersal, see also migration, 230, 250, 307, 389 balanced, 395, 402 breeding, 528 Daphnia, 61~$ causes of evolution, 309 condition-dependent, 307, 322 cost and benefits, 234, 322 curve, 460 density-dependent, 251,314-5, 329 directed, 530 effects of, 397, 401 ES rate, 231,233,249-50 evolution of, 233, 255 exploratory movement, 534 forces selecting for, 230 functions, 313, 462, 514 habitat heterogeneity, 310 in metapopulations, 230, 528 inbreeding, 313 kernel, 52, 85-6 kernel, Laplace, 379 landscape structure, 312 long-distance, 462 morphs, 319 mortality due to predation, 529 natal and breeding, 309 natural selection on, 402 negative density dependence, 329 one-way, 394 parental control, 323 passive, 86, 402, 405 pattern, 402, 409 physiological and behavioral control of, 320 see plant predation, 312, 323 presaturation, 308 propensity, 234, 235, 403 propensity, evolution of, 234 proximate control of, 319 rate, 248, 390, 394, 409 rate, habitat-specific, 311 rate, optimal, 253 rate, plasticity of, 403 saturation, 308 social factors, 314 species interactions, 332 state-dependent, 313, 318-21 stepping-stone, 534 strategy, conditional, 253 success, 28-30, 32 swamping effect of, 406 symmetric, 397 threshold, 30 distribution cauchy, 462 gaussian, 462 negative exponential, 462 686 disturbance, 145 effects of, 250 effect on dispersal rate evolution, 249 rate, 144 diversification dynamics of, 283 patterns of, 298 phase, 286 time homogeneous, 289 diversity, 140, 149 dominance, 165 coefficients, 162 effects, 362 drift load, 169 dynamic probability method, 554, 556 problem, 550 dynamic theory of island biogeography, 6-9, 83 E ecological niches, 408 evolutionary dynamics of, 408 ecotone, 396 edge effects, 38, 56 sensitivity, 38, 39 effective colonization rate, 94-5 density, 328 dispersal, 328 extinction rate, 93, 95 isolation, 27, 28 metapopulation size, 94-7, 166, 328 migration rate, 171,215 number of habitat patches, 95, 98 population size, 154, 156-7, 391 size, 155, 175, 193, 326, 370 size with source-sink structure, 159 elasticity of long-term growth rate, 590 emigration, 361 main proximate cause of extinction, 534 probability, 330 rate, 266-7 endangered species, 566 recovery planning for, 596 endogenous heterogeneity, 45 environment coarse-grained, 390 spatially heterogeneous, 392 environmental deterioration, population responses to, 356 heterogeneity, 55 INDEX stochasticity, 84-5, 339-40, 344 variance, 203, 340-1 epidemic theory demographic stochasticity, 420 traveling waves, 426 epidemics, 67 epidemiology, 102, 416 models, 103 theory, 12 epistasis, 200-1,262 epistatic interaction, 164 equilibrium assumption, 495 eradication threshold, 442 ESU, 581-2, 585-6 Euphydryas aurinia, 130, 494-5 editha, 9, 338, 400-1,491,503, 505, 507, 514 European nuthatch, 32-3 evolution of dispersal, 317, 467 dispersal, kin selection model, 231 dispersal, multiple factors, 317 dispersal, theoretical studies, 230 host-parasite interactions, 468 interactions among individuals, 260 life span in a metapopulation, 237 migration rate, 18, 102, 317, 467 species' ranges, 18-9 evolutionary equilibrium, 408 genetic theory, 259, 262 load, 349 significant units, 581 evolutionary dynamics in metapopulations, 275 of reproductive systems, 467 exogenous heterogeneity, 45 extinction, 20, 345 caused by disease, 519 correlates of, 526 dataset, 116 debt, 41, 94, 452, 494, 496, 532 effect of habitat patch size, 357 genetic effects of, 157 mechanisms of, 355, 365 probability of, 252 related to patch area, 115 rate, 344, 521 rate and emigration, 521 risk, 34, 346, 357 risk, effect of gene flow, 358, 362 risk, effect of migration, 358, 362 INDEX risk in dynamic landscapes, 40 risk, scaling with carrying capacity, 343 threshold, 34-7, 42, 89, 94, 98-9, 423, 465, 490, 494, 496 time, 80 vortices, 355 extinction-colonization probabilities, 107-8, 111, 114 processes, 230 rates, 83 stochasticity, 96, 98, 122, 533 F fadeouts, 421 fecundity, 248 fence effect, 308 field vole (Microtus agrestis), 130, 517, 520 Fisherian quantitative genetics, 201,223 fitness, 350 measure of, 404 sensitivity of, 404 variation, 263 fixation deleterious alleles, 169 index, 180 rate, 285 flour beetles, 267 foot-and-mouth epidemic, 428 forest models, 48 founder effect, 241 fractal landscape, 30-1, 35 fragmentation, 35 effects on extinction threshold, 36 effects on species richness, 145 fragmented communities, 144 frequency dependence, 163 frequency-dependent transmission, 418 Fst, 153-4, 160, 162, 173, 187, 180, 368-9 as an estimator of dispersal, 173 fugitive species, 135, 139-40, 145 G gametic disequilibrium, 371 equilibrium, 371 gene by deme interaction, 209-10 flow, 359, 361,405 for gene (GFG) resistance, 468 frequency change, 154 genealogies, 174-5 genealogies, effect of metapopulation structure on, 175, 189 687 interaction and speciation, 220 genealogical approaches, 368 branching patterns, 177, 190 histories, simulations of, 183 genealogies, branching patterns of, 177, 190 genealogy of a sample, 176 generation overlap, 349 genetic architecture, 297 architecture of sex, 241 background, 200 basis of dispersal, 319 contexts, intraindividual, 262 data to directly estimate immigration, 535 differentiation between populations, 405 differentiation between source and sink, 405 diversification, 281 diversity, 252, 349 diversity, effects of extinction and recolonization, 535 drift, 153-4, 230, 348 effects of population structure, described, 173 factors, relative importance, 338 load, 361 load in subdivided populations, 168 mixture coefficients, 376 neighborhoods, 474 rescue effect, 359, 365 resemblance, 242 stock identification method, see GSI structure of metapopulations, 16 threats, 347 variability, 348-9 variability, habitat fragmentation, 537 variability in metapopulations, 539 variability, loss of, 347 variance, 348 variance among demes, 208 variance components, 203 variance partitioning, 216 variance, fragmented populations, 536 variation, 207, 347, 499 variation in a sample, effect of metapopulation structure, 193 variation, effect of extinction, 187 variation, effect of recolonization, 187 variation, loss of, 350 variation, maintenance of, 412 genotype fitness, 160, 163 genotype-by-environment interactions, 263 genotypic value, 205, 216 ~8 INDEX I-I heterogeneous environments, 65 landscapes, 28 metapopulation, 95 patch networks, 81 SPOM, 11, 76 heterosis, 171,323, 327, 357 heterozygosity, 348 loss of, 357 heterozygous advantage, 351 highly fragmented landscapes, 15, 73-5, 496 history of metapopulation biology, holey adaptive landscapes, 18, 280, 297 host colonization, 480 extinction, 480 occurrence, 476 population, 415 host-pathogen system, simulation of, 486 habitat and metapopulation approaches, 497 area, 392 choice, active dispersal, 403 choice, evolution of, 412 configuration, 463 connectivity, 32, 456, 459 corridors, 312, 465 deterioration, 490 fragmentation, 99, 234, 492 fragmentation and extinction thresholds, 35 fragmentation, reduced species richness, 459 heterogeneity, 140 loss, 34, 99, 364, 493, 545, 563 loss and fragmentation, 42 loss, lagged response, 37 management, 508 networks, 500 patch, 75 quality, 129, 315, 392, 497, 508, 512 quality threshold, 497, 512 selection, 311 specialization, 137, 140 value of patches, 99 Halicoerus grypus, 375 hard selection, 160-1,166 haystack model, heritability, 203 of dispersal, 319 heritable variation, 262 hermaphrodites, 245 Hesperia comma, 235, 491,498, 507-9, 513 ideal free distribution, 311,332 ideal population, 155, 347 IGE, 259, 264, 268, 273 effects of, 269 immigration, 254 density-dependent, 517 negative density-dependent, 533-4 inbreeding, 245 avoidance, 314 coefficient, 206, 213, 351,538 decreasing resistance, 458 depression, 162, 314, 317, 350-2, 355 depression, effect of environmental conditions on, 352 depression, purging of, 351 effect of, 457 incidence dataset, 116 function model (IFM), 33, 82, 87, 90, 100-1, 108, 112, 116, 428, 453, 492, 495, 503, 508-9, 518, 521,558 matrix, 141 indirect genetic effects, 18, 259, 272 in metapopulations, 266 individual by deme interaction, 209 movement, 27, 505 individual-based model, 46-7, 98-9 infinite sites model, 179 integrodifference equations, 50 interacting particle systems, 48 interdemic selection, strength of, 265 geographical information systems, 43 distributions, 508 Glanville fritillary butterfly (Melitaea cinxia), 36, 74, 83, 89-90, 92-3, 129, 146-7, 229, 233,237, 315, 358, 491-3,500-1,503, 505 model, 95-6 global eradication (extinction) of disease, 423 graph theory, 32-3 gray seal, 375 great tit, 340 growth rate long-term, 341 stochastic, 341 GSI, 371,373, 376-7 gynodioecious species, 239 I INDEX interspecific competition, 65, 273 intragenomic conflicts, 241,243 introductions of Melitaea cinxia, 506 species, 504 invasion capacity of a network, 89-90 invasions, 65 island biogeographic theory, 6-9, 83 island model, 11, 15-6, 74, 76, 155, 168, 172, 187, 213, 368 of dispersal, 328 isolation by distance, 370 iteroparity, 238 K key patches, 100 kin competition, 231,235, 316, 319 cooperation, 316 selection, 231,255 kin-based interactions, 317 Krebs effect, 308 L Lactuca muralis, 467 lacunarity index, 31 Lande's model, 79, 81 landscape change, 551 connectivity, 24-9, 32-3 dynamics, 44, 563 ecological perspective, 25 ecology, 7-8, 11-2, 23, 102 fragmentation, evolutionary consequences of, 234 matrix, 24 mosaics, 24 structure, 34, 36, 76, 98, 101,491 lattice model, 46, 48, 68, 98 Leadbeater's possum, 99 Levins model, 74, 77, 79, 81, 93, 101 rule, 81, 100 Levins-type models, 90 life span, 235, 237 table, 38 life history, 347 density-dependent, 251 evolution, effect of genetic structure on, 255 evolution, in metapopulations, 228, 255-6 evolution, metapopulation effect on, 228 strategies, 252 689 strategies, evolution of, 246 strategies, evolutionarily stable, 255 syndromes, 237, 250 theory, 227, 229, 237 traits, 227 traits, evolution of, 243, 254 life-time reproductive contribution, 340 success, 396 likelihood ratio test, 373, 380 linkage disequilibrium, 371 Linum marginale, 468 load in subdivided populations, 171 local adaptation, 326, 407, 457 and regional patterns of diversity, 142 average effects, 207, 209, 212, 218 breeding value, 205-6 breeding value in metapopulations, 209 context, 261 diversity, 144-6, 148 drift load, 171 dynamics, 248 extinction, 84 extinction, effect of inbreeding on, 352 extinction, evolutionary effects of, 275 extinction in the metapopulation context, 358 mate competition, 317 population decline rate, 570 population size, 234 population size distribution, 570 population trends, differences in, 596 resource competition, 317 selection, 405 specialization, 501 logistic regression, 552 long-term dynamics, 132 Lotka-Volterra model, 52 M macroparasites, 416 mainland-island epidemic metapopulation, 426 metapopulation model, 413, 531 metapopulation structure, 490 major histocompatibility complex, 350 maladaptation, 411 in sink habitats, 410 management actions, 587 harvest, 588 local population level, 589 river hydropower systems, 588 690 INDEX Mantel test, partial, 385 marginal habitats, adaptation to, 404 Markov Chain, 80 Monte Carlo method, see MCMC Markov process, 80 mark-release-recapture, 505 of seeds, 461 techniques, 28 marsh fritillary butterfly, see Euphydryas aurinia mass effects, 140 mate choice, 310 maternal effects, 325 and state-dependent dispersal cues, 321 matrix, 313 population models, 12, 102 MCMC, 117-8, 183 mean additive genetic variance, within demes, 206 additive genetic variance, effect of drift on, 217 allele frequency, 160 local average effect, 210 phenotypic values, 200 relatedness of individuals, 161 time to extinction, 78, 353-4 measles, 420-1,423,426, 429-30, 432, 436 megapopulation, 500 Melitaea cinxia, see Glanville fritillary diamina, 233-4, 559-61 metacommunity, 133, 270 diversity, 140 indirect effects in, 270 mass effects perspective, 136 neutral perspective, 136 patch-dynamic perspective, 134 species-sorting perspective, 135 metapopulation approach, 3, 9, 11,514 approach, criticism of, 14-5 biology, fundamental processes, 367 citations to, classic, see classic metapopulation capacity, 35, 88-91, 93, 98, 465, 495, 500-1,506, 512 concept, 243, 413 concept, plant-specific problems, 451 decline, 492, 567, 570 diffusion approximation, 566 dynamic connectivity, 87 ecology, effect, 467 endangered, 565 evolution, 18 extinction, 40, 78, 98,~100, 311,362, 364, 496 extinction time, 127, 558 genealogies of samples from, 188 genetic structure, 273 genetics, 15 level management actions, 587 meltdown, 363 model, stochastic, 567 models, structured by the sizes of local populations, 101 of birds, of butterflies, of fishes, of mammals, of plants, patterns, 6, 494, 499 persistence, 34, 42, 129, 498 processes, processes, butterflies, 14 processes on population genetic structure, 536 quantitative genetics, 200, 205, 223 site selection, 559 size, 91,328 size distribution, 570 spatially realistic theory, 9, 11, 76, 83 stochastic theory, 94 structures, theory 8, 228 trajectories, 573 viability metrics, 577, 584, 587 with very small local populations, 538 Metropolis-Hastings algorithm, 184 MHC loci, 350, 352 Microbotryum violaceum, 473,487 microsatellite DNA polymorphism, 536 genotyping, 528 Microtus oeconomus, 530 rossiomeridionalis, 532 migrant pool, 158 model, 369 migration, see also dispersal, 153, 360, 505 corridor, 360 cost of, 505 costs and benefits of, 323 density dependence, 528 diversity, 144 in a metapopulation context, 385 691 INDEX limitation, 140 load, 170 process, 367 rate, 372 rate and diversity, 139 migrational meltdown, 361,364 minimum set coverage formulation, 547 missing data, 118-20 years, 117 moment closure, 58-9 moment equation approaches, 68 Moran effect, 426 mosaic models, 456 movement parameters, habitat-specific, 27 multilocus BDM model, properties of, 282 genotype approaches, 368 genotype methods, 20, 370 multiple equilibria, 90 multitrophic interactions, 459 mutation, 17, 162, 179 accumulation, 353 and genealogical process, 181 average fitness effects, 354 load, 168 number of, 181 parameter, 179 rate, genome-wide, 355 rate, per-genome, 354 mutational meltdown, 170, 327, 354, 356, 363, 365 mutation-selection balance, 162 mutualistic relationship, 459 N natural experiments, 401 selection, 162 selection in metapopulations, 259 Ne, 156, 173, 349, 354 effect of population structure on, 157 network connectivity, 501 neutral landscape models, 32, 34 metacommunity perspective, 143 theories of community structure, 10 neutrality tests, 180, 185 non-equilibrium systems, 490 non-linear interaction, 260 non-random mating, 155, 163 number of alleles, 180 of species, 276-7, 292 O Oncorhynchus species, 581 open and closed population structures, 490 opposite-sex attraction, 534 Orkney Isles, 376 outbred vigor, 360 outbreeding depression, 361-2 overdominance, 163, 351 P pair approximation, 49 panmictic population, 175 parapatric speciation, 289 Pararge aegeria, 510 parasitoid-prey system, 51, 61 parentage assignment, 528 parental control of offspring dispersal, 323 passive propagule dispersal, 86 patch areas, 76 connectivity, 26 dynamics, 139, 456 network, homogeneous, 77 occupancies, 114 occupancy models, 77 turnover, 40 values, 88, 91, 93, 100 pathogen colonization, 485 extinction, 485 occurrence, 476 percolation, 29 theory, 26, 31, 100 threshold, 29-30, 32 phase locking, 425 phenotypic effects of alleles, 200 of genes, 262 phenotypic plasticity, 457 variance, 201,203, 263 variation, partitioning of, 263 philopatry, 316 phylogenetic envelope, 391 pitcher plants, 145, 147-8 plant dispersal, 460-1 dispersal, molecular markers, 462 dispersal, distance curve, 460 metapopulations, 449, 475 plant-animal interactions, 459 plant-pathogen metapopulation, 471 692 plant-specific problems with metapopulation concept, 451 plasticity, 251 Plebejus argus, 131, 234, 491,501-4, 512, 552, 556-7 point speciation model, 277 pollinators, 458 polymorphic nucleotide sites, number of, 181 sites, expected number of, 177 sites, expected value, 181 sites, total number of, 181 polymorphism, protected, 247 pool frog (Rana lessonae), 129 population biology, perspective in, 338 population age-structured, 395 bottleneck, 348-50 connectivity, 498 density, 246, 387 differentiation, 213,223 dynamics, 250, 345-6 dynamics, transient, 256 extinction, 337, 355 extinction, ceiling model, 343 extinction, ecological factors, 339 extinction, genetic factors, 347 extinction rate, 358 fluctuations, 340, 344 genetic methods, 368 genetic processes, 172 growth rate, 344, 346 history, inferences about, 182 mean fitness, 353 neighborhood, 63-4 persistence, effects of source-sink structure, 398 regulation, 337-8, 390 size, effect of source-sink structure, 397 stability, effects of source-sink structure, 398 stage-structured, 395 structure, inferences about, 182 synchrony, 329 turnover, 455, 502-3, 524, 526 turnover, genetic effects of, 368 viability analysis, 13, 20, 338, 565, 595 portfolio effect, 421 posterior distribution, 374 predator-prey interaction, 66 models, 54 INDEX prior distribution, effect of, 384 probability of coexistence, 332 common origin, 370 extinction, 355 fixation, 165 fixation, effect of population structure on, 168 fixation of beneficial alleles, 165-7 fixation of additive alleles, 165 fixation of neutral mutations, 285 fixation of new mutations, 165-6 Proclossiana eunomia, 491 productivity, 142, 248 index, 378 propagule-pool colonization, 535 model, 369-70 pattern, 538 propagules, colonizing, 369 protected polymorphism, 247 PVA, 565-6 Q quantitative genetic parameters, effect of migration on, 213 quantitative genetics, 199 quasiequilibrium, 495 quasistationarity, 114, 122, 131 assumption, 122 quasistationary distribution, 78-80, 82, 97 equilibrium, 116 state, 107-8, 124 II radiation adaptive, 298 duration of, 289, 297 dynamics of, 289 time to beginning of, 297 random fission model, 277 random walk, 86 model, 28 range boundaries, 507 expansion, 509 margins, 507 rate of expansion, 508, 510, 512 habitat destruction, 41 reaction norm, 252 reaction-diffusion, 50 693 INDEX reciprocal transplant experiments, 457 recolonization, 241,243, 245, 254, 526 genetic effects of, 157 recombination, 153 genealogies and, 178 recruitment from seed rain vs seed bank, 455 Red List criteria, 566 regional coexistence, 146 diversity, 144, 146, 148 dynamics, 248 stochasticity, 89, 96, 100, 121, 130, 363, 514 stochasticity, migration and, 363 relatedness, 242 remnant populations, 452-3, 455-6 representativeness, 542 reproduction assurance, 241 reproductive effort, 246, 248, 250, 252 age-specific, 238 ES, 235, 248 evolution of, 235 in a metapopulation, 235 in a metapopulation, evolution of, 246 reproductive isolation, 281,284, 296 genetic architecture of, 279 genetics of, 278 threshold effect, 297 reproductive success in sink habitats, 410 reproductive value, 349, 391,404-5 habitat specific, 391 rescue effect, 13, 87, 90, 101, 107-8, 111-2, 115, 130, 310, 328, 464, 518-9, 529, 531,534 reserve aggregation, 550, 554-5, 562 reserve network, 541 design, 542 design, landscape dynamics, 546 design, spatial population dynamics, 546 dispersal ability, 560 patch cost, 560 reserve selection, 541 algorithms, 542, 544 algorithms, persistence, 546 amount of resources, 561 resistance, fitness costs of, 473 resource partitioning, 137 restoration efforts, 596 Reversible Jump Markov Chain Monte Carlo, 381 risk metric confidence intervals for, 578 uncertainty, 578 variability of, 595 risk of extinction, 340, 344, 353, 356-7, 359 long-term, 596 short-term, 596 RJMCMC, 381 $ salmon, 581 scattering phase, 189, 191-3 scoring approaches, 543 seasonal forcing, 426 seed banks, 454, 456 dimorphism, 319 sowing, 451 segregating sites expected number of, 177 expected value, 181 segregation load, 169 SEIR model, 422 selection, 153, 219, 230, 237 among local populations, 18 balancing, 163 differential, 203 directional, 221 diversifying, 200 hard, 160-1, 166 heterogeneous, 164 in metapopulations, 264 in subdivided populations, 159 individual, 266, 268, 270 interdemic, 266, 268-70 response to, 160, 200, 203, 266, 404 soft, 160-1, 166 spatially heterogeneous, 159, 164, 245 stabilizing, 165 strength of, 160 temporal heterogeneity in, 245 uniform, 159-60 selfing 245 semelparous organisms, 246 semi-independent patch network (SIN), 499-500 Senecio jacobaea, 146 settlement decision, 334 pattern, 529 success, 324-5 694 sex allocation, 238, 243, 245 inheritance, mode of, 242 ratio, 241-2 ratio, evolution, 239 ratio, evolution in a metapopulation, 239 ratio variation, 239 sex-biased dispersal, 313-4, 317, 521 sexual selection, 313 sexually transmitted diseases, 427 shifting balance theory, 18, 186 shrews, 130, 344, 347, 531 Silene alba, 360 lati[olia, 457, 473, 487 Silene-Microbotryum metapopulation, 475, 485 sink, 40 absolute, 388 black-hole, 394 habitat, 388, 392, 398, 401 populations, 136, 141, 158 relative, 388 SIR model, 416, 427, 442 SIS model, 427 site selection algorithms, 542 stochastic metapopulation model, 558 dispersal ability, 559 site-frequency distribution, 185 size-structured populations, 102 small mammals, 515 small-population paradigm, 339, 365 snapshot data, 112, 114-5 social contact networks, 67 context, average, 264 context, genetic variation in, 265 fence, 329 fence effect, 534 soft selection, 160-1,166 Sorex, 344 araneus, 130, 531 caecutiens, 130, 531 minutus, 130, 531 source habitat, 388, 392 populations, 136, 141,158 source-sink concept, applied aspects of, 414 dynamics, 37, 143, 497 dynamics, ecological consequences of, 397, 399 INDEX dynamics, influence on population size, 401 metapopulations, 17, 19, 311 metapopulations, evolutionary models of, 408 models, predictions of, 402 population dynamics, 387-8, 408 populations, 404, 453 potential, 37-8 relations, 139 structure, 387, 398-400 structure, concept of, 413 structure, ecological consequences of, 400 structure, effect of, 158 structure, effect on adaptive evolution, 405 structure, evolutionary consequences, 403 structure, evolutionary stability of, 402 structure, patch model of, 389 structure, reverse, 394, 410 system, experimental, 401 system, polygenic model of evolution in, 406 systems, 358 systems, laboratory, 412 spatial aggregation, configuration, 497 correlation, 97 coupling in epidemic metapopulations, 433 ecology, 4, genetic structure, 327 locations, 76 logistic model, 64 moment equations, 46, 51, 98 point process, 47 population viability analysis, 559 reserve design, 547 synchrony, 50 spatially correlated environmental conditions, 96 correlated extinction, 518, 527, 532-3 explicit approach, 396 explicit models, 47, 395 explicit models, consequences for ecological processes, 471-2 explicit models, consequences for genetic processes, 471-2 realistic Levins model, 82, 84 realistic metapopulation theory, 9, 11, 76, 83 realistic models, 234 speciation, 221,223, 278, 296, 298 by random drift and mutation, 300 dynamics, 296 in metapopulations, 275 mechanisms, 221 INDEX probability of, 278 rate of, 291 species extinction, 364 range, 364, 397 sorting, 139-40, 147 species-area curves, 276-7 relationship, SPOM, 11, 74-75, 105-6, 112, 428 classification, 89 deterministic approximation of, 76, 82 homogeneous, 11, 76 homogeneous, deterministic approximation of, 11 SRLM, 93-4 stable equilibria, 346 stationary state, 244 statistical genetics, 20 methods, limitations of, 375 stepping-stone dispersal, 534 model, 16 stochastic and seasonal forcing, 420 cellular automata, 48 forcing of epidemics, 420 logistic model, 78-9, 81, 98 logistic model, deterministic mean-field approximation of, 77 metapopulation equilibrium, 82 metapopulation model, effects of management, 587 metapopulation, trajectory of, 596 patch occupancy models, see SPOM theory, 94 structured coalescent, 187-8 metapopulation model, 12, 440 populations, genetics of, 186 successional habitats, 513 susceptible-infected-susceptible (SIS) model, 103 T tagging experiments, 582 temporal variation, 97 temporally varying environmental conditions, 96 territoriality, 314 threatened species, recovery planning for, 596 threats to populations, 365 695 threshold condition, 79, 81, 89 in connectivity, 29 response, 31 value, 80 time delay (lag), 454-6, 465, 494 in metapopulation response, 494 time to extinction, 91, 344, 452, 495-6, 499, 503 most recent common ancestor, 192 total coalescence times, expected, 178 total length of the genealogy, distribution, 177 trade-off, 137-8, 146, 148-9 between competitive ability and predator tolerance, 148 between colonization and competitive abilities, 135, 146 transient dynamics, 24, 93 dynamics, evolutionary importance of, 254 dynamics of speciation, 288 growth, 248 metapopulation dynamics, 490 period in population dynamics, 243-4 population dynamics, 250 transmission model density-dependent, 482 frequency-dependent, 482 transplantations, 451 traveling waves of infection, 434 tree frog (Hyla arborea), 106, 109-10, 122-4, 128 Tribolium castaneum, 266, 271-2 trophic cascades, 143 turnover, 125, 127, 132 data, 114-5, 117-8 events, 506 underestimate of, 132 U unstable equilibrium, 346 Urophora cardui, 129 Urtica dioica, 459 Uta stansburiana, 316, 320 V vaccination pulse, 442-4 random, 443 strategies, 442 Vancouver Island marmot (Marmota vancouverensis), 532 696 variance in local average effects, 210 local average effects, effect of drift on, 217 reproductive success, 155-7 viability analysis, 565 metric validation, simulations for, 579 of total metapopulation, 38, 569 INDEX W water vole metapopulations, 522 wavelet phase analysis, 434 white-backed woodpecker (Dendrocopos leucotos), 81 whooping cough, 421-2 Wright-Fisher model, 176 ... entitled one of the five main sections of his Population Biology of Plants as "The effects of neighbours." It has been less obvious that spatial positions of individuals matter in the case of mobile... and other interactions Likewise, we may use the term spatial population biology to emphasize the influence of the spatial positions of individuals and populations on their genetic and evolutionary... animals, which may form more or less random-mating (panmictic) populations However, from the point of view of ecological interactions, spatial positions often matter even in mobile animals One

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