Population dynamics L5 English in Natural Science 自然科学の英語 2006 自然科学の英語-ENS-L5 Abundance Birds in river forests (Spain) (Sanchez-Bayo, F. 1985) 2006 自然科学の英語-ENS-L5 Abundance vs body size Mammals Log(Y) = 1.3-0.66*log[X] Birds Log(Y) = 0.22-0.54*log[X] 350 mammal and 552 bird species (Silva et al., 1997) • Small animals are more abundant than large ones • Birds are less abundant than mammals 2006 自然科学の英語-ENS-L5 Abundance and distribution Jim Brown (1984) “Population densities Western grey kangaroo (Macropus fuliginosus) (Caughley et al. 1987) decrease towards the boundary of the geographical range of a species” Ilkka Hanski (1982) “Widespread species tend to be more abundant” 1. Sampling artifact 2. Specialization Generalist - large area Specialist - small area 2006 自然科学の英語-ENS-L5 3. Metapopulations and dispersal Abundance vs distribution range Rapoport’s rule (1975) “Geographic range size decreases from polar to equatorial latitudes, with smallest range sizes in the tropics” Why? 523 North American mammals (Pagel et al. 1991) 1. Tolerance 2. Dispersal favours generalist species 3. Competition Tolerance Dispersal + 2006 自然科学の英語-ENS-L5 + + Competition Population dynamics Big mammals • Parameters – Natality (fertility) rate Birds • Offspring, reproduction r x = bx ÷ nx Fish invertebrates Small mammals – Mortality rate • Life expectancy (longevity) qx = dx ÷ nx • Life tables – Mortality/cohort – Age, sex structure 2006 自然科学の英語-ENS-L5 Intrinsic capacity for increase r (Lotka, 1925) • Exponential – constant rate (%) Nt = N0 ert Exponential N population – Finite rate of increase = er Linear individual – Doubling time: time for a quantity to double Dt = 70 ÷ r • Logistic K = carrying capacity Nt = K ÷ (1+ ea-rt) 2006 • Linear 自然科学の英語-ENS-L5 – constant amount y=x+A Natural processes • Exponential reduction Exponential growth – Populations – Radioactive residues – Chemical concentration (human, r = 1.7% year) – Food consumption – Waste production – Economy (Japan: 1-2% year) (USA: 5% year) (China: 7% year) (eg. pesticides, pollutants) – Forest destruction 1 2 3 Linear growth 2006 Gone! 4 – [CO2] atmosphere 5 – Food production (?) 6 – industry 自然科学の英語-ENS-L5 Limits to population growth • Food resource – carrying capacity (K) T.R. Malthus (1766-1834) • Predators • Abiotic factors – Temperature – Water availability Reindeer Daphnia rosea (Scheffer, 1951) 2006 自然科学の英語-ENS-L5 (Walters et al., 1990) Life strategies – Generalist niche – Unstable populations – Quick recovery • Low r (K strategy) – Specialist niche – Stable populations – Prone to extinction • Decisive factor: – Mortality rate High Low 2006 Repeated reproduction (K strategy) Reproductive success r unrelated to abundance • High r (r strategy) Big-bang reproduction (r strategy) Reproductive effort • How to increase r ? – Larger offspring size (r) – Increase longevity (K) r K • more times to reproduce – Younger reproductive age (both r and K) 自然科学の英語-ENS-L5 Stationary distribution No population increase in time Fertility rate = mortality rate r = qx 2006 自然科学の英語-ENS-L5 Competition • Resource competition – Inter or intraspecific • Interference competition (contest) – Usually intraspecific – Sex: males only 2006 Resources • Plants – Water – Light – Nutrients in soil • Animals – Food – Space 自然科学の英語-ENS-L5 Competition: Mathematical models Species 1 wins Species 2 wins Lotka (1925) and Volterra (1926) Species 1 dN1 K1-N1-aN2 =r1N1 dt K1 Coexistence Exclusion Species 2 dN2 K2-N2-bN1 =r2N2 dt K2 2006 自然科学の英語-ENS-L5 Zero growth 2006 Tilman model (1990) 1 Neither species can live 2 Only species A can live 3 Species A wins 4 Species A & B co-exist 5 Species B wins 6 Only species B can live • Equilibrium point depends on rate of consumption of resources 1 and 2 R1: rate A > rate B 自然科学の英語-ENS-L5 A wins Co-existence Species must occupy different niches (Gause, 1934) 1. resource partitioning (share) Saccharomyces + Schizosaccharomyces yeast (Gause, 1932) 2. Spatial segregation outside inside 2006 自然科学の英語-ENS-L5 Grain beetles in wheat (Birch,1953) Segregation • Efficient utilization of the same resource – Habitat (space) – Size of prey (diet) – Time • Day - night • Seasons (migration) • Mechanism of evolution Tern species in Christmas Island (Ashmole, 1968) 2006 – r and K selection theory (MacArthur & Wilson, 1967) 自然科学の英語-ENS-L5 Predation & parasitism • Natural agents to control populations – Exponential increase logistic model • Exponential reproduction ‘biomass waste’ – Producers: plants, phytoplankton – Predation: one species eats another • Herbivores: eat plants • Carnivores/parasites: eat herbivores (prey) • Predators/parasites USE that extra biomass • Predators – External – Big size 2006 • Parasites – Internal - live on host – Small size (i.e.larvae) 自然科学の英語-ENS-L5 Predators and parasites depend on prey/host Abundance Prey (lemming) abundance bird predators 2006 自然科学の英語-ENS-L5 Models • Discrete populations: one generation/year – Prey Nt+1 = (1-B zt)Nt-C NtPt – Predator Pt+1 = Q NeqPt B = prey reproductive rate C = predator efficiency Q = predator reproductive rate Parasitic wasp Prey = Host (Utida, 1957) 2006 自然科学の英語-ENS-L5 Continuous generations • Lotka (1925) and Volterra (1926): unrealistic • Rosenzweig-MacArthur (1963) Predator equilibrium Predation Food shortage equilibrium Environmental pressure (Carrying capacity) Predator density (P) Intraspecific competition Prey population density (N) Caribou (Bergerud 1980; Sinclair 1989) 2006 自然科学の英語-ENS-L5 Net reproductive rate (R0) = number of female offspring / female / generation Population regulation Birth rate (b) UP Death rate (d) DOWN Stochastic variation Humans R0 = 1.1 Stable populations Birth rate (b) DOWN Death rate (d) UP predation disease food shortage 2006 自然科学の英語-ENS-L5 Extinction • Species ceases to exist • Causes – Habitat loss – Introduced species (competition, predation) – Overkill – stochasticity • Human impact – Habitat destruction – Overkill (e.g. Dodo, Mammoth, Moa) 2006 Probability of extinction (Pielou, 1969) P = (d/b)N0 d = death rate b = birth rate N0 = initial population size 1) b > d 2) b < d 3) b = d P > 1.0 survival P < 1.0 extinction P = 1.0 extinction because of stochastic changes in a lifetime (e.g. disease, climate) 自然科学の英語-ENS-L5 Natural extinction • Geological eras and periods – Characterised by changes in biodiversity • Extinction of old forms • Apparition of new forms • Natural causes – Atmospheric composition • Plants increased O2 and decreased CO2 – Astronomic - Milankovitch cycles • Climate variation (i.e. iceage) – Catastrophes (Cuvier, 1769-1832) • Five major extinction events • Cause: asteroids? Earth’s geochemistry? 2006 自然科学の英語-ENS-L5 Historical extinction events 15% families 50% genera 52% families 95% species 2006 自然科学の英語-ENS-L5 Cretaceous-Triasic boundary extraterrestrial iridium layer Italy Caribbean 2006 (Alvarez et al. 1980, 82) 自然科学の英語-ENS-L5 meteorite Denmark (Kastner et al. 1984) Mass extinctions…recovery 2006 自然科学の英語-ENS-L5 Evolution and extinction • Extinction is an irreversible process • Extinction events have a founder effect – New taxa appear – Biodiversity flourishes, even more than before • Eventually all species go extinct – Evolve to generate another species (average lifetime of species is 10 m years) – Stop existing - gone! 2006 自然科学の英語-ENS-L5 References • Charles J. Krebs. 2001. Ecology 5th ed. / 応用動物昆虫学 B-226 • Tokeshi M. 1999. Species coexistence: ecological and evolutionary perspectives / 応用動物昆虫学 B-207 • Alvarez, L. W., W. Alvarez, et al. 1980. Extraterrestrial cause for the CretaceousTertiary extinction. Science 208: 1095-1108 2006 自然科学の英語-ENS-L5