Population Dynamics Focus on births (B) & deaths (D) B = bNt , where b = per capita rate (births per individual per time) D = dNt N = bNt – dNt = (b-d)Nt Exponential Growth • Density-independent growth models Discrete birth intervals (Birth Pulse) vs. Continuous breeding (Birth Flow) G e o m e t r i c G r o w t h ( B i r t h P u l s e ) 1 4 0 1 2 0 N = N t 0 1 0 0 = 2 , N = 2 0 t >1 <1 =1 N 8 0 6 0 4 0 Nt = N0 t 2 0 0 0123456 T i m e Geometric Growth • When generations do not overlap, growth can be modeled geometrically. Nt = Noλt – – – – Nt = Number of individuals at time t. No = Initial number of individuals. λ = Geometric rate of increase. t = Number of time intervals or generations. Exponential Growth Birth Pulse Population (Geometric Growth) e.g., woodchucks (10 individuals to 20 indivuals) N0 = 10, N1 = 20 N1 = N0 , where = growth multiplier = finite rate of increase >1 <1 =1 Exponential Growth Birth Pulse Population N2 = 40 = N1 N2 = (N0 ) = N0 2 Nt = N0 t Nt+1 = Nt Exponential Growth • Density-independent growth models Discrete birth intervals (Birth Pulse) vs. Continuous breeding (Birth Flow) Exponential Growth • Continuous population growth in an unlimited environment can be modeled exponentially. dN / dt = rN • Appropriate for populations with overlapping generations. – As population size (N) increases, rate of population increase (dN/dt) gets larger. Exponential Growth • For an exponentially growing population, size at any time can be calculated as: Nt = Noert • • • • • Nt = Number individuals at time t. N0 = Initial number of individuals. e = Base of natural logarithms = 2.718281828459 r = Per capita rate of increase. t = Number of time intervals. Exponential Population Growth Exponential Population Growth ln(N t) l n ( N ) = l n ( N ) + r t t 0 r = e p lo s Nt = N0ert Difference Eqn Note: λ = er l n ( N ) 0 T i m e Exponential growth and change over time N = N0ert Number (N) Slope (dN/dt) dN/dt = rN Time (t) Slope = (change in N) / (change in time) = dN / dt Number (N) ON THE MEANING OF r rm - intrinsic rate of increase – unlimited resourses rmax – absolute maximal rm r - also called rc = observed r>0 r<0 r=0 G e o g r a p h i c R a n g e a n d V a l u e s o f r m r < 0 m x r > 0 m r > > 0 m x r m a x Intrinsic Rates of Increase • On average, small organisms have higher rates of per capita increase and more variable populations than large organisms. Growth of a Whale Population • Pacific gray whale (Eschrichtius robustus) divided into Western and Eastern Pacific subpopulations. – Rice and Wolman estimated average annual mortality rate of 0.089 and calculated annual birth rate of 0.13. Growth of a Whale Population • Reilly et.al. used annual migration counts from 1967-1980 to obtain 2.5% growth rate. – Thus from 1967-1980, pattern of growth in California gray whale population fit the exponential model: Nt = Noe0.025t Logistic Population Growth • As resources are depleted, population growth rate slows and eventually stops • Sigmoid (S-shaped) population growth curve – Carrying capacity (K): Logistic Population Growth Logistic Population Growth dN / dt = rN dN/dt = rN(1-N/K) • r = per capita rate of increase • When N nears K, the right side of the equation nears zero – As population size increases, logistic growth rate becomes a small fraction of growth rate Exponential & Logistic Growth (J & S Curve) Logistic Growth Actual Growth Populations Fluctuate Limits to Population Growth • Environment limits population growth by altering birth and death rates – Density-dependent factors – Density-independent factors Galapagos Finch Population Growth Readings • • • • • 8.3 through 8.13; pp. 162-177 Field Studies, pp. 164-165 Ecological Issues, p. 175 Ecological Issues, p. 219 Field Studies, pp. 230-231 Logistic Population Model A. Discrete equation Nt = 2, R = 0.15, K = 450 Logistic Population Growth - Built in time lag = 1 - Nt+1 depends on Nt N(t) 500.0 400.0 300.0 200.0 100.0 0.0 0 20 40 60 Time Nt N t 1 N t r N t 1 K 80 100 I. B. Logistic Population Model Density Dependence I. Logistic Population Model C. Assumptions • No immigration or emigration • No age or stage structure to influence births and deaths • No genetic structure to influence births and deaths • No time lags in continuous model I. Logistic Population Model C. Assumptions Per Capita Growth Rate (N(t+1) - N(t))/N(t) • Linear relationship of per capita growth rate and population size (linear DD) 0.2000 0.1500 0.1000 0.0500 0.0000 K 0.0 100.0 200.0 300.0 N(t) 400.0 500.0 I. Logistic Population Model C. Assumptions • Linear relationship of per capita growth rate and population size (linear DD) I. Logistic Population Model Discrete equation Nt = 2, r = 1.9, K = 450 N(t) Logistic Population Growth 600.0 500.0 400.0 300.0 200.0 100.0 0.0 Damped Oscillations r <2.0 0 20 40 60 Time 80 100 I. Logistic Population Model Discrete equation N(t) Logistic Population Growth Nt = 2, r = 2.5, K = 450 600.0 500.0 400.0 300.0 200.0 100.0 0.0 0 20 40 60 Time 80 100 Stable Limit Cycles 2.0 < r < 2.57 * K = midpoint I. Logistic Population Model Discrete equation Nt = 2, r = 2.9, K = 450 Logistic Population Growth 800.0 N(t) 600.0 400.0 200.0 0.0 0 20 40 60 Time 80 100 Chaos r > 2.57 • Not random change • Due to DD feedback and time lag in model Underpopulation or Allee Effect b=d b=d d Vital rate b<d r<0 b N* N K I. Review of Logistic Population Model D. Deterministic vs. Stochastic Models Random R0 and K Logistic Growth Model Nt = 1, r = 2, K = 100 120.000 100.000 N(t) 80.000 60.000 40.000 20.000 0.000 0 20 40 60 Time (t) 80 100 120 I. Review of Logistic Population Model D. Deterministic vs. Stochastic Random R0 and K Models Logistic Growth Model Nt = 1, r = 0.15, SD = 0.1; K = 100, SD = 20 120.000 100.000 N(t) 80.000 60.000 40.000 20.000 0.000 0 20 40 60 Time (t) 80 100 120 * Stochastic model, r and K change at random each time step I. Review of Logistic Population Model D. Deterministic vs. Stochastic Random R0 and K Models Logistic Growth Model Nt = 1, r = 0.15, SD = 0.1; K = 100, SD = 20 120.000 100.000 N(t) 80.000 60.000 40.000 * Stochastic model 20.000 0.000 0 20 40 60 Time (t) 80 100 120 I. Review of Logistic Population Model D. Deterministic vs. Stochastic Random R0 and K Models Logistic Growth Model Nt = 1, r = 0.15, SD = 0.1; K = 100, SD = 20 120.000 100.000 N(t) 80.000 60.000 40.000 * Stochastic model 20.000 0.000 0 20 40 60 Time (t) 80 100 120 II. Environmental Stochasticity A. Defined • Unpredictable change in environment occurring in time & space • Random “good” or “bad” years in terms of changes in r and/or K • Random variation in environmental conditions in separate populations • Catastrophes = extreme form of environmental variation such as floods, fires, droughts • High variability can lead to dramatic fluctuations in populations, perhaps leading to extinction II. Environmental Stochasiticity B. Examples – variable fecundity Relation Dec-Apr rainfall and number of juvenile California quail per adult (Botsford et al. 1988 in Akcakaya et al. 1999) II. Environmental Stochasiticity B. Examples - variable survivorship Relation total rainfall pre-nesting and proportion of Scrub Jay nests to fledge (Woolfenden and Fitzpatrik 1984 in Akcakaya et al. 1999) II. Environmental Stochasiticity B. Examples – variable rate of increase 6 5 Muskox population on Nunivak Island, 1947-1964 (Akcakaya et al. 1999) 4 3 2 1 0 0.925 0.975 1.025 1.075 1.125 1.175 1.225 1.275 Growth Rate Number of Animals II. Environmental Stochasiticity - Abundance Example of random of Wildebeest in the K Serengeti 1600000 1400000 1200000 1000000 800000 600000 400000 200000 0 1954 1959 1964 1969 1974 1979 1984 1989 Years • Serengeti wildebeest data set – recovering from Rinderpest outbreak – Fluctuations around K possibly related to rainfall Exponential vs. Logistic dN N rN 1 dt K dN rN dt No DD All populations same DD All populations same No Spatial component Incorporating Space Metapopulation: a population of subpopulations linked by dispersal of organisms Two processes = extinction & recolonization • subpopulations separated by unsuitable habitat (“oceanic island-like”) • subpopulations can differ in population size & distance between Metapopulation Model (Look familiar?) dp cp 1 p ep dt p = habitat patch (subpopulation) c = colonization e = extinction Another Population Model Source-sink Dynamics: grouping of multiple subpopulations, some are sinks & some are sources Source Population = births > deaths = net exporter Sink Population = births < deaths >1 <1 <1 The probability of dispersal from one patch to another depends on: • Distance between patches – inverse relation • Nature of habitat corridors linking the patches • Ability of the species to disperse (vagility or mobility) – dependent on body size – positive relation