Number of Animals Population Ecology 72 73 74 75 76 77 78 79 80 Year Goal of Population Ecology is to Describe the Composition of Populations Through Time and Understand Population Fluctuations Describing Population Composition Sex Ratio Age Ratio Genetic Composition Spatial Structuring Sex Ratio Indicates Important Processes in Population Sex Ratio (Males:Females) in Flock of Pinyon Jays Population growth potential--greater male bias = less growth ability (sexual species) Breeding System Dispersal (Data from Marzluff and Balda 1992) 72 73 74 75 76 77 78 79 80 Age Pyramids Summarize Age Structure Differ for Increasing, Steady, and Declining Populations Indicate Bad Years, Bottlenecks in Reproduction, etc. Proportion in each age class Increasing Population Stable Population Declining Population Pinyon Jays Were Studied for 20 Years Long-term studies of marked animals are needed to get accurate population growth and composition information. Age Structure Reflects Relative Productivity of Cohorts Young (cohort) from productive years constitute large proportion of population for many years (1977, 1978) A poor year of reproduction continues to be echoed in population as a missing cohort (1976) 300 Number of Jays in Flock 50 73 1978 1977 1976 74 75 76 (Marzluff & Balda 1992) 77 78 YEAR 79 80 81 82 Importance of Indirect and Direct Selection Depends on Genetic Composition of Population Number of Relatives in Flock (Marzluff & Balda 1992) Age of Focal Individual Density of Great Tits in 4 Areas Describing Change in Population Size (Lack 1966) Year Managers are usually concerned with monitoring population SIZE---So, How do WE Quantify CHANGE in Population Size?? Population size and rates of growth Population size: Nt = population size at time t Nt+1 = population size at time t+1 Nt+1 = Nt + Births + Immigration – Deaths -Emigration Growth rates: r = exponential growth rate λ (‘lambda’) = intrinsic population growth rate Population growth Reproduction, births, natality (B) Immigration (I) Population Mortality, death (D) “BIDE” Emigration (E) Age-specific birth rates A fecundity schedule for Chamois from New Zealand. Age (yrs) N # Female births per pregnant female _____________________________________________ 0 0.000 1 60 2 0.017 2 36 14 0.194 3 70 52 0.371 4 48 45 0.469 5 26 19 0.365 6 19 16 0.421 7 6 5 0.417 >7 10 7 0.350 ___________________________________________ Survivorship curves for male & female moose on Isle Royale females Survivors (lx) males 2 4 6 8 10 12 14 16 18 20 Age at Death (years) Emigration and Immigration Juvenile dispersal: movement from place of birth to place of breeding Breeding dispersal: movement by adults from one place of breeding to another Birds: Female dispersing sex Mammals: Male dispersing sex # # # ## # # # American Robin post-fledging movements # # ## # # # # # # # # # # # # # # # # # ## # ## ## # # # ## ## # ## # ### # 0 500 1000 1500 2000 2500 3000 3500 4000 Meters Population Growth Carrying capacity (k) Exponential N k N Logistic time Classic growth curve, unlimited resources time Classic growth curve, limited resources (k) The Simplest Quantification of Population Growth Assumes Exponential Growth Nt=N0ert-----let t = 1 year N1=N0er er=N1/N0===Lambda, Finite rate of Increase Lambda goes from 0 (extinction) to 1 (stable growth) to positive infinity (Exponential growth of various magnitude) Exponent Indicates the Magnitude of Change er=N2/N1---Take ln (natural log, loge) of both sides to get: r = ln(N2/N1) varies from negative infinity (decrease) to 0 (Stable) to positive infinity (increase) r, the exponential multiplier, = Intrinsic (instantaneous) rate of increase Exponents provide consistent quantification of magnitude of change Doubling and halving of population produces same exponent multiplier of change----sign of multiplier changes N1=50 ---N2=100--doubling er = (lamda) = 100/50 = 2 r = ln (2) = .693 N1=100--N2=50---halving er = (lamda) = 50/100 = 0.5 r = ln (0.5) = -.693 Units of r and lambda Units of lambda are obvious numbers per unit time restricted to the unit it was calculated over t = 1 year, then rate is change per year Units of r not obvious it is a multiplier, not a rate “growth multiplier of ln(#s) per unit time” not restricted to unit it was calculated over r from 1 year can be transformed to r for each day by dividing by 365, etc. Lambda and r Both present the same information in varying formats Population increases at lambda per unit time or r at any instant in time r is useful because it can be transformed to fit time interval of interest, lambda is more intuitive Unlimited Growth Australian rabbit (European hare) • 1859: 24 hares introduced (for human food?) • 1865: over 20,000 hares were harvested, actual population much greater. • Mid-1800’s to mid-1900’s: major problem with too many hares; caused habitat destruction and reduction in native mammals • 2000: still present, local problems Carrying capacity Rabbits exceeded k Rabbit-proof fence No rabbits Carrying capacity Carrying capacity (k): the number of organisms that can be supported by a given area; the actual number of organisms fluctuates near this # of Animals (N) k time Adding A Limit to Population Growth More Realistic than Exponential Growth Growth is adjusted as population approaches carrying capacity (K) of the environment Population growth simply stops at K Population crashes after resource is consumed Population growth is under negative feedback as it approaches K and gradually reaches K Population Growth is Gradually Reduced as Carrying Capacity is Reached; Resources Renew Independently of Population Size Logistic Growth Inflection Point #s K simple favorite in wildlife management Rate of Increase is only a function of Population Density Assumes resources are not damaged by large populations Time Wildebeast don’t affect grass roots Logistic Math Verhulst (1838) and Pearl & Reed (1920) independently derived equation Verhulst-Pearl Equation (Sigmoidal Growth) dN/dt = derivative form of change in N with respect to time dN/dt = rmN(1-N/K) dN/dt = rmN = exponential growth As N approaches K, N/K approaches 1. Therefore rmN(1N/K) approaches 0 With K and Typical Seasonal Patterns of Reproduction, There is Often A “Doomed Surplus” Good Sites Poor Sites High Pops Low Pops Deep Water (Errington 1946) Dry Upland Mink control distribution of muskrats those in poor sites including dispersers are eaten Predators often take the young, homeless, sick, injured, dispersing, or old individuals so effect on species or community is less Logistic Growth Model May be Used to Calculate Harvest K #s Maximum Yield =1/2 K Time Maximum Sustainable Yield is at Inflection Point Growth is Maximum and Population is at Largest Size Larger Populations Start to Have Slower Growth Another View of Logistic Growth Inflection Point Max Sustainable Yield Growth rate starts slow, peaks, and ends slow dN/dt K N Maximum Sustainable Yield is at rate of fastest population growth Assumptions of Logistic Growth All individuals contribute equally to population growth--equal reproduction regardless of age or sex Growth rate is constant regardless of environmental variation K is constant--not affected by growth Reduction in growth as population approaches K is linear and instantaneous (no time lags) Populations fluctuate due to Density dependent factors Ex: Predation, competition, habitat availability change population growth in predictable ways N is driven by population density Density independent factors Random or Stochastic events Ex. Weather, accidents Breeding 14 aug 2007 Reindeer (caribou) # young produced Bighorn sheep Population density (top) or size (bottom) Population regulation: food High food addition Low food addition Townsend’s vole No food added Shaded area is winter Population regulation: food Population cycles: Ex. peaks in lynx populations show time lag behind peaks in snowshoe hare populations Population size Snowshoe hare Lynx Time (years) Population regulation: climate Population regulation: competition Competition – demand by 2 or more individuals of the same or different species for a common resource Between 2 individuals of same species: Intraspecific Between 2 individuals of different species: Interspecific Limited supply of resource: Exploitation Not limited but interaction detrimental: Interference Inter- or Intraspecific competition? Exploitation or Interference competition? Population regulation: competition