Stano Pekár“Populační ekologie živočichů“ dN = Nr dt a major sub-field of ecology which deals with description and the dynamics of populations within species, and the interactions of populations with environmental factors expanding field (Price & Hunter 1995): - populations 52 %, communities 9 %, ecosystems 10 % main focus on - Demography = description of populations that gave rise to Life-history theory - Population dynamics = describe the change in the numbers of individuals in a population populations of member species may show a range of dynamic patterns in time and space central question: “WHAT DOES REGULATE POPULATIONS?“ Change in abundance of Lynx and Lepus in Canada density independent factors, food supply, intraspecific competition, interspecific competition, predators, parasites, diseases 1. Conservation biology World Conservation Union (IUCN) uses several criterions (population size, generation length, population decline, fragmentation, fluctuation) to assess species status by means of Population viability analysis (PVA) estimates the extinction probability of a taxon based on known life history, habitat requirements, threats and any specified management options Saiga tatarica critical: 50% probability of extinction within 5 years endangered: 20% probability of extinction within 20 years vulnerable: 10% probability of extinction within 100 years 2. Biological control to assess ability of a natural enemy to control a pest in 1880 Icerya purchasi was causing infestations so severe in California citrus groves that growers were burning their trees in winter 1888-1889 Rodolia cardinalis and Cryptochaetum were introduced into California from Australia, growers took the initiative and applied the natural enemies themselves by fall 1889 the pest was completely controlled Rodolia cardinalis has been exported to many other parts of the world the interest of growers and the public in this project was due to its spectacular success: the pest itself was showy and its damage was obvious and critical; the destruction of the pest and the recovery of the trees was evident within months Rodolia cardinalis (Coccinellidae) eating Icerya purchasi (Hemiptera) 3. Epidemiology to predict the diffusion of a disease and to plan a vaccination phocine distemper virus was identified in 1988 and caused death of 18 000 common seals in Europe during 4 months the disease travelled from Denmark to the UK the population of common seals in the UK declined by about half 0 50 100 150 200 250 300 350 400 0 2 4 6 8 10 12 14 16 18 20 week no.ofdeadseals observed predicted Observed and predicted epidemic curves for virus in common seals in the UK Grenfell et al. (1992) 4. Harvesting to predict maximum sustainable harvest in fisheries and forestry but also used to regulate whale or elephant hunting when population is growing most rapidly (K/2) then part of population can be harvested without causing extinction Beddington (1979) Relationship between capture and fishing effort Panulirus cygnus Population + environment = population system population conditions resources enemies molecules → organels → cells → tissues → organs → organ systems → organisms → populations → communities → ecosystem → landscape → biosphere a group of organisms of the same species that occupies a particular area at the same time and is characterised by an average characteristic (e.g., mortality) characteristics: Individual → Population Stage structure Age structure Size structure Sex ratio Spatial distribution Developmental stage Age Size Sex Territorial behaviour Event – an identifiable change in a population Process – a series of identical events • rate of a process – number of events per unit time Natality (birth rate) Mortality (mortality rate) Growth (growth rate) Population increase (rate of increase) Consumption (consumption rate) Birth [inds] Death [inds] Increment [gram] Increment [number] Acquisition of food [gram] ProcessEvent inherent characteristics of the evironment (pH, salinity, temperature, moisture, wind speed, etc.) not modified by populations not consumed by population ⇒ no feedback mechanisms ⇒ do not regulate population size limit population size optimal suboptimal unfavourable reproduction growth survival performance conditions any entity whose quantity is reduced (food, space, water, minerals, oxygen, sun radiation, etc.) modified (reduced) by populations defended by individuals (interference competition) regulate population size non-renewable resources - space Renewable resources - regeneration centre outside the population system ⇒ no effect of the consumer (e.g., oxygen, water) - regeneration centre inside of the population system ⇒ influenced by the consumer (e.g., prey) competitors, predators, parasites, pathogens negative effect on the population top-down regulation of the population Absolute - number of individuals per unit area - number of individuals per unit of habitat (leaf, plant, host) - sieving, sweeping, extraction, etc. Relative - number of individuals - trapping, fishing, pooting Capture-recapture method Assumptions: - marked individuals are not affected and marks will not be lost - marked animals become mixed in the population - all individuals have same probability of capture - capture time must be short Closed population - population do not change over sampling period - no death, births, immigration, emigration Petersen-Lincoln estimator: N - number of individuals in population a - total number of marked individuals r - total number of recaptured marked individuals n - total number of individuals recaptured r an N = 3 2 )( r rnna − =νVariance: Open population - changes due to death, births, immigration, emigration - at least 3 sampling periods Stochastic Jolly-Seber method Ni - estimate of population on day ai - number of marked individuals on day i ni - total number of individuals captured on day i ri - sum of recaptured marked individuals on day i Zi - sum of marked individuals before day i Ri - sum of all marked individuals on day i i i ii i r R Za M += i ii i r nM N = where Stano Pekár“Populační ekologie živočichů“ dN = Nr dt aim: to simulate (predict) what can happen models are tested by comparison with observed dynamic realistic models - complex (many parameters), realistic, used to simulate real situations strategic models - simple (few parameters), unrealistic, used for understanding of model behaviour a model should be: 1. a satisfactory description of diverse systems 2. an aid to enlighten aspects of population dynamics 3. a system that can be incorporated into more complex models deterministic models - everything is predictable stochastic models - including random events discrete models: - time is composed of discrete intervals or measured in generations - used for populations with synchronised reproduction (annual species) - modelled by difference equations continuous models: - time is continual (very short intervals) thus change is instantaneous - used for populations with asynchronous and continuous overlapping reproduction - modelled by differential equations STABILITY stable equilibrium is a state (population density) to which a population will move after a perturbation stable equilibrium unstable equilibrium focus on rates of population processes number of cockroaches in a living room increases: - influx of cockroaches from adjoining rooms → immigration [I] - cockroaches were born → birth [B] number of cockroaches declines: - dispersal of cockroaches → emigration [E] - cockroaches died → death [D] population increases if I + B > E + D rate of increase is a summary of all events (I + B - E - D) growth models are based on B and D spatial models are based on I and E Blatta orientalisEDBINN tt −−++=+1 Assumptions: immigration and emigration are ignored all individuals are identical reproduction is asexual resources are infinite for population with discrete generations (annual reproduction) if births and deaths do not depend on population size exponential (geometric) growth Malthus (1834) realised that any species can potentially increase in numbers according to a geometric series N0 .. initial density b .. birth rate (per capita) d .. death rate (per capita) Discrete (difference) model λ1−= tt NN 11 −− −=∆ tt dNbNN 11 )( −− −=− ttt NdbNN 1)1( −−+= tt NdbN λ=−+ db1where λ < 1 .. population declines λ > 1 .. population increases λ = 1 .. population does not change time0 Nt t t NN λ0= λλλ 012 NNN == population number in generations t is equal to number of individuals is multiplied each time - the larger the population the larger the increase λ = finite growth-rate, per capita rate of growth λ = 1.23 .. 23% increase R ..average of finite growth rates t t tt i iR 1 21 1 1 )...( λλλλ =      = ∏= Comparison of discrete and continuous generations populations that are continuously reproducing when change in population number is permanent Nt time Continuous (differential) model t t NN λ0= )ln()ln()ln( 0 λtNNt += )ln()ln()ln( 0 λtNNt =− )ln( 1 λ= Ndt dN )ln(λN dt dN = Nr dt dN =)ln(λ=rif Solution of the differential equation: - analytical or numerical at each point it is possible to determine the rate of change by differentiation (slope of the tangent) when t is large approximated by the exponential function r < 0 .. population declines r > 0 .. population increases r = 0 .. population does not change time N r - intrinsic rate of natural increase, instantaneous per capita growth rate Nr dt dN = r Ndt dN = 1 ∫∫ = TT rdtdN N 00 1 )0()ln()ln( 0 −=− TrNNT rT N NT =        0 ln rt t eNN 0= rTT e N N = 0 t t NN λ0= rt t eNN 0= rtt e=λ )ln(λ=r r versus λ r is symmetric around 0, λ is not r = 0.5 ... λ = 1.65 r = -0.5 ... λ = 0.61 doubling time: time required for a population to double r t )2ln( = MODULARIZACE VÝUKY EVOLUČNÍ A EKOLOGICKÉ BIOLOGIE CZ.1.07/2.2.00/15.0204 S. Pekár podzim 2011 Cvičení z Populační ekologie Capture Day ni ai 1 2 3 ri 1 60 58 2 125 123 15 15 3 154 150 5 38 43 4 189 187 9 20 45 Ri 58 45 Zi+1 14 20 Recapture day Cockroaches were captured using traps with baits for 4 consecutive days. Each day the specimens were marked and released. 1. Estimate population density assuming closed population. 2. Estimate population density assuming open population. 3. Estimate average population size for both closed and open populations. R2 = 58, R3 = 45Z2 = 14, Z3 = 20 r2 = 15, r3 = 43 Population density of the true bugs Coreus marginatus was recorded for 10 years. Here are the densities: 160, 172, 188, 154, 176, 185, 168, 194, 170, 169 Does population increase or decrease? What is the average population growth (R)? Project population for another 10 years using R and N0 = 90. Simulate population growth for the next 20 years using observed finite-growth rates. Population density of the mite Acarus siro was recorded every 3 days during 28 days. The following densities were found: 165, 145, 139, 125, 105, 101, 88, 81, 73, 69 What is the intrinsic rate of increase (r) and what was the initial density ? How long it takes for a population to decrease to half size? Project population growth for another 5 weeks using estimated r and N0 = 69. What would be the estimated rate if you know the initial and final density?