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Population
Re gulation
Chapter 5
Preconditions…
 Populations change over time
 Populations cannot grow indefinitely
 Logistic curve
 Logistic equation represents equilibrium view
  of population regulation (if perturbed,
  population returns to equilibrium value, K)
 Other views see population fluctuations as
  random over time, without returning to
  equilibrium (due to disturbance)
Background
 Population regulation: fluctuations in
  abundance with feedback mechanisms to
  increase or decrease density toward K
 Population control: ecological mechanisms
  which control upper limit of density
 Density is a result of combination of factors
 In general: ΔN = (b + i) – (d + e), where N is
  population size, b is births, d is deaths, i is
  immigrants, e is emigrants
Patter ns of
Population
Fluctuation
Small-magnitude irregular
fluctuations, Large-scale irregular
fluctuations, Cycles, Irruptions
Small-magnitude irregular
 fluctuations
 Small random
  changes in density
  of one order of
  magnitude or less
Large-scale irregular
 fluctuations

 Large random
  changes in
  density of
  several orders
  of magnitude
Cycles

  Regular interval changes in population
   density
Irruptions

  Occasional, unpredictable population
   explosions
Equilibrium Theories
 Central difference among theories lies in the
  relative importance of density-dependent
  factors and density-independent factors.
 Density-dependent factors have an
  increasing effect with increasing density
 Density-independent factors have an effect
  that does not vary with density
Extrinsic Biotic School
 Accepts importance of density-dependent
  factors
 Emphasizes external biotic factors

   Food supply
   Predation
   Disease
Food supply
 Evidence shows that food-supply is a strong
  determinant of density.
 Birds frequently die of starvation.
 Areas with high food supplies tend to have
  high bird densities. (correlation Vs.
  causation)
 Artificially supplemented food studies
 Naturally supplemented food studies
Predation
 Difficult to establish (need to know density
  differences of predators with varying prey
  densities)
 Studies indicate that predator species
  depress prey populations
 Removal experiments yield ambiguous
  results

 “Top-down” or “bottom-up” controversy
Disease and parasitism
 Increased densities may increase the rate of
  transmission
 Increased density frequently correlates with
  increased disease rate
 However, correlation may not indicate
  causation (food supply, red grouse)
The Intrinsic School
 Based on mechanisms intrinsic to the
  population
 Aka the population is self-regulated
 Also relies on density-dependence

 Stress, territoriality, genetic polymorphism
  hypothesis, dispersal
Stress, Territoriality
 Stress may regulate density by causing
  physiological reactions to high densities

 Territoriality may regulate density by
  excluding some individuals from reproducing
Genetic Polymorphism
 Hypothesis, dispersal

 Genetic composition changes in response to
  density

 Saturation dispersal, presaturation dispersal
  (reduces inbreeding)
Nonequilibrium
theories of
population
re gulation
Abiotic Extrinsic Regulation,
Metapopulations, Chaos theory
Abiotic Extrinsic Regulation
 Density-independent, abiotic factors
 Weather, temperature, moisture, sun-
  exposure, rainfall, etc…
 These factors are sufficient to explain density
  variations. Populations do not encounter
  ideal conditions long enough for density-
  dependent factors to be of importance.
Metapopulations
 Population consisting of several patches of
  populations linked by dispersal.
 Patches vary, may go extinct; not in
  equilibrium, but overall population survives
  due to dispersal among patches
 Metapopulations are particularly important in
  fragmented habitats
Chaos Theory
 Unpredictable patterns of population growth
 Particularly interesting with r values above
  2.69
 Pattern depends on initial conditions
 Not stochastic
 Property of the growth itself (growth
  equation)
Recapitulating Population
 Regulation
 There are equilibrium and non-equilibrium
  populations
 Density-dependent and density-independent
  factors affect populations (biotic and abiotic
  factors)
 It is undeniable that there is no single
  explanation: rather, a combination of theories
  applies. To what extent in each case is the
  relative contribution becomes the question.
Invasions
 Four stages: Transport, Introduction,
  Establishment, Spread
 Invasions follow the logistic curve, usually
  with longer lag phase, followed by
  exponential growth
 Invasions reach high densities (e.g. zebra
  mussels, Opuntia cactus and cactoblastis
  moth)
 Escape from density-dependent factors?
  Probably not. Other possibilities.
 Zebra mussel figure
Anywhere, everywhere!
Extinction and Risk Analysis
 Extinction is a natural component of
  populations (strongly aggravated by humans)
 Birth rate decreases, mortality increases
 Very low populations suffer the Allee effect
 Anthropogenic habitat loss creates three risk
  factors: demographic accidents, habitat
  fragmentation, genetic risk
Demographic accidents
 Habitat loss creates population decrease
 With smaller populations, risk of extinction
  increases, due to demographic accidents
 Chance events have a greater impact on
  small populations
 Severe winter, epidemic, predators, etc…
Habitat fragmentation
 Habitat loss frequently leads to habitat
  fragmentation
 This leads to a metapopulation structure
 Single patches may not be large enough to
  support a breeding population
 Dispersal may not be possible to support
  supplying of extinct patches
 Patches may go extinct simultaneously
Genetic risks
 Smaller populations have increased
  inbreeding and genetic drift
 Both lead to increased homozygosity
  (bottlenecking effect leads to loss of alleles)
 Increased homozygosity decreases fitness,
  and thus places population at risk
Heath hen on Martha’s
 Vineyard
 Overhunting caused massive population
  decline until 1907
 Population increased moderately thereafter
  (genetic risks?)
 In 1916, fire, storm, cold winter, invasion
  reduced population to 50 pairs (demographic
  accidents-more genetic risk)
 Subsequent years showed sex-ratio skewed
  toward males (demographic accident)
 Extinct by 1932 (any habitat fragmentation?)
T he end.

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Ch. 5 population regulation part

  • 2. Preconditions…  Populations change over time  Populations cannot grow indefinitely  Logistic curve  Logistic equation represents equilibrium view of population regulation (if perturbed, population returns to equilibrium value, K)  Other views see population fluctuations as random over time, without returning to equilibrium (due to disturbance)
  • 3. Background  Population regulation: fluctuations in abundance with feedback mechanisms to increase or decrease density toward K  Population control: ecological mechanisms which control upper limit of density  Density is a result of combination of factors  In general: ΔN = (b + i) – (d + e), where N is population size, b is births, d is deaths, i is immigrants, e is emigrants
  • 4. Patter ns of Population Fluctuation Small-magnitude irregular fluctuations, Large-scale irregular fluctuations, Cycles, Irruptions
  • 5. Small-magnitude irregular fluctuations  Small random changes in density of one order of magnitude or less
  • 6. Large-scale irregular fluctuations  Large random changes in density of several orders of magnitude
  • 7. Cycles  Regular interval changes in population density
  • 8. Irruptions  Occasional, unpredictable population explosions
  • 9. Equilibrium Theories  Central difference among theories lies in the relative importance of density-dependent factors and density-independent factors.  Density-dependent factors have an increasing effect with increasing density  Density-independent factors have an effect that does not vary with density
  • 10.
  • 11. Extrinsic Biotic School  Accepts importance of density-dependent factors  Emphasizes external biotic factors  Food supply  Predation  Disease
  • 12. Food supply  Evidence shows that food-supply is a strong determinant of density.  Birds frequently die of starvation.  Areas with high food supplies tend to have high bird densities. (correlation Vs. causation)  Artificially supplemented food studies  Naturally supplemented food studies
  • 13. Predation  Difficult to establish (need to know density differences of predators with varying prey densities)  Studies indicate that predator species depress prey populations  Removal experiments yield ambiguous results  “Top-down” or “bottom-up” controversy
  • 14. Disease and parasitism  Increased densities may increase the rate of transmission  Increased density frequently correlates with increased disease rate  However, correlation may not indicate causation (food supply, red grouse)
  • 15. The Intrinsic School  Based on mechanisms intrinsic to the population  Aka the population is self-regulated  Also relies on density-dependence  Stress, territoriality, genetic polymorphism hypothesis, dispersal
  • 16. Stress, Territoriality  Stress may regulate density by causing physiological reactions to high densities  Territoriality may regulate density by excluding some individuals from reproducing
  • 17. Genetic Polymorphism Hypothesis, dispersal  Genetic composition changes in response to density  Saturation dispersal, presaturation dispersal (reduces inbreeding)
  • 18. Nonequilibrium theories of population re gulation Abiotic Extrinsic Regulation, Metapopulations, Chaos theory
  • 19. Abiotic Extrinsic Regulation  Density-independent, abiotic factors  Weather, temperature, moisture, sun- exposure, rainfall, etc…  These factors are sufficient to explain density variations. Populations do not encounter ideal conditions long enough for density- dependent factors to be of importance.
  • 20. Metapopulations  Population consisting of several patches of populations linked by dispersal.  Patches vary, may go extinct; not in equilibrium, but overall population survives due to dispersal among patches  Metapopulations are particularly important in fragmented habitats
  • 21.
  • 22. Chaos Theory  Unpredictable patterns of population growth  Particularly interesting with r values above 2.69  Pattern depends on initial conditions  Not stochastic  Property of the growth itself (growth equation)
  • 23.
  • 24. Recapitulating Population Regulation  There are equilibrium and non-equilibrium populations  Density-dependent and density-independent factors affect populations (biotic and abiotic factors)  It is undeniable that there is no single explanation: rather, a combination of theories applies. To what extent in each case is the relative contribution becomes the question.
  • 25. Invasions  Four stages: Transport, Introduction, Establishment, Spread  Invasions follow the logistic curve, usually with longer lag phase, followed by exponential growth  Invasions reach high densities (e.g. zebra mussels, Opuntia cactus and cactoblastis moth)  Escape from density-dependent factors? Probably not. Other possibilities.
  • 26.
  • 29.
  • 30. Extinction and Risk Analysis  Extinction is a natural component of populations (strongly aggravated by humans)  Birth rate decreases, mortality increases  Very low populations suffer the Allee effect  Anthropogenic habitat loss creates three risk factors: demographic accidents, habitat fragmentation, genetic risk
  • 31. Demographic accidents  Habitat loss creates population decrease  With smaller populations, risk of extinction increases, due to demographic accidents  Chance events have a greater impact on small populations  Severe winter, epidemic, predators, etc…
  • 32. Habitat fragmentation  Habitat loss frequently leads to habitat fragmentation  This leads to a metapopulation structure  Single patches may not be large enough to support a breeding population  Dispersal may not be possible to support supplying of extinct patches  Patches may go extinct simultaneously
  • 33. Genetic risks  Smaller populations have increased inbreeding and genetic drift  Both lead to increased homozygosity (bottlenecking effect leads to loss of alleles)  Increased homozygosity decreases fitness, and thus places population at risk
  • 34. Heath hen on Martha’s Vineyard  Overhunting caused massive population decline until 1907  Population increased moderately thereafter (genetic risks?)  In 1916, fire, storm, cold winter, invasion reduced population to 50 pairs (demographic accidents-more genetic risk)  Subsequent years showed sex-ratio skewed toward males (demographic accident)  Extinct by 1932 (any habitat fragmentation?)

Editor's Notes

  1. D-shaped, gets its name from the stripes, usually around 2 cm, up to 3 cm.