increasing vulnerability in small...
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Increasing Vulnerability in Small Populations
Brook Milligan
Department of BiologyNew Mexico State University
Las Cruces, New Mexico [email protected]
Fall 2009
Brook Milligan Increasing Vulnerability in Small Populations
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Designation versus protection
Recognition of species at risk
9 IUCN categoriescan lead to official designation, e.g., Federally-listedendangered species, which conveys legal status
Biological status
unchanged by designationmay continue to deterioratemitigation of detrimental factors required in addition todesignation
Brook Milligan Increasing Vulnerability in Small Populations
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Core Concepts
Core concepts
Minimum viable population size (MVP)Minimum dynamic area (MDA)
Populations below a certain threshold exhibit much higherprobability of extinction, while those above are much morelikely to persist
Brook Milligan Increasing Vulnerability in Small Populations
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MVP: Bighorn Sheep
Brook Milligan Increasing Vulnerability in Small Populations
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MVP: Channel Island Birds
Brook Milligan Increasing Vulnerability in Small Populations
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MVP: Ipomopsis aggregata
Brook Milligan Increasing Vulnerability in Small Populations
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Which Factors Cause Extinction?Heath hen: Tympanuchus cupido cupido
Once fairly common from New England to Virginia
Declined steadily with European settlement
1876: remained only on Martha’s Vineyard
1900: fewer than 100 survivors
1907: refuge on Martha’s Vineyard and predator control
1916: increase to over 800 birds
1916: fire destroyed most nests and habitat
1916 winter: high predation by goshawks (Accipiter gentilis)
1917: reduced population to 100–150 individuals
1920: 200 individuals, but disease reduced population tobelow 100
1920s: increasingly sterile, male skewed sex ratio
1932: extinct
Brook Milligan Increasing Vulnerability in Small Populations
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General Use of MVP and MDA
Result: quantitative assessment of risk in face of uncertaintyand stochasticity as a function of population size
Analogous to flood prediction
Framework for evaluating alternative management options
Factors leading to population decline and extinction (Shaffer,1981)
demographic fluctuationenvironmental fluctuationloss of genetic variability
Brook Milligan Increasing Vulnerability in Small Populations
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MVP and Types of Stochasticity
Brook Milligan Increasing Vulnerability in Small Populations
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Demographic Stochasticity
Variation among individuals in survival or reproductive success
“Arises from chance events in the survival and reproductivesuccess of a finite number of individuals” (Shaffer, 1981)
Measurable as the variance in fitness among individualscompared with the mean of the population
Brook Milligan Increasing Vulnerability in Small Populations
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Environmental Stochasticity and Natural Catastrophes
Temporal variation in survival or reproductive successaffecting entire population
Two typesenvironmental stochasticity
“due to temporal variation of habitat parameters and thepopulations of competitors, predators, parasites, and diseases”(Shaffer, 1981)measurable as the variance through time in the populationmean fitness
natural catastrophes
“such as floods, fires, droughts, etc., which may occur atrandom intervals through time” (Shaffer, 1981)measurable as the variance through time in mean acrosspopulations
Brook Milligan Increasing Vulnerability in Small Populations
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Loss of Genetic Variability
Inbreeding: increase in common ancestry
Heterozygosity: decline in variation
Effective population size: Ne
Inbreeding depression: decrease in fitness
Brook Milligan Increasing Vulnerability in Small Populations
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Loss of Genetic Variability: Inbreeding
Increase of homozygosity or identity by descent
F (t + 1) = F (t)(1) + (1− F (t))
(1
2Ne
)(1)
∆F (t) = (1− F (t))
(1
2Ne
)(2)
Brook Milligan Increasing Vulnerability in Small Populations
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Loss of Genetic Variability: Heterozygosity
Loss of heterozygosity
H(t + 1) =
(1− 1
2Ne
)H(t)
= λH(t) (3)
∆H = H(t + 1)− H(t)
=
(1− 1
2Ne
)H(t)− H(t)
= − 1
2Ne(4)
H(t) =
(1− 1
2Ne
)t
H(0)
= λtH(0)
≈ H(0) exp
(− t
2Ne
)(5)
Brook Milligan Increasing Vulnerability in Small Populations
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Loss of Heterozygosity
Brook Milligan Increasing Vulnerability in Small Populations
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Loss of Genetic Variability: Effective Population Size
Effective population size: Ne
unequal sex ratio
Ne =4NmNf
Nm + Nf(6)
variation in reproductive outputpopulation fluctuation
1
Ne=
1
t
t∑i=1
1
Ni(7)
1
Ne=
1
5
(1
10+
1
20+
1
100+
1
20+
1
10
)=
1
5· 31
100
=1
16.1(8)
Brook Milligan Increasing Vulnerability in Small Populations
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Effective Population Size
Brook Milligan Increasing Vulnerability in Small Populations
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Loss of Genetic Variability: Inbreeding Depression
Inbreeding leads to greater homozygosity
Increased homozygosity exposes more recessive deleteriousalleles
Greater expression of recessive deleterious alleles leads toreduction in fitness
Brook Milligan Increasing Vulnerability in Small Populations
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Inbreeding Depression
Brook Milligan Increasing Vulnerability in Small Populations
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Inbreeding Depression
Brook Milligan Increasing Vulnerability in Small Populations
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Detrimental Positive Feedback: Extinction Vortices
Any reduction in population size increases probability offurther reductions
increases demographic stochasticityincreases vulnerability to environmental stochasticityincreases loss of genetic variation
Example: mutational meltdown
fixation of deleterious allelesreduction in population growth rateincreased risk of fixation of deleterious alleles
Brook Milligan Increasing Vulnerability in Small Populations
References
Shaffer, M. L. 1981. Minimum population sizes for speciesconservation. BioScience, 31:131–134.
Brook Milligan Increasing Vulnerability in Small Populations