Habitat loss and fragmentation I Bio 415/615 Questions 1. What does FST measure? 2. How does FST relate to fire management and collared lizards in the Ozarks? 3. How are genetic drift and selection different? 4. Why is FST similar to beta diversity? Fragmentation & Genetics • Ne decreases and distance increases • Instantaneous loss on sampling • Fast loss with reduced gene flow (sink genes) • Slower loss with random genetic drift, decreasing allelic diversity (rare alleles) within populations and causing populations to diverge (depends on time at low Ne and thus on generation time) • Inbreeding increases Processes of species loss and allele loss are similar Number of alleles Instantaneous Fast Slow Genetic drift Extinction debt Fragmentation and gene flow • How do genes move between populations? – Movement of animals – Dispersal of plants (seeds, spores) – Movement of sperm (pollen) • Is gene flow equal to colonization rate? – Individuals may die before reproducing (e.g., territoriality in animals) – Some insects invade areas where they don’t reproduce… Ecological dispersal is not equal to genetic dispersal Fragmentation and gene flow • How do we estimate the effects of fragmentation on gene flow? Fst describes the proportion of genetic variance in the whole population (all patches) attributed to differences between patches. Eg: if there are 10 alleles at a locus in a population of 10 subpopulations, and each has a different allele, Fst = 1 Fragmentation and gene flow • Fst is defined more formally as the variance in allele frequencies between populations (standardized by the mean) It is also estimated as: Fst ~ 1 / (4Ne*m + 1) Migration rate (of alleles): gene flow Effective population size (why important?) As gene flow goes up, Fst goes down; as local population size decreases, Fst goes up Consider similarity to alpha and beta diversity gene flow genetic drift Population 1 Gene Flow ↓ FST gene flow Population 2 Local adaptation ↑ FST genetic drift Population 3 gene flow Drift ↑ FST genetic drift Consider similarity to alpha and beta diversity β diversity α diversity Colonization Rate ↓ β diversity Population 1 β diversity Population 2 α diversity Population 3 α diversity β diversity Isolation ↑ β diversity Endangered Rutidosis (daisy) in Australia: genetic diversity and Ne Young et al. 1999 Con. Bio. 13:256-265. Rare alleles are those lost when Ne is small; heterozygosity was not related to Ne… so small populations do not appear to suffer inbreeding depression % loci with >1 allele allozymes High Gene Flow: Fst=.17 Case study: Missouri glades Eastern collared lizard Fragmentation Natural burn regime: every 5 years Fire suppression since 1950: glades are disappearing, juniper expanding Why do these lizards inhabit these ‘desert’like environments? Fragmentation Q: What did forest regrowth do to the lizard population? A: Some populations were lost, the lizards stopped moving (how do they know?), and Fst (.40) indicated major genetic drift. Q: Why do they think the high Fst was the result of drift rather than selection (local adaptation)? A: Genetic similarity was not related to spatial distance (why does this matter?). Q: What was the management response? A: BURN http://www.youtube.com/watch?v=N01BygHu-T0 Response to management The forest opens up Are they moving? Conclusion Q: What happened to the genetic structure? A: Eventually Fst did go down (not reported here). Q: How do the authors respond to the suggestion that population isolation is ‘good’ because it eventually produces new species (‘shifting balance theory’)? A: It assumes that population size will again grow. Should we expect this from fragmentation? Q: What, the authors conclude, should management focus on? A: Maintain the ecological processes (eg burn regime).