I Biodiversity Definitions and Assessment A Definitions 2

I. Biodiversity – Definitions and Assessment A. Definitions 2. Components of Biodiversity • a. b. c. The term “biodiversity” often is used incorrectly or incompletely • Not synonymous with “species diversity” • Encompasses three measures Species Diversity 1) Species richness – Total number of species • Often cited incorrectly as “biodiversity” • Fairly simple to estimate from rarefaction curves 2) Evenness – Proportions of species in a community • More difficult to determine (requires more complete survey) Genetic Diversity – Variety of genotypes Ecosystem Diversity – Variety of habitat types

I. Biodiversity – Definitions and Assessment A. Definitions 2. Components of Biodiversity • a. b. c. The term “biodiversity” often is used incorrectly or incompletely • Not synonymous with “species diversity” • Encompasses three measures Species Diversity 1) Species richness – Total number of species • Often cited incorrectly as “biodiversity” • Fairly simple to estimate from rarefaction curves 2) Evenness – Proportions of species in a community • More difficult to determine (requires more complete survey) Genetic Diversity – Variety of genotypes Keystone Species

I. Biodiversity – Definitions and Assessment B. Estimates of Biodiversity • Described species ~ 1. 8 million • • • Insects > 1, 000 species Plants > 290, 000 species Probably an underestimate • Only ~5000 species of bacteria Less conspicuous species studied less often Estimates range from 5 – 30 million • • • Around 300 new species described each day Average estimate ~ 17. 5 million Splitting of taxa more common than lumping • • Tendency to increase number of described species Cryptic species

I. Biodiversity – Definitions and Assessment C. Estimates of Extinction Rates • Geological history • Periods of extinction followed by periods of rapid speciation (every ~ 26 million years) How do we estimate rates of extinction? ? • 1. Problems a. b. c. d. Difficult to know when a species is extinct • Ex – Coelacanth, ivory billed woodpecker, giant lemur Species distributed unevenly (patchy distribution) • Species affected unevenly by habitat loss Extinctions may not happen immediately • Short-lived species show effects rapidly • Long-lived species may appear to be unaffected for long periods of time • “Biologically extinct” – Populations not self-sustaining • “Living dead” - Janzen Uncertainty about number of species in an area • Wilson – “No precise estimate can be made of the numbers of species being extinguished in the rain forests or in other major habitats, for the simple reason that we do not know the numbers of species originally present”

I. Biodiversity – Definitions and Assessment C. Estimates of Extinction Rates 2. Estimation Methods • a. b. c. d. • • Area-species relationship (Mac. Arthur & Wilson) Estimate biodiversity for a small area Extrapolate estimate to area of habitat • Species ~ Area 0. 25 (0. 15 -0. 35) • Increase area 10 X Increase species 2 X Estimate rate at which ecosystem area is being reduced Calculate extinction rate based on predicted reduction in species richness from reduction in habitat area Current estimate ~ 17, 500 species year-1 • 1 out of every 1000 species on Earth each year “Background” rate from fossil record • 1 out of every 1 -10 million species on Earth each year

I. Biodiversity – Definitions and Assessment C. Estimates of Extinction Rates 2. Estimation Methods • a. b. c. d. • • Area-species relationship (Mac. Arthur & Wilson) Estimate biodiversity for a small area Extrapolate estimate to area of habitat • Species ~ Area 0. 25 (0. 15 -0. 35) • Increase area 10 X Increase species 2 X Estimate rate at which ecosystem area is being reduced Calculate extinction rate based on predicted reduction in species richness from reduction in habitat area Current estimates ~ 17, 500 species year-1 • 1 out of every 1000 species on Earth each year • Myers – 40, 000 year-1 • Lomborg – 1033 documented from 1600 – 1998 • The Skeptical Environmentalist “Background” rate from fossil record • 1 out of every 1 -10 million species year-1

I. Biodiversity – Definitions and Assessment C. Estimates of Extinction Rates • Point: Estimates may be unreliable and thus invalid • • No action should be taken until biodiversity loss is demonstrated and shown to be harmful Counterpoint: Wilson – Projections using areaspecies relationships in tropical settings (where most of biodiversity loss currently is happening) are conservative • • Tropical species have localized distributions that make them especially vulnerable to habitat loss Damaging loss of genetic diversity may occur, even if outright extinction of a species doesn’t happen

I. Biodiversity – Definitions and Assessment D. Biodiversity Hotspots • Myers – Up to 20% of the world’s plant species and more than 20% of the animal species are confined to 0. 5% of the land surface Biodiversity Hotspot – Area with high degree of • • • Biodiversity Endemism Risk of habitat degradation/loss Concept originally intended for tropical and subtropical areas • Endemism less prevalent in temperate and polar regions

II. Biodiversity – Factors A. Nutrient Availability 1. Oligotrophic • • 2. Dominated by a few species able to survive on limited nutrients Low diversity, Low biomass Mesotrophic • • 3. Support greater numbers of species • Rapid colonizers held in check by nutrient limitation • Less aggressive species capable of surviving High diversity, Medium biomass Eutrophic • • Dominated by a few species able to grow and/or colonize rapidly with abundant nutrients Low diversity, High biomass

II. Biodiversity – Factors B. Selective Colonization/Mortality 1. Colonization • 2. Excellent colonizers (r-selected) may dominate newly available habitats Mortality • • Predation • Ex – Birds with colorful plumage • Ex – Sea urchins (sushi) Species-specific diseases/pests • Ex – Dutch elm disease • Ex – Western bark beetles

II. Biodiversity – Factors C. Habitat Disturbance • Non-selective habitat disturbance has potential to increase diversity • • Prevents competitive exclusion Intermediate disturbance Maximum diversity

II. Biodiversity – Factors C. Habitat Disturbance • Fire and fire-dependent species • • • Ex – Peter’s Mountain Mallow (Iliamna corei) Discovered in 1927 (50 plants) Endemic to meadow in western Virginia 1986 - Three plants remaining • Not setting seed • Listed as endangered Research on seeds indicated importance of fire • Cracks hard seed coat, aiding germination • Removes competing vegetation • Had been suppressed in the area Controlled burns in 1992 and 1993 led to appearance of 500+ seedlings

II. Biodiversity – Factors D. Habitat Fragmentation/Destruction • • • Most significant factor causing species loss Smaller habitats support fewer species and smaller populations than large habitats Population sizes tend to fluctuate more in smaller habitats than large habitats Reduced population Lower genetic diversity Behavior of territorial species changes in fragments, esp. when territory size ~ fragment size Fragments may not support self-sustaining populations (rely on immigration from outside)

• Mount Hood National Forest, Oregon • Patches due to timber removal

II. Biodiversity – Factors D. Habitat Fragmentation/Destruction • • • Most significant factor causing species loss Smaller habitats support fewer species and smaller populations than large habitats Population sizes tend to fluctuate more in smaller habitats than large habitats Reduced population Lower genetic diversity Behavior of territorial species changes in fragments, esp. when territory size ~ fragment size Fragments may not support self-sustaining populations (rely on immigration from outside)

II. Biodiversity – Factors D. Habitat Fragmentation/Destruction • Fragmentation increases edge effects • • • Positive effects • Increased light to plant species at edges Negative effects • Increased predation by animals foraging at habitat edge Ex – Nesting success among migratory birds in Midwestern forests lower in fragments due to increased nest predation and parasitism by cowbirds

II. Biodiversity – Factors E. Exotic Species • Species invasions may profoundly affect ecosystems Detrimental exotic species usually are • • • Superior competitors • Ex – Argentine ants, starlings, zebra mussels Effective predators • Ex – Nile perch, mongeese

II. Biodiversity – Factors E. Exotic Species 1. Zebra mussel • • Competitor in Great Lakes and elsewhere Transported from Europe in ballast water Fouling organism • Restricts movement of water through intake pipes • Colonizes boat hulls, pier pilings, buoys, etc. • Fouls other organisms (clams, mussels) Filter feeder – removes larvae and particulate material • Outcompetes native shellfish species for food and space • Removes larvae from water

II. Biodiversity – Factors E. Exotic Species 2. Mongoose • • • 3. Predator in Hawaii Introduced in 1883 to combat rat population Prey on native birds Lionfish • • Venomous predator Introduced in Caribbean/W Atlantic ca. early/mid 1990’s Preys on 65+ spp. of fishes No natural predators

Nile perch – Lake Victoria Argentine ants - California Brown tree snake - Guam Caulerpa taxifolia - California

III. Biodiversity – Value A. Value to Humans • Economic • • Ex – Lomborg: $3 -33 trillion annually Biodiversity loss could lead to removal of species that benefit humans but aren’t currently known to do so • • Ex – Chapin et al. suggest increased frequency of Lyme disease in 20 th century may have been related to increase in abundance of tick-bearing mice (once controlled by food competition with passenger pigeons) Species extinction reduces potential pool of species containing chemical compounds with pharmaceutical or industrial applications • Counter – Many pharmaceutical companies now use directed design to search for new drugs

III. Biodiversity – Value A. Value to Humans • Problem – Benefits may not be obvious • • • Difficult to convince people that it’s important to preserve something with no immediately apparent intrinsic value to them (charisma? ) Ex – Economic value of viral resistance added to commercial strains of perennial corn through hybridization with teosinte (Mexican wild grass) is ~ $230 -300 million Ex – Weedy tomatoes from Peru Discovered in 1962 during search for potatoes Seeds sent to researcher at UC Davis who used plants to breed with other tomatoes In 1980 after nearly 10 generations of crossing and backcrossing, new strains were produced with larger fruit, improved pigmentation and increased concentrations of sugars and soluble solids

III. Biodiversity – Value B. Ecosystem Value • 1. Biodiversity can have large effects on ecosystem stability and productivity Benefits of biodiversity a. b. Productivity • Halving species richness reduces productivity by 10 -20% (Tilman) • Average plot with one plant species is less than half as productive as a plot with 24 -32 species • Question – Can these results be extrapolated to other systems and time/space scales? Nutrient retention • Loss of nutrients through leaching is reduced when diversity is high • Caveat – Studies to date have focused on low diversity communities (Why? ); can those results be generalized?

III. Biodiversity – Value B. Ecosystem Value 1. Benefits of biodiversity c. • • Ecosystem stability Mechanism • Multiple species within a trophic level compete for resources • If abundance of one species declines due to perturbation, competing species may increase in abundance • Individual species abundances may vary, but community as a whole is more stable with more species Consequences • High diversity doesn’t guarantee that individual populations won’t fluctuate • Ex – Higher diversity (unfertilized) plots of native plant species maintained more biomass during drought than lower diversity (fertilized) plots • High diversity may confer greater resistance to pests and diseases • Ex – Higher diversity plots of native plant species had greater resistance to fungal diseases, reduced predation by herbivorous insects and reduced invasion by weeds

III. Biodiversity – Value B. Ecosystem Value 2. Considerations a. • b. • • • Species richness vs. Species evenness Simple species richness may be deceptive as an indicator of biodiversity and ecosystem stability • Evenness usually responds more rapidly to perturbation than richness and may have important ecosystem consequences • Richness is typical focus of studies and policy decisions Importance of individual species Charismatic megafauna: What about non-charismatic species? Different species affect ecosystems in different ways (keystone species vs. non-keystone species) • Ex – Sea otters/Sea urchins/Kelp forests in eastern Pacific Ocean Question: How many species are required to maintain “normal” ecosystem function and stability? • No magic number • Losing one ant species in a tropical forest may have less immediate impact than losing one species of fungus that is crucial to nutrient cycling in the soil

IV. Biodiversity – Management • Strategies outlined in Convention on Biological Diversity • • • Developed between 1988 and 1992 Opened for ratification at UN Conference on Environment and Development (Rio “Earth Summit”) Ratified by 168 nations; went into force in Dec 1992 Objectives – “…the conservation of biological diversity, the sustainable use of its components and the fair and equitable sharing of the benefits arising out of the utilization of genetic resources…” Articles 8 -9 specify a combination of in situ and ex situ conservation measures • • Primary use of in situ conservation Use of ex situ measures as a complement