The Sixth Extinction? Community effects on ecosystem processes CMM Chap The context: altered biodiversity. 2a. Loss of Global Biodiveristy:

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Community effects on ecosystem processes CMM Chap. 12 A.1. State factors and interactive controls: Species effects on interactive controls determine ecosystem consequences I. Introduction A. The context 1. ClORPT 2. Loss of biodiversity 3. Species invasions B. What is biodiversity? C. What is ecosystem functioning? II. Organism effects on ecosystem processes A. Community determinants B. Functional determinants: interactive controls III. Effects of diversity on ecosystem processes A. Magnitudes of processes B. Stability of processes C. Synthesis 1.3 Powerpoint modified from Chapin (http://www.faculty.uaf.edu/fffsc/) How will changing biodiversity affect ecosystem functioning? 2. The context: altered biodiversity 2a. Loss of Global Biodiveristy: Current extinction rates are 10-100 fold higher than past Chapin et al. 2000 12.1 The Sixth Extinction? 2b. Loss of Regional Biodiveristy: Problems can occur long before species go extinct. 2. Devonian 370 mya 5. Cretaceous 65 mya 1. Ordovician 440 mya 3. Permian 250 mya 4. Triassic 210 mya National Geographic 1999 National Geographic Magazine 2007 1

3. Introductions of exotic species Washington examples (to name just a few): Shifts in species can be difficult, expensive, or impossible to reverse. Cheatgrass (Bromus tectorum) (eastern WA) http://www.cabnr.unr.edu/cabnr/newsletter/fullstory.aspx?storyid=41 Spartina alterniflora (cordgrass) in Gray s Harbor http://baycenter.wordpress.com/2007/10/17/43/ European Green crab (Carcinus maenas) http://massbay.mit.edu/exoticspecies/exoticmaps/images/carcinus_big.jpg Reed canary grass, Whatcom County Not all introduced species become problems in their new habitat, but those that do can cause extensive ecological and economic damage. Introductions Establishment Invasions Noxious pests B.What is biodiversity? Genetic Species Functional groups Effect Response Community types Ecosystem arrangement on the landscape Salmon - Genetic diversity maintains local adaptation Functional group diversity: Need to understand how responses link to effects on processes. Risk of extinction of Chinook stocks http://www.pnl.gov/news/ Augerot 2005 Suding et al. 2008 2

Understanding interactions Cows are good?! Ecosystem distribution matters for some services Bay checkerspot Wetlands & riparian buffers: Flood control, Nutrient filtration Plantago erecta Invasive grasses L. Gonzalez photos 2005 D. Hooper photos 2005 http://extension.umd.edu/environment/index.cfm C. What is ecosystem functioning? Ecosystem properties (non-qualitative) (production, nutrient cycling, energy flux, etc.) Ecosystem goods ( Provisioning services ) directly marketable Ecosystem services benefit human endeavor, but harder to quantify in economic terms Regulating: pollination, pest control, water purification, maintenance of soil fertility, climate regulation, etc.; Cultural: recreational, educational, spiritual, etc. II. Organism effects on ecosystem processes A. Community context Impact of organisms on ecosystems depends on: B. Functional Effect Traits D. Species interactions Chapin et al. 2000 3

Species effects are an important interactive control Affect most ecosystem processes 1. Species effects on resource supply: Invasion of exotic N fixer augments N cycling Myrica faya is a N-fixing exotic tree in Hawaii Deep-rooted species access more resources Enhanced NPP Alters ecosystem structure and functioning Deep-rooted species access more water Augments water available to support NPP Native perennial bunch grasses of Calif. Have been replaced either by European annual grasses or by deep-rooted Eucalyptus trees that can access more water and nutrients Relative growth rate is key trait with strong ecosystem impacts Determines NPP Influences nutrient requirement and litter quality Plant species differ predictably in litter quality High-resource-adapted leaves decompose quickly Enhance nutrient cycling rates 4

Species effects on nitrogen mineralization Associated with differences in litter quality 2. Species effects on climate a. in tropical forests Climate changes predicted for the Amazon Basin if rain forests were replaced by pasture. 12.5 2. Species effects on climate b. in boreal forest 3. Species effects on disturbance a. Many animals are ecosystem engineers Soil disturbance (worms, gophers, ants) Landscape disturbance (beavers) Harvester ant mound http://www.cwp.mines.edu/~norm/photos/cbautumn04/images/cement%20creek%20beaver%20dam.jpg 3. Species effects on disturbance b. Plants influence disturbance Decrease: Stabilize soils (e.g., sand dunes) Increase: Fire probability Linked response and effect? Species effects on succession 5

Boreal conifer forests are born to burn After fire are replaced by less flammable deciduous forests (effect and response functional groups differ) Fire in California chaparral (effect and response groups the same) http://www.werc.usgs.gov/fire/ http://www.csulb.edu/depts/geography/jpegs/woodsfirefreq.jpg Fire in Hawaiian woodlands (effects of invasive grasses) - Schizachrium scoparium is an invasive grass that both promotes fire and resprouts quickly after burning. - Effect and response groups are the same positive feedback. - Native Hawaiian shrubs and trees are not adapted to frequent fire and are eliminated. III. Effects of species diversity A. Magnitudes of processes 1. Enhance efficiency of resource use 2. Primary productivity 3. Mechanisms B. Stability of ecosystem processes C. Synthesis 1. Efficiency of resource capture/retention - greater plant species richness led to lower pool sizes of available N - from greater plant uptake? 2. Responses of Primary Productivity to Changing Plant Diversity Tilman et al. 1996 Hector et al. 2001 yr) 12.8 Hooper & Vitousek 1997 Figure from Inchausti 2001 Hooper and Vitousek 1997 6

3. Mechanisms of diversity effects Complementarity - niche differentiation that leads to greater overall resource use in more diverse communities. Complementarity in plants with strong functional differences Light interception Rooting depth E L (Vandermeer 1990) (Gulmon et al. 1983) 3. Mechanisms of diversity effects Detecting complementarity Complementarity - niche differentiation that leads to greater overall resource use in more diverse communities. Sampling - statistical probability of choosing a highly productive species. Sampling for dominant Complementarity Tilman et al. 1997 ANPP (g/m2/yr) 300 250 200 150 100 50 0 Aboveground Productivity 1999 P L E N EP LP EL ELP ELPN y = 21.22x + 100.67 R 2 = 0.1815 p = 0.25 0 1 2 3 4 5 Number of Functional Groups B. Stability of ecosystem properties 1. Different kinds of stability a. Resistance b. Resilience 2. Mechanisms of diversity effects a. Compensation Process rate 12.2 Species richness 7

Response variation in species leads to less variation in ecosystems Many species with a continuum of trait differences can lead to compensation. Portfolio effect 12.9 There may be little compensation for species with very distinct functional roles (e.g., keystone species) Cottingham et al. 2001 12.2 B.3. Experimental tests of diversity effects on stability of ecosystem properties Microbial diversity decreases ecosystem variability 12.10 What else could cause this pattern? McGrady-Steed et al. 1997 Report Warns of Global Collapse of Fishing By CORNELIA DEAN Published: November 2, 2006 Correlated response and effect traits: large ecosystem change Extinction Collapse Land use change and pollinator loss: effects on pollination efficiency (Worm et al. 2006 Science) Suding et al. 2008 (data from Larsen et al. 2005) 8

C. Synthesis of diversity effects Figure from Field 1996 One year/one site Zone accessible to intensive management + site to site variation + year to year variation + different successional stages Without knowledge of exact functional roles Summary Differences in species functional traits can influence ecosystem properties (effect and response groups) Direct effects and effects on interactive controls (climate, disturbance, resource availability) Diversity effects - Magnitudes of processes - Stability of processes - Understanding mechanisms needs critical evaluation of alternative hypotheses. 9