10/24/2013 Bio122 F13: Lecture 9 (10/24/13): Communities Community Structure I. Communities (Ch 15) A. Defined B. Food webs II. Community structure A. Species diversity B. Measuring species diversity 1. Diversity indices 2. Rank abundance curves 3. Species accumulation curves C. Species composition III. Processes that shape community structure: Interactions A. Direct & Indirect effects 1. Types of indirect effects 2. Interaction strength 3. Dominant species 4. Ecosystem engineers 5. Keystone species IV. Community Succession(Ch 16) A. Succession & its drivers Functional diversity in coral reef herbivores: Impact upon ecosystem structure Michelle Paddack Predators Herbivores Resilience ? Reef Builders University of East Anglia, Norwich UK Simon Fraser University, BC Canada X NCORE Grazing on coral reefs Fishes Number of species Great Barrier Reef Caribbean Functional Group: Species that perform a similar role in ecosystem processes, irrespective of taxonomic affinities Bellwood et al. Nature 2004 • Macro-herbivores have high grazing rates – 100 % of daily algal production • Algal communities in lowbiomass, highly-productive states (algal turfs: filamentous, < 1 cm tall) • Macroalgal biomass historically negligible 1 10/24/2013 Functional Groups: Herbivores Scarids (Parrotfishes) 13 spp. Pomacentrids (Damselfishes) 6 spp. Urchins (Diadema) Acanthurids (Surgeonfishes) 3 spp. Are herbivorous fishes a single functional group? Kyphosids (Chubs) 1 sp. Acanthurus sp. Purcell & Bellwood 1993 Scarus spp. Sparisoma spp. Bellwood & Choat 1990 Goals 1. Define herbivore functional sub-groups 2. Assign species to sub-groups 3. Determine whether correlations with benthic variables differ among sub-groups Methods: Defining & designating Caribbean herbivore functional sub-groups (23 fish species) Field observations: Feeding behavior Food preferences Laboratory observations: Randall 1967 Gut contents: type of algae Gut contents: sediment 2 10/24/2013 bites minute-1 Grazing rate variation among Caribbean herbivorous fishes Photo by R. Steneck 40 35 30 25 20 15 10 5 0 1. Define herbivore functional sub-groups Feeding strategies & ecosystem function Diet Disturbance to benthos Impact on coral Browser (Grazer) Macroalgae Very little sediment in gut Low - Crop algal fronds Reduce shading, abrasion, overgrowth Scraper Epilithic algal turf & associated surface sediment/detritus High gut sediment content Moderate – Partial clearing of surface Facilitate growth & survival of CCA & corals Epilithic and endolithic Excavator (Bioeroder) algae & sediment/detritus High – Remove substrate, visible scar Clear space for CCA & coral settlement Eat coral recruits Gardener High/Patchy – modify algal community in territories No coral recruitment in territories High gut sediment content “Farmed” epilithic algae & associated sediment/detritus Low gut sediment Svir Saur Scroi Svet Abah 3. Examine sub-group correlations: Data Atlantic & Gulf Rapid Reef Assessment http://www.agrra.org/ Fish Counts: 30 x 2 m belt transect Min. 10 per site All Scarids (≥ 5 cm) All Acanthurids Microspathodon chrysurus Sized in 5 cm bins Excavators: Scarus vetula Sparisoma viride Algal cover: (original methodology) 25 x 25 cm quadrat 5 per transect Target algal-covered area % Cover Macroalgae Algal height (avg) # Coral Recruits (≤ 2cm) AGRRA data: Sub-country means Fore reef, 3 – 9 m Relative % Cover of Macroalgae Costa Rica removed (1 site) r2 = 0.278 p = 0.001 100 Rel % macroalgae 80 60 40 20 0 0 1 2 3 4 5 7 Total herbivore biomass (kg 100m-2) 100 80 60 40 20 6 r2 = 0.204 p = 0.006 80 60 40 20 0 0.0 0.5 1.0 1.5 2.0 2.5 3.0 3.5 Excavator biomass (kg 100m-2) 100 r2 = 0.117 p = 0.050 0 0.0 0.5 1.0 1.5 2.0 2.5 3.0 Browser biomass (kg 100m-2) 80 r2 = 0.006 p = 0.687 60 40 20 0 0.0 0.1 0.2 0.3 0.4 0.5 0.6 Gardner biomass (kg 100m-2) Rel % macroalgae 100 Rel % macroalgae AGRRA Fore reef sites (135) 15 countries 3 – 9 m depth Rel % macroalgae Gardeners: Stegastes spp. Microspathadon chrysurus Scrapers: Scarus coelestinus Scarus guacamaia Scarus taeniopteris Scarus iserti Scarus aurofrenatum Acanthurus bahianus Acanthurus chirurgus Rel % macroalgae Browsers: Kyphosus sectatrix Acanthurus coeruleus Sparisoma chrysopterum Sparisoma rubripinne Scarus coeruleus Sparisoma atomarium Sparisoma radians Crypototomus roseus Significant negative correlations for Excavators & Scrapers 100 80 60 r2 = 0.229 p = 0.004 40 20 0 0.0 0.5 1.0 1.5 2.0 2.5 Scraper biomass (kg 100m-2) 3 10/24/2013 Costa Rica removed (1 site) Canopy height (mm) 0 1 2 3 4 5 6 7 Total herbivore biomass (kg 100m-2) 7 6 5 4 3 2 1 0 0.0 r2 = 0.125 p = 0.040 0.5 1.0 1.5 60 Significant negative correlation for All Herbivores, Excavators & Scrapers 2.0 7 6 r2 = 0.229 p = 0.004 5 4 3 2 1 0 0.0 0.5 1.0 1.5 2.0 2.5 3.0 3.5 Excavator biomass (kg 100m-2) Fish biomass (kg m-2) Functional group composition relative to total biomass 1.8 Browser (8 spp.) 1.5 Excavator (2 spp.) 1.2 Scraper (7 spp.) 0.0 40 r2 = 0.002 p = 0.808 All herbivores: N/S relationship 30 20 Significant positive correlation for Excavators 10 0 2 4 6 8 10 12 Total herbivore biomass (g 100 m-2) 60 50 40 r2 = 0.270 p = 0.001 30 20 10 0 0.0 0.5 1.0 1.5 2.0 2.5 3.0 3.5 Excavator biomass (g 100m-2) Findings: • Few sites with high herbivore biomass in Caribbean 38% • Herbivorous fishes are functionally diverse; distinct sub-groups vary in impact on benthos • Scrapers & excavators appear to have a particularly strong impact Gardener (6 spp.) • Only 2 excavator species in the Caribbean 0.6 0.3 50 0 2.5 Scraper biomass (kg 100m-2) 0.9 AGRRA data: Sub-country means Fore reef, 3 – 9 m % Live Coral Cover % coral cover r2 = 0.181 p = 0.012 AGRRA data: Sub-country means Fore reef, 3 – 9 m % coral cover 7 6 5 4 3 2 1 0 Canopy height (mm) Canopy height (mm) Macroalgal canopy height • Excavators decline quickly on heavily fished reefs 24% < 1.1 (n=9) • Low biodiversity -> vulnerability -> decreased resilience? 1.1 - 2 (n=8) 2.1 - 3 (n=11) > 3 (n=7) Total herbivorous fish biomass (kg m-2) • To manage for resilience, need to identify & focus on important & vulnerable functional groups St Thomas 2005 Photo: Jen Schull Food chain Trophic levels Trophē = feed 4 10/24/2013 Rank Abundance Diagrams Rank Abundance Diagrams indicate differences in species richness & evenness Birds of Great Lakes Basin http://cfpub.epa.gov/ncer_abstracts/index.cfm/fuseaction/display.abstractDetail/abstract/7441/report/F Species accumulation curve 5 10/24/2013 Foundation species Important species • Keystone species – disproportionately large effect on its environment relative to its abundance Keystone Species Oculina arbuscula Elephants Keystone species Ochre star (Pisaster) Sea otter Community dynamics— succession Disturbance •Humans can change the flow of energy. •Fishing down the food web. 6 10/24/2013 Figure 16.5 A Theoretical Model of Succession 37 38 Community Succession and Species Richness ? 39 Study questions 1. Define community 2. Why is a community more than the sum of its parts? 3. Define guild. 4. What is the difference between a trophic web and an interaction web? 5. Compare direct from indirect interactions. 6. What is the difference between bottom-up and top-down control within a food web? 7. Describe a rank-abundance curve, noting what is plotted along each axis. How would the curves differ for communities with relatively higher vs. lower species richness and for higher vs. lower species evenness? 41 Study questions 8. What 2 pieces of information are used in calculating species diversity (such as the Shannon index)? 9. What would cause a community with low species richness have a higher species diversity than another community with higher species richness? 10. Define: foundation species, ecosystem engineer, & keystone species. Provide an example of each. 11. Define succession. Provide an example of the steps of succession in a community, being clear as to what happens to allow each step to proceed into the next. 12. At what point in community succession is species richness the greatest? Why? 42 7 10/24/2013 Study questions 13. What role does disturbance play in a community? 14. What is meant by “space for time substitution” of succession? 15. Define alternate stable states and give an example. 43 8
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