Effect * of species 1 on species 2

1. Effect * of species 1 on species 2 - + - COMPETITION PREDATION Effect of species 2 on species 1 + MUTUALISM PREDATION * On per capita growth rate

2. Two ways to Classify Mutualisms I. By Function: • Trophic Mutualisms • Protective Mutualisms • Dispersive Mutualisms

3. Two ways to Classify Mutualisms II. By relationship of mutualists for one another: Obligate: species are so dependent on their mutualistic relationship that cannot live in its absence. e.g., symbionts lichens leaf-cutter ants & fungus termites and gut fauna Facultative: species can live without their mutualistic partner e.g., oxpeckers cleaner-wrasse many diffuse mutualists Diffuse Mutualisms: Mutualistic interactions that where the “partner” can be one of any species, e.g., pollinators, seed dispersers.

4. Trophic Mutualisms – involve a mutual exchange of energy/nutrients – specialized to the point of obligates Mycorrhizal fungi Lichens

5. Nitrification processes carried out only by specialized bacteria NH3 NO2- NO2- NO3- Nitrosomonas (soil) Nitrosoccus (marine) Nitrobacter (soil) Nitrococcus (marine) Denitrification processes carried out only by specialized bacteria, such as Pseudomonas denitificans NO3- NO2-  NO Nitrogen fixation offsets Denitification and is accomplished by free-living bacteria, such as Azotobacter, symbiotic bacteria, such as Rhizobium occurring in roots of legumes, and cyanobacteria. Nitrogen-fixing bacteria in root nodules of soybeans

8. Protective Mutualisms – involve an exchange of protection from predators/parasites often for an exchange of resources

9. Ant-leafhopper-goldenrod interactions

10. #alarm signals before • and after predator • contact Alarm Calling • prey-2 predator • Tree hopper-ant mutualisms

11. More ants patrolling Fewer LB remain • Mechanically mimicking the tactile alarm call

12. Dispersive Mutualisms – involve an exchange of a reproductive service often for an exchange of resources Magnificent Hummingbird

13. Seed Dispersal BANFF NATIONAL PARK: Friendly Relations Between Clark’s Nutcracker and Whitebark Pine

14. Seed dispersal American robin and Viburnum

15. Experimental tests of mutualisms Beltian bodies Janzen’s study of Psuedomyrmex and the bullshorn acacia.

16. A comparison of acacia with ants and those with ants removed Growth (cm) by bullshorn acacia Survival (%) by bullshorn acacia 15-5     Source: Janzen 1966

17. The benefit to the plant is fewer herbivorous insects

18. Mutualisms…..Really?

19. 2000 interactions/day w/queue • Eat parasites (coop) & mucosal • tissue/scales (defect) • The latter leads to client “jolt” • so an observable cue

20. Jolts are predictors: • 12-18/min among biting cleaners • 2-3/min among coop cleaners • Do clients image score? • If previous client ends w/conflict • (jolt/chasing): • 100% bystanders (queue) depart • w/o conflict 100% approach • Audience effects? • 17 vs. 6 cleaners spp showed (-) • relationship between jolts and • bystander presence • And also observed in Sergeant major

21. More ants patrolling Fewer LB remain • Mechanically mimicking the tactile alarm call

22. Keystone Mutualists(Eco Eng, Foundation Spp) Plant species Avian frugivores Virola surinamensis 7 Tetragastris panamensis 12 Virola sebifera 6 Caesearia corymbosa ** 22 Gurarea glabra 19 Didymopanax morototoni 37 Miconia argentea 46 ** Considered a Keystone species by Howe (1977) because it bears a fruit crop for 10 week period including December, the time of lowest fruit production in La Selva forest

23. Casearia corymbosa Red-lored Amazon eats the arils and drops the fruit w/o dispersing

24. Chestnut-mandibled Toucan Eats huge quantities of seeds, but a poor disperser. Toucans however depend on Casearia seeds in December when all their preferred foods are gone.

25. The only known dependable disperser for Casearia seeds. Like the toucan, it appears to have no alternative food to eat in December. Disappearance of key fruit would push the tityra, the toucan, and other species to local extinction. Without its main disperser, Casearia, Virola (primarily dispersed by the Chestnut-mandibled toucan), and other trees dependent on these birds for dispersal would slowly lose their competitive position and diversity would slowly erode away in the tropical rain forest in La Selva.

26. Alternative *forms* of mutualisms (1) Physical Ecosystem Engineers (after Jones et al. 1997) What does a tree do in the forest? • living and dead tissues are eaten by animals (+,-) • competes with other plants for water and nutrients (-,-) • branches, bark, roots, and leaf surfaces make shelter, resting locations, • and living space • small pools for organisms are created where water gets channeled into • crotches • soil cavities that from when roots grow provide places to live and cache food • leaves and branches cast shade, reduce the impact of rain and wind, moderate • temperature extremes, and increase humidity in the under story and soil • root growth aerates the soil, alters its texture, and affects the infiltration of • water • And more…..

27. Jones et al. called these modifications ecosystem engineering And there are many, many, many examples. Some almost trivial, such as plants creating shade?? Others are very obvious….

28. And EE have both positive and negative effects on individual Species or the Sometimes it’s the non-living structure left behind that provides the benefit, other times it may be new structures created as a result of an animal’s activity.

29. Coral reefs Kelp forest Sometimes Eco. Engineers are called Foundation species when they form the base from which a whole community of species is built