BIOS 5970: Plant-Herbivore Interactions Dr. Stephen Malcolm, Department of Biological Sciences • Week 9. Ecology & Evolution of Mutualisms: – Lecture summary: • Mutualism: a world of cheerful cooperation? • Energetics of bee foraging • Factors limiting plant reproduction • Fruit profitability • Dispersal probability • Consequences of seed dispersal Bombus vagans http://bugguide.net/node/view/470601 BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 1 2. Mutualisms: • Charles Darwin noted that Victorians liked to – “behold the face of nature bright with gladness” in a world of cheerful cooperation. • Darwin used this attitude to contrast with the reality in which a mistletoe exploits a bird for dispersal. • But the bird eats the fruit for food and leaves the sticky seeds in clumps where they compete intraspecifically for limited resources. BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 2 3. Mutualism and optimality: • Mutualisms between plants and animals are widespread. • Howe & Westley consider the reason why. • They use an analysis of costs and benefits to the interactants. • Ideas about optimization borrowed from economics. BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 3 4. Balancing costs and benefits: • Bernd Heinrich estimated the energetic costs and benefits of bee foraging for nectar and pollen of flowers. • Bees with longer tongues were more efficient at extracting nectar than shorttongued bees (Fig. 7-1) BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 4 5. Bumblebee foraging at flowers: • 4 Bombus species foraging at: – Long-corolla flowers. – Short-corolla flowers (<3mm). – Pollen flowers. • Partition resources according to tongue length (Fig. 7-2). – – – – B. vagans >8mm, stippled B. fervidus >8mm, open B. terricola <7mm, lined B. ternarius<7mm, black BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 5 6. Diversity of pollinator species: • Many pollinator species may visit related flower species, • E.g. 55 species of sunflowers (Asteraceae) in Illinois: – Frequency distribution of species (Fig. 7-3). BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 6 7. Majoring and minoring by bumblebees • In the absence of competition bees narrow their foraging range of flower species (Fig. 7.4A). • In the presence of competition bees visit the same most profitable flowers on subsequent trips (Fig. 7.4B). BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 7 8. Experience and improved handling: • Foraging bumblebees learn to handle jewelweed (Impatiens) flowers more accurately with experience (Fig. 7-5). BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 8 9. Flower patchiness and residence times: • Foraging bees stay longer in patches of flowers with most nectar. • They do this by travelling short distances after encountering high nectar volumes (Fig. 7-6). BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 9 10. Bee Foraging: • 6 options for coping with competition: – (1) Visit more flower species. – (2) Improve nectar-handling efficiency. – (3) Extend foraging range rather than switch to minor foods. – (4) Territoriality to defend food resources. – (5) Respond to high rewards with short movements. – (6) Respond to lower reward variances rather than high reward means. BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 10 11. Limiting factors for plant reproduction: • Ovule-limited seed production: – When abundant pollen grains fail to fertilize eggs. • Pollen-limited seed production: – When some ovules are not fertilized (like Fig. 7-8). • Resource-limited seed production: – When fertilized eggs are aborted because light or soil resources are insufficient. • Herbivore-limited seed production: – Fertilized ovules eaten by consumers before maturity. BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 11 12. Fruit profitability & dispersal probability: • Fruit size and fruit disperser size both influence fruit profitability and the likelihood that a particular fruit will be dispersed by a particular disperser. – Fig 7-10 - profitability as a function of fruit size and bird size (a = 17g, b = 250g, c = 500 g birds) – Profitability P depends on pulp weight (Fm) nutritional content (Fn) and the basal metabolic weight (M0.61)/ exploitation time (T). BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 12 13. Fruit diameter and gape width: • Maximum fruit diameters are positively correlated with bill gape in 36 bird species in the mountains of Costa Rica (Fig. 7-11). BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 13 14. Fruit size and bird disperser diversity: • Large fruited plants have fewer bird species visiting than small fruited plants (Table 7-3). BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 14 15. Ecological Consequences of Seed Dispersal: BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 15 16. Seedling recruitment: • Product of dispersed seed shadow and probability of seed/seedling survival (Fig. 7-15). BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 16 17. Pollen dispersion: • Pollen follows a negative binomial distribution with most pollen deposited near its flower source (Fig. 8-2). • Higher probability of same species fertilization BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 17 18. Consequent gene dispersal: • Most butterfly pollinator flights are <3m between plants of Phlox pilosa (Fig. 8-3). BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 18 19. Patterns of pollen dispersion in plant communities: • Pollinators moved dyed pollen further in populations of chickweed alone – (Fig. 8-4 top). • Dyed pollen had more restricted movement in chickweed when grown mixed with spring beauty – (Fig. 8-4 bottom). BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 19 20. Constraints on coevolution between plants and their mutualists: BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 20 21. Diversity of symbiotic mutualisms: BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 21 Figure 7-8: Pollinator (fungus gnat) limitation in jack-in-the-pulpit. BIOS 5970: Plant-Herbivore Interactions - Dr. S. Malcolm. Week 9: Ecology & Evolution of Mutualisms Slide - 22