1. Population Interactions Competition (--) when both species suffer from an association Predation (+-) when one benefits and one suffers Commensalism (+0) when one species benefits from another and it is unaffected Amensalism (-0) when one species negatively affects another and it is unaffected Mutualism (++) when both species benefit from another

2. Competition

3. Liebig’s Law of the minimum Guiding principle: Liebigs's Law of the Minimum The distribution of a species will be controlled by that environmental factor for which the organism has the narrowest range of adaptability or control. The growth of a population of organisms increases until the supply of a critical resource becomes limiting

4. Liebig’s Law of the Minimum: an example “growth of phytoplankton is dependent on the minimum amount of nutrients/light present” whatever is in shortest supply will limit (and may stop) growth –nitrate, silica, phosphate, or iron limitation –light limitation if a nutrient (or light) is at low levels, it may be limiting growth

5. Limiting Resources: an example Space is a limiting resource to these colonies of colonial ascidians

6. Intraspecific competition: Background Individuals of same species Shared resource demands and use of a limiting resource- food, shelter, mates As individuals compete for resources some are deprived- resulting in reduced fitness, reflected by lower growth, fecundity and survivorship

7. Intraspecific competition: an example with limpets Patella cochlear

8. Intraspecific Competition Intrapspecific competition among limpets

9. Intraspecific Competition (Self Thinning) Reduced individual fitness Reduced body size Increased mortality

10. Gause’s Competitive Exclusion Principle When two species compete for identical resources, one will be more successful and will eventually eliminate the other

11. Forms of Interspecific Competition Interference (Contest) Competition –access to a resource is limited or denied by the dominant species –examples include antibiotics secreted by microorganisms, or territorial behavior Exploitative (Scramble) Competition –the direct use of a resource before a competitor can use it, thus reducing its availability simply by elimination

12. Nature of the interspecific competitive interaction Direct competition- Interference competition –Dominate resource –Need for space - e.g. Sessile & territorial –Prevent use by others, “winner takes all” Indirect competition- Exploitation competition –Competitors can not dominate the resource –Reduces resource availability by using it up –Resource is “shared” - no outright “winner”

13. Competition Where do you see evidence of competition in the oceans ? –Overgrowth –Aggressive behavior What’s the result of interspecific competition? –Dominance or monopoly by a single species in a given habitat –Competitive exclusion

14. Persistence among competing species Behavioral acclimations- learn to feed when competitors are not present Character displacement-through time two closely related species tend to be more distinct morphologically and therefore use different portions of limiting resources Change in habitat utilization

15. Competition in unusual forms Overgrowth competition-one species overgrows a second species –Some corals and sponges Chemicals are used to defend access to a shared resource –Big impacts on settlement –Allelopathy in some sponges

16. How to assess competitive effects Measure: –Resource dynamics –Number of competing individuals Disadvantageous impacts on : –survival rate –growth rate –adult weight –fecundity

17. How do we measure competition in the field? 1. Observe the patterns of distribution and see if they conform to predictions of competition theory white shrimp brown shrimp Estuarine salinity gradient One conclusion is that white and brown shrimp are in competition with each other (competition theory would predict this distribution) Weak argument: there are multiple explanations for this distribution Low High

18. Classic study in Experimental Ecology Connell (1961) Experiments with intertidal barnacles Connell (1961) Experiments with intertidal barnacles Observations Two species – Chthamalus stellatus and Balanus balanoides (now Semibalanus balanoides) Chthamalus adults in upper zone, juveniles in both upper and lower zone. Balanus in lower zone Observations Two species – Chthamalus stellatus and Balanus balanoides (now Semibalanus balanoides) Chthamalus adults in upper zone, juveniles in both upper and lower zone. Balanus in lower zone

19. Observations Chthamalus juveniles Chthamalus adults Balanus

20. Model H a : Competition for space with Balanus prevents adult Chthalamus from occurring in the lower area H 0 : When present together Balanus has no effect on Chthalamus

21. Experiment Transplanted stones with Chthamalus to lower level Followed settlement of Balanus, removed them from one half of each stone Recorded the fate of individual barnacles Transplanted stones with Chthamalus to lower level Followed settlement of Balanus, removed them from one half of each stone Recorded the fate of individual barnacles Balanus Chthamalus

22. Results Chthamalus survival was much greater where Balanus was excluded. Most Chthamalus killed by being overgrown or undercut by Balanus Chthamalus survival was much greater where Balanus was excluded. Most Chthamalus killed by being overgrown or undercut by Balanus

23. Experimental conclusions Balanus –upper limit set by physical environment –lower limit set by Thais predation Chthamalus –upper limit probably set by physical environment –lower limit set by interspecific competition Asymmetry

24. growth rate Location in intertidal zone low high middle Chthamalus alone competitive release – niche of the competitively-inferior species expands in the absence of the competitively-superior species fundamental niche realized niche Chthamalus with Balanus competitive release

25. Character displacement When two species occur in sympatry natural selection should favor the evolution of mechanisms that reduce competition if resources are limiting This often takes the form of character displacement, where the two competing species diverge in a trait that reduces the strength of interspecific competition Use Resource

26. Character displace- ment: mud snails Hydrobia ventrosa Hydrobia ulvae A A S S

27. The importance of intra- and interspecific competition Can have strong negative impact on the population growth of inferior competitor Reduces the geographic distribution of competing species Alter evolutionary trajectories.

28. The Niche Concept and Competition in Evolutionary Time Niche - the role of a species in a community, defined in practice by measuring all possible resources used and tolerance limits Niche Breadth - The amount of a resource used by an organism; this amount may change when new species are introduced or removed from a community

29. Niches and Types of Species Generalist species have large niches, tolerate wide range of environmental variations, and do better during changing environmental conditions Specialist species have narrow niches; they are more likely to become endangered; these do better under consistent environmental conditions

30. Niche Breadth (width or size) Some plants and animals are more specialized than others, and measures of niche breadth attempt to measure this quantitatively

31. Diet breadth consumes only one prey type consumes many prey types broad diet narrow diet specialist generalist

32. Niches and Natural Selection Region of niche overlap Generalist species with a broad niche Specialist species with a narrow niche Niche breadth Niche separation Number of individuals Resource use

35. Fundamental niche depends on physical (abiotic) conditions. Realized niche depends on biotic as well as abiotic conditions. What is the realized niche of each barnacle? What is the fundamental niche of each?

36. growth rate Location in intertidal zone lowhighmiddle How can we determine the realized niche of each barnacle? Where do they grow when allowed to compete? Balanus realized niche Chthamalus realized niche Balanus and Chthamalus