Reproduction, Migration and Dispersal

1. Lecture #3 Review Reproduction, Migration and Dispersal 1) Sex is nearly universal, despite its considerable costs to organisms. Its value is the maintenance of genetic diversity, which increases adaptability to environmental challenges. 2) Sexes may be separate (in gonochoristic species), simultaneous, or sequential in the same body. The contributions of different sexes to the next generation determine the value of hermaphroditism, as well as the relative sizes of males and females. For example, if the relative advantage of a sequential hermaphrodite functioning as a large female and small male have an advantage in maximizing offspring production, this will select for male-female transitions..

2. Reproduction, Migration and Dispersal(2) 3) Fertilization success is affected by the mode of sperm transfer, the volume of gamete production, the distance between males and females, water turbulence, the timing of spawning, and behavior. Free spawning has a number of costs, and planktonic gametes have special problems in ensuring fertilization, but specialized sperm attractants exist. 4) Marine species differ in parentalcare. Most species have free-swimming larvae, but some guard or brood eggs, brood young within body cavities, or have live birth like mammals. 5) Inalgae and corals, nonsexual reproduction involves colonial individuals (modules) that are connected to each other and exchange nutrients.

3. Reproduction, Migration and Dispersal (3) 6) Gamete production and larval life must be timed to allow settlement and promote dispersal, to avoid being swept to inappropriate habitats, and to counter predation. Egg size decreases with increasing egg numbers. 7) Marine invertebrate offspring may be (1) brooded or released as small adults, (2) dispersed to a small degree by means of short-lived, yolk-dependent lecithotrophic larvae, or (3) dispersed great distances by longer-lived plankton-feeding planktotrophic larvae. 8) Despite the potential for dispersal, planktonic larvae often settle near their origin, owing to behavior and cyclonic currents. On the microscale, larvae use a number of cues to find their settlement site. 9) The geographic range of species with planktonic dispersal is greater than the range of species without planktonic larvae. Genetic variation can be used to identify barriers to coastal zone dispersal.

4. Reproduction, Migration and Dispersal (4) 10)Migratory patterns include: Anadromy - breeding in freshwater and living in seawater (e.g., salmon, shad, sea lamprey); catadromy - Adults living in freshwater then migrate to seawater to spawn (e.g., eels); and oceanodromy- living totally in seawater but migrating from adult habitat to spawning habitat )e.g., herring and cod)

5. REVIEW QUESTIONS • Why are not all sexual species hermaphroditic? • What are the “costs” and “benefits” of sex? • Under what circumstances does it make sense for a hermaphrodite to be protandrous? Protogynous? • What might be the value, if any, of long-distance dispersal across an ocean? • What are the advantages of having planktotrophic larvae capable of settling and metamorphosing upon any substratum? What are the disadvantages?

6. REVIEW QUESTIONS • Why are species with planktotrophic larvae more common in the tropics than at polar latitudes? • What is the value of planktonic feeding larval development? • What are the potential sources of mortality for planktonic larvae? • What effect does planktotrophic larval dispersal tend to have on the geographic range of a coastal marine invertebrate species? • Anemones often occur in clones of large numbers that have arisen by fission from a founder individual. What experiment might be performed to determine the benefit of large numbers of adjacent anemones as opposed to smaller groups or solitary individuals?

7. 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

8. Competition

9. 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

10. 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

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

12. 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

13. Intraspecific competition: an example with limpets Patella cochlear

14. Intraspecific Competition • Intrapspecific competition among limpets

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

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

17. A classic interspecific competition experiment two species of Paramecium predict the outcome of interspecific competition P. aurelia P. caudata Gause (1934)

18. α12 – effect of an indv of species 2 on an indv of species 1 α21 – effect of an indv of species 1 on an indv of species 2 α21 & α21 = competition coefficients How does interspecific competition affect N? dN1 dt r1N1 K1-N1 K1 species 1 = dN2 dt r2N2 K2-N2 K2 species 2 =

19. intraspecific competition interspecific competition How does interspecific competition affect N? Lotka-Volterra equations dN1 dt r1N1 K1-N1 - α12N2 K1 species 1 = dN2 dt r2N2 K2-N2- α21N1 K2 species 2 = What does it mean ifα12= 1? What ifα12= 0.5?

20. 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

21. 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 obvious “winner”

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

23. Persistence among competing species • Behavioral acclimations- learn to feed when competitors are not present (only partially successful) • 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

24. 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

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

26. white shrimp brown shrimp Estuarine salinity gradient How do we measure competition in nature? 1. Observe the patterns of distribution and see if they conform to predictions of competition theory Low High 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

27. Classic study in Experimental Ecology Connell (1961) Experiments with intertidal barnacles Observations Two species – Chthamalusstellatus and Balanusbalanoides(now Semibalanusbalanoides) Chthamalus adults in upper zone, juveniles in both upper and lower zone. Balanusonly in lower zone

28. Chthamalus juveniles Chthamalus adults Balanus Observations

29. Model Ha: Competition for space with Balanus prevents adult Chthalamus from occurring in the lower area H0: When present together Balanus has no effect on Chthalamus

30. 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 Balanus Chthamalus

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

32. Experimental conclusions • Balanus • upper limit set by physical environment • lower limit set by snail (Thais) predation • Chthamalus • upper limit probably set by physical environment • lower limit set by interspecific competition • Asymmetry

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

34. 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 Use Resource

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

36. 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.

37. 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

38. 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

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

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

41. Niche separation Generalistspecies with a broad niche Number of individuals Specialistspecies with a narrow niche Niche breadth Region of niche overlap Resource use Niches and Natural Selection

42. Competition for resources causes evolution CS Fig. 4.7 42

45. Lotka-Volterra competition equations = including interspecific competition in the logistic model. Competition coefficients (’s) show per capita competitive effect of each species on the other. When ’s are less than 1, stable coexistence is possible.  implies that niches don’t completely overlap