1. Lecture 8 – INTERTIDAL - ZONATION PHYSICAL FACTORS

2. 1. DESICCATION Experiment (Frank, 1965. Ecol. 46: 831) Barnacle Fucus Before artificial seep

3. 1. DESICCATION Experiment (Frank, 1965. Ecol. 46: 831) After artificial seep Barnacle Fucus Before artificial seep

4. 1. DESICCATION A second kind of experiment (Foster ‘71, J. Anim. Ecol. 40:33)

5. 1. DESICCATION A second kind of experiment (Foster ‘71, J. Anim. Ecol. 40:33)

6. 1. DESICCATION Avoiding drying -seeking refuge (Kensler, 1967, Carefoot, 1977) Inner Region Middle Region Outer Region Transient species Highest diversity Very few inhabitants Clay, fine silt, sand Gravel, shells, coarse sand

7. 1. DESICCATION Avoiding drying 1. Barnacles - trap water CO 2 O2O2 2. Mussels - Airgape - open valves repeatedly during low tide

8. 1. DESICCATION Coping with oxygen depletion Fucus resubmerge Percentage of initial water retained Percentage of initial water lost O 2 consump- tion 100 50 100

9. 2. TEMPERATURE

11. METABOLIC RATE ºC INTERTIDAL INVERTEBRATES DEEPER WATER INVERTEBRATES

12. 2. TEMPERATURE Upper Lethal Temperature Median lethal time (hrs) 40 35 30 Balanus crenatus S. balanoides Chthalamus 125102050

13. 2. TEMPERATURE Upper Lethal Temperature Median lethal time (mins) 45 40 35 30 50100 Asterias Ophioderma Arbacia Uca Ilyanasa

14. 2. TEMPERATURE -effects of substrate and crowding TISSUE ºC EXPOSURE TIME solitary cobble crowded cobble solitary boulder crowded boulder solitary cobble crowded cobble solitary boulder crowded boulder High intertidal Low intertidal

15. 2. TEMPERATURE -effects of shading Surface ºC TIME 40 10 Canopy removed Under canopy

16. 2. TEMPERATURE Latitudinal effects Helmuth et al, Ecol. Monogr. 2006

17. 2. TEMPERATURE -low temperature

18. 2. TEMPERATURE -low temperature Dendronotus frondosus (Gionet & Aiken, 1992) % Survivorship Temperature (4 hr exposure) 0-4-8-10-12 100 50 0

19. 3. WAVE STRESS a. Limitation of size Water flow 100% 90% Boundary layer

20. 3. WAVE STRESS a. Limitation of size Water flow

21. 3. WAVE STRESS b. Holding on Keyhole limpet

22. 3. WAVE STRESS b. Holding on - body orientation Water flow

23. 3. WAVE STRESS b. Holding on - body orientation <.5 m/s -90090 >.5 m/s Freq Orientation (º to flow)

24. 3. WAVE STRESS b. Holding on - tenacity What is “tenacity”? 1. Suction? Atmospheric pressure ≈ 1 kg/cm 2 Patella ≈ 5 - 7 kg/cm 2

25. 3. WAVE STRESS b. Holding on - tenacity What is “tenacity”? Patella

26. 3. WAVE STRESS b. Holding on - tenacity What is “tenacity”? 2. Adhesion F = 2 A S d area surface tension thickness Theoretical adhesion = 600 kg/cm 2

27. 3. WAVE STRESS b. Holding on - tenacity What is “tenacity”? 2. Adhesion F 1d1d Tenacity (kg/cm 2 to detach) Weight of mucous

28. 3. WAVE STRESS a.Limitation of size - plants Laminaria

29. 3. WAVE STRESS - How plants deal with it current Movement of plant – dissipates E Reaction force Inertial force

30. 3. WAVE STRESS -can extend intertidal zones Upper limit of barnacles Upper limit of mussels Upper limit of fucoids Upper limit of kelp ELWS EHWS ExposedSheltered

32. Effects on limpet distribution Todgham et al, 1997

33. Effects on limpet distribution Todgham et al, 1997 HYPOTHESES 1. Greater density of limpets the wave-exposed site. 2. Limpets will be found more frequently in habitats with refuges. 3. Limpets will be found less frequently in wave protected habitats with refuges.

34. Effects on limpet distribution Todgham et al, 1997 Habitats ExposedProtected

35. Effects on limpet distribution Todgham et al, 1997 Wave Velocity Recorder

36. Effects on limpet distribution Todgham et al, 1997 Lottia digitalisLottia paradigitalis Lottia pelta Tectura personnaTectura scutum

37. Effects on limpet distribution Todgham et al, 1997 At each site recorded: 1.Species 2. Size class - Small, Medium, Large 3.Microhabitat a.Bare rock b.Bare rock with barnacles (Balanus) c.On/under algae d.Crevices

38. Effects on limpet distribution Todgham et al, 1997 SpeciesProtectedExposedP-value Lottia digitalis 50 ± 7.362.4 ± 6.44NS L. paradigitalis 18.6 ± 3.728.0 ± 2.7NS L. pelta19.6 ± 3.3318.7± 1.44NS Tectura scutum 25.4 ± 3.1133.9 ± 2.880.009 T. personna25.9 ± 3.8Not foundXXXX

39. Effects on limpet distribution Todgham et al, 1997 Low tideHigh tide L. digitalis L. paradigitalis L. pelta T. personna T. scutum

40. Effects on limpet distribution Todgham et al, 1997 Wave protected L. digitalis frequency Habitat frequency Bare rock Rock/Barnacle Cover Crevice Bare rock Rock/Barnacle Cover Crevice Wave exposed Lottia digitalis

41. Effects on limpet distribution Todgham et al, 1997 Distribution of size classes in all species

42. Effects on limpet distribution Blanchette, 1997 Growth and survival of Fucus gardneri

43. Effects on Fucus Blanchette, 1997 Growth and survival of Fucus gardneri

44. Effects on Fucus Blanchette, 1997

45. Growth and survival of Fucus gardneri Effects on Fucus Blanchette, 1997

46. Growth and survival of Fucus gardneri March August February Planiform area m 2 Exposed Protected Effects on Fucus Blanchette, 1997

47. Growth and survival of Fucus gardneri March August February Mean Length Exposed Protected Effects on Fucus Blanchette, 1997

48. Growth and survival of Fucus gardneri March August February Mean Mass Exposed Protected Effects on Fucus Blanchette, 1997

49. Transplants Effects on Fucus Blanchette, 1997

50. Transplants P to P P to E E to P E to E P to P P to EE to P E to E Mean area Maximum area Mean area Sept Effects on Fucus Blanchette, 1997

51. Transplants P to P P to E E to P E to E Reproductive Status (number of blades with mature receptacles) Effects on Fucus Blanchette, 1997

52. Next time Intertidal Zonation - Biological Factors