1. Lecture 8 – INTERTIDAL - ZONATION PHYSICAL FACTORS

2. Studies of intertidal ecology Descriptive phase Understand process Understand interactions Investigate physiological/genetic/cellular mechanisms

3. Studies of intertidal ecology Organism A Distribution on the intertidal Effects of physical factors Organism B Distribution on the intertidal Effects of physical factors

4. Immersion time Emersion time Temperature & desiccation Wave action Physical Factors on the Intertidal

5. Barnacle life cycle Nauplius Cyprid Settle Adult Benthic Pelagic

6. 1. Desiccation Barnacle cyprids Foster 1971. Mar. Biol. 8: 12-29 Time (hours) 0369121518 Percentage 100 50 0 mortality water loss

7. 1. Desiccation Foster 1971. Mar. Biol. 8: 12-29 Median lethal time (hours) 020406080100120140 Basal diameter (mm) 12 10 8 6 4 2 0 Balanus crenatus Semibalanus balanoides Elminius modestus

8. Why are larger animals more resistant to desiccation? 1 cm Surface area = 6 cm 2 Volume = 1 cm 3 2 cm Surface area = 24 cm 2 Volume = 8 cm 3 Ratio = 6:1 Ratio = 3:1

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

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

11. 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

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

13. 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

14. 2. TEMPERATURE

16. METABOLIC RATE ºC INTERTIDAL INVERTEBRATES DEEPER WATER INVERTEBRATES

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

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

19. 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

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

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

22. Weather for St. Andrews 19 Jan – 1 Feb

23. 2. TEMPERATURE Tolerances within genera LT 50 45 40 35 30 25 20 P. eriomerus P. cinctipes 1020304050 LT 50 = MHT 19 species of Petrolisthes Mean Habitat Temperature (MHT) Somero. 2002. Int.Comp. Biol. 42:780 P. cinctipes P. eriomerus

24. 2. TEMPERATURE Tolerances within genera Somero. 2002. Int.Comp. Biol. 42:780 T. funebralisT. brunneaT. montereyi L. sctulataL. keenae

25. Temperature and Aggregation Chapperon & Seuront. 2012. J. Therm. Biol 37: 640

26. Temperature and Aggregation Chapperon & Seuront. 2012. J. Therm. Biol 37: 640

27. Temperature and Aggregation Chapperon & Seuront. 2012. J. Therm. Biol 37: 640

28. Desiccation and Aggregation Coleman 2010. J.Exp.Mar.Biol.Ecol. 386:113 No significant differences

29. Coleman 2010. J.Exp.Mar.Biol.Ecol. 386:113 Desiccation and Aggregation No significant differences

30. 2. TEMPERATURE -low temperature

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

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

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

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

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

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

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

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

39. 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

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

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

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

43. 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

45. Effects on limpet distribution Todgham et al, 1997

46. 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.

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

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

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

50. 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

51. 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

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

53. 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

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

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

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

57. Effects on Fucus Blanchette, 1997

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

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

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

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

62. Transplants Effects on Fucus Blanchette, 1997

63. 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

64. 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

65. Next time Intertidal Zonation - Biological Factors