1. Lecture 17 ROCKY SUBTIDAL

3. Substrate Types CobbleWalls Horizontal bedrock Sandy

4. ZONATION CHART DATUM infralittoral fringe littoral About 200 m Infralittoral Circalittoral Adequate light for photosynthesis Infralittoral + Circalittoral = Subtidal

5. ZONATION CHART DATUM infralittoral fringe littoral Upper infralittoral Lower infralittoral Upper circalittoral Lower circalittoral Laminaria forest + foliose algae + few animals Foliose algae + few animals Few foliose algae + many animals No algae many animals

6. Rocky Subtidal - Zones 1) Sublittoral fringe Laminaria Bryozoan Anemone (Metridium)

7. Rocky Subtidal - Zones 2) Infralittoral Laminaria Anemone (Metridium) Strongylocentrotus droebachiensis Urchin barrens

8. Rocky Subtidal - Zones 3) Circalittoral Anemone (Metridium) tunicates Sea stars Lamp shells

9. Determinants of Zonation Slope HorizontalVertical

10. Determinants of Zonation Slope Differences in species’ patterns 1. Higher light levels on horizontal surfaces 2. Higher sedimentation on horizontal surfaces 3. Photonegative larvae select walls/overhangs 4. More shelter from disturbance

11. Determinants of Zonation Depth Density Depth 33 m42 m % Cover Depth 28 m45 m Brachiopods Sponges

12. Local Processes Affecting the Community Temperature 0 5 10 15 20 25 30 0 5 10 15 20 25 30 111213 192325 Maine (Machias Seal Is) Aug. 1995 Galapagos (May1999)

13. Local Processes Affecting the Community Temperature Internal wavesUpwelling

14. Physical Disturbance Depth LowHigh

15. Physical Disturbance

16. Creation of Patches Type I Disturbance -death of some residents -patch surrounded by survivors Type II Disturbance -widespread destruction Recovery - vegetative growth & asexual budding Recovery - long distance dispersers -more stochastic

17. Physical Disturbance Creation of Patches Wittman (‘98) Patch of anemones Predation by nudibranchs Got Type I patches

18. Physical Disturbance Creation of Patches Wittman (‘98) ExpectGot Recruitment of planktonic bryozoans, polychaetes and ascidians Asexual/vegetative growth of neighbours

19. Physical Disturbance Flow Patterns - Reidl ‘71 Primary - effect of flow directly on organisms Secondary - effect of flow on transport of nutrients, gasses, foods Tertiary - effect of flow on substrate

20. Physical Disturbance Flow Patterns 20406080100120140 cm/s 2 4 6 8 10 Minimum Maximum DepthDepth

21. Physical Disturbance Sedimentation Sedimentation rate = 1/flow 0 5 10 15 20 25 30

22. Light 0 5 10 15 20 25 30 35 40 45 50 Depth Percent surface light.01.1110100 Turbid coastal water Clear open ocean water Limit for Laminaria Limit for foliose algae

23. Rocky Subtidal - communities Terebratulina septentrionalis Community Terebratulina septentrionalis Chiton

24. Rocky Subtidal - Communities Crustose coralline algae Community Terebratulina septentrionalis Chiton Bryozoans

25. Rocky Subtidal - Biotic Factors

26. Biotic Factors on the Subtidal Predictions: 1. Predation - reduced in harsh environments 2. Biotic factors set lower distribution limits 3. Physical factors set upper distribution limits

27. Biotic Factors on the Subtidal 1. Predation - reduced in harsh environments +5 0 5 10 15 20 LowHigh Predator pressure

28. Biotic Factors on the Subtidal 1. Predation - reduced in harsh environments PREDATOR SURVIVAL Number of waves 0 100 200 Time (Months) 0 6 12 Denny et al, 1985 Survivorship L r /2 LrLr 2L r L r /2 LrLr 2L r Colisella digitalis

29. Predator density Galapagos Seychelles Gulf of Maine Biotic Factors on the Subtidal 1. Predation - reduced in harsh environments Absence of predators

30. Biotic Factors on the Subtidal 1. Predation - reduced in harsh environments minuteshoursdays Birds Crabs/lobster Snails/urchins /seastars Impact of predator on prey (per capita prey removal rate) Return time of limiting stress

31. Biotic Factors on the Subtidal 1. Predation - reduced in harsh environments New Zealand

32. Biotic Factors on the Subtidal 1. Predation - reduced in harsh environments urchins, sea stars, lobsters freshwater mussels New Zealand

33. Biotic Factors on the Subtidal Predictions: 1. Predation - reduced in harsh environments 2. Biotic factors set lower distribution limits 3. Physical factors set upper distribution limits √

34. Biotic Factors on the Subtidal Predictions: 1. Predation - reduced in harsh environments 2. Biotic factors set lower distribution limits 3. Physical factors set upper distribution limits

35. Biotic Factors on the Subtidal 2. Biotic factors set lower distribution limits Anderson Ledge, New Hampshire Laminaria - limit dictated by urchin grazing Urchins - stay near bottom of Laminaria - physical stress too high to go shoreward Mytilus - colonized areas grazed by urchins Sea stars - prevent Mytilus recruitment lower in subtidal

36. Biotic Factors on the Subtidal 2. Biotic factors set lower distribution limits Scheibling et al, 1999 Mill Cove Little Duck Island

37. Biotic Factors on the Subtidal 2. Biotic factors set lower distribution limits Strongylocentrotus droebachiensis Laminaria longicruris

38. Biotic Factors on the Subtidal 2. Biotic factors set lower distribution limits Scheibling et al, 1999 kelp grazed edge

39. Biotic Factors on the Subtidal 2. Biotic factors set lower distribution limits Scheibling et al, 1999

40. Biotic Factors on the Subtidal 2. Biotic factors set lower distribution limits But …. within limits set by physical factors Vadas & Steneck, 1988 - depth greater without grazers

41. Biotic Factors on the Subtidal 2. Biotic factors set lower distribution limits But …. within limits set by physical factors Miller and Hay, 1996 Oculina

42. Biotic Factors on the Subtidal Predictions: 1. Predation - reduced in harsh environments 2. Biotic factors set lower distribution limits 3. Physical factors set upper distribution limits √ (with qualifications)

43. Biotic Factors on the Subtidal Predictions: 1. Predation - reduced in harsh environments 2. Biotic factors set lower distribution limits 3. Physical factors set upper distribution limits √ (with qualifications)

44. Biotic Factors on the Subtidal 2. Competition +5 0 5 10 15 20 LowHigh Competition for space

45. Biotic Factors on the Subtidal 2. Competition Competition for space (among algae) a. allelopathy

46. Biotic Factors on the Subtidal 2. Competition Competition for space (among algae) b. overgrowth

47. Biotic Factors on the Subtidal 2. Competition Competition for space (among algae) c. overtopping

48. Biotic Factors on the Subtidal 2. Competition Competition for space (among algae) d. Algal whiplash

49. Biotic Factors on the Subtidal 2. Competition Competition for space e. Aggression (Cnidaria) acrorhagi

50. Biotic Factors on the Subtidal 3. Recruitment +5 0 5 10 15 20 LowHigh % space covered

51. Biotic Factors on the Subtidal 3. Recruitment +5 0 5 10 15 20 LowHigh % space covered effect of predators

52. Biotic Factors on the Subtidal 3. Recruitment Osman & Whitlach, 1995 Mitrella Anachis Without predators With predators

53. Mesoscale Processes High flow Low flow km’s

54. Other mesoscale processes Disease Productivity Thermoclines