Pacific Outer Continental Shelf Region 

•

OCS Study 

MMS-2005-071 

Monitoring of Rocky Intertidal Resources 

Along the Central and Southern 

California Mainland 

Comprehensive Report (1992-2003) for San Luis 

Obispo, Santa Barbara, Ventura, Los Angeles, and 

Orange Counties 
OCS Study

MMS-2005-071

Monitoring of Rocky Intertidal Resources 

Along the Central and Southern 

California Mainland 

Comprehensive Report (1992-2003) for San Luis 

Obispo, Santa Barbara, Ventura, Los Angeles, and 

Orange Counties 

Authored by: 

C. Melissa Miner 

Peter T. Raimondi 

Richard F. Ambrose 

John M. Engle 

Steven N. Murray

Submitted by: 

Department of Ecology & Evolutionary Biology Marine Science Institute 

Center for Ocean Health/Long Marine Lab University of California 

University of California Santa Barbara, CA 93106 

Santa Cruz, CA, 95060 

Environmental Science and Engineering Program Department of Biological Sciences 

University of California California State University

Box 951772 800 N. State College Blvd. 

Los Angeles, CA 90095-1772 Fullerton, CA 92834-6850 

Prepared under MMS Cooperative Agreement No. 1435-01-02-CA-85144 

U.S. Department of the Interior Camarillo 

Minerals Management Service November 2005 

Pacific OCS Region
DISCLAIMER 

This report has been reviewed by the Pacific Outer Continental Shelf Region, Minerals Management 

Service, U.S. Department of the Interior and approved for publication. The opinions, findings, 

conclusions, or recommendations expressed in the report are those of the authors, and do not necessarily 

reflect the views of the Minerals Management Service. Mention of trade names or commercial products 

does not constitute endorsement or recommendations for use. This report has not been edited for 

conformity with Minerals Management Service editorial standards. 
PROJECT ORGANIZATION 

Project Technical Officer:

Mary Elaine Dunaway, Minerals Management Service (MMS) 

Report Authors:

C. Melissa Miner, University of California, Santa Cruz 

Peter T. Raimondi, University of California, Santa Cruz 

Richard F. Ambrose, University of California, Los Angeles 

John M. Engle, University of California, Santa Barbara 

Steven N. Murray, California State University, Fullerton 

Key Project Personnel:

Principal Investigators: Peter T. Raimondi 

Richard F. Ambrose 

John M. Engle 

Steven N. Murray 

Project Staff: Stevie Adams 

Sean Bergquist 

Aimee Bullard 

 Letitia Conway-Cranos 

 Lisa Gilbane 

 Janine Kido 

 Steven Lee 

 Dan Martin 

 Melissa Miner 

 Christy Roe 

 Rafe Sagarin 

 Kim Whiteside 
TABLE OF CONTENTS 

TABLE OF CONTENTS.....................................................................................................1 

LIST OF FIGURES .............................................................................................................2 

LIST OF TABLES...............................................................................................................4 

ACKNOWLEDGMENTS ...................................................................................................5 

TECHNICAL SUMMARY .................................................................................................6 

INTRODUCTION .............................................................................................................17 

REPORT ORGANIZATION.............................................................................................17 

SAMPLING METHODS...................................................................................................18 

DESCRIPTION OF REGIONS .........................................................................................21 

RESULTS ..........................................................................................................................26 

DISCUSSION....................................................................................................................34 

LITERATURE CITED......................................................................................................98 

APPENDIX A: CATALINA ISLAND SITES.............................................................. A-1 

APPENDIX B.: NATURAL HISTORY OF TARGET SPECIES .................................B-1 

APPENDIX C.: RAW DATA TABLES ........................................................................C-1 

1
LIST OF FIGURES 

Figure 1. Location of rocky intertidal monitoring sites along the central and southern 

California Coast. . ..............................................................................................................20 

Figure 2a. Species abundances in Silvetia plots................................................................39 

Figure 2b. Species abundances in Silvetia plots. ..............................................................40 

Figure 2c. Species abundances in Silvetia plots................................................................41 

Figure 3. Species abundances in Hesperophycus plots.....................................................42 

Figure 4a. Species abundances in Endocladia plots. ........................................................43 

Figure 4b. Species abundances in Endocladia plots.........................................................44 

Figure 4c. Species abundances in Endocladia plots. ........................................................45 

Figure 5. Species abundances in Mastocarpus plots. .......................................................46 

Figure 6. Species abundances in Mazzaella plots.............................................................47 

Figure 7a. Species abundances in Anthopleura plots........................................................48 

Figure 7b. Species abundances in Anthopleura plots. ......................................................49 

Figure 8a. Species abundances in Barnacle plots. ............................................................50 

Figure 8b. Species abundances in Barnacle plots. ............................................................51 

Figure 8c. Species abundances in Barnacle plots. ............................................................52 

Figure 8d. Species abundances in Barnacle plots. ............................................................53 

Figure 8e. Species abundances in Barnacle plots. ............................................................54 

Figure 9. Species abundances in Pollicipes plots. ............................................................55 

Figure 10a. Species abundances in Mytilus plots..............................................................56 

Figure 10b. Species abundances in Mytilus plots. ............................................................57 

Figure 10c. Species abundances in Mytilus plots..............................................................58 

Figure 10d. Species abundances in Mytilus plots. ............................................................59 

Figure 10e. Species abundances in Mytilus plots..............................................................60 

2
Figure 11. Species abundances in Recovery plots............................................................61 

Figure 12a. Species abundances in Phyllospadix transects. .............................................62 

Figure 12b. Species abundances in Phyllospadix transects. .............................................63 

Figure 12c. Species abundances in Phyllospadix transects. .............................................64 

Figure 12d. Species abundances in Phyllospadix transects. .............................................65 

Figure 13a. Seastar abundances and Pisaster ochraceus colors.......................................66 

Figure 13b. Seastar abundances and Pisaster ochraceus colors.......................................67 

Figure 13c. Seastar abundances and Pisaster ochraceus colors.......................................68 

Figure 13d. Seastar abundances and Pisaster ochraceus colors.......................................69 

Figure 13e. Seastar abundances and Pisaster ochraceus colors.......................................70 

Figure 14a. Pisaster ochraceus mean size........................................................................71 

Figure 14b. Pisaster ochraceus mean size........................................................................72 

Figure 15a. Pisaster ochraceus size distributions by color. .............................................73 

Figure 15b. Pisaster ochraceus size distributions by color. .............................................74 

Figure 15c. Pisaster ochraceus size distributions by color. .............................................75 

Figure 16a. Lottia gigantea abundances. ..........................................................................76 

Figure 16b. Lottia gigantea abundances...........................................................................77 

Figure 16c. Lottia gigantea abundances. ..........................................................................78 

Figure 17a. Lottia gigantea mean size..............................................................................79 

Figure 17b. Lottia gigantea mean size..............................................................................80 

Figure 18a. Lottia gigantea size distributions...................................................................81 

Figure 18b. Lottia gigantea size distributions. .................................................................82 

Figure 18c. Lottia gigantea size distributions...................................................................83 

Figure 19a. Black abalone abundances.............................................................................84 

Figure 19b. Black abalone abundances.............................................................................85 

3
Figure 20. Black abalone mean size. ................................................................................86 

Figure 21a. Black abalone size distributions. ...................................................................87 

Figure 21b. Black abalone size distributions. ...................................................................88 

Figure 22. Common motile invertebrates in Silvetia plots. ..............................................89 

Figure 23. Common motile invertebrates in Endocladia plots.........................................90 

Figure 24. Common motile invertebrates in Barnacle plots. ............................................91 

Figure 25. Common motile invertebrates in Mytilus plots. ..............................................92 

Figure 26a. Nucella emarginata/ostrina size distributions in mussel plots......................93 

Figure 26b. Nucella emarginata/ostrina size distributions in mussel plots......................94 

Figure 27a. Tegula funebralis size distributions in mussel and Silvetia plots..................95 

Figure 27b. Tegula funebralis size distributions in mussel and Silvetia plots..................96 

Figure 27c. Tegula funebralis size distributions in mussel and Silvetia plots..................97 

LIST OF TABLES

Table 1. Summary of key species monitored at all sites, including survey methods and 

number of replicate plots. ..................................................................................................25 

4
ACKNOWLEDGMENTS 

This study would not have been possible without the assistance and understanding of the Project Technical

Officer, Mary Elaine Dunaway. 

We also could not have completed the work without a large number of volunteers who helped with field 

work, including Loana Addessi, Katherine Anderson, Katie Arkema, Mike Behrens, Irene Beers, Anne

Boettcher, Melissa Boggs, Anthony Boxshall, Karleen Boyle, Debbie Boylen, Charlene Burge, Tyra Byers, 

Jackie Campbell, Don Canestro, Miranda Canestro, Jay Carroll, Jim Castle, Sarah Chaney, Lisa Conti, Noel 

Davis, Bill Douros, Cathie Dunkel, Rosylnn Dunn, Ginny Eckert, Chris Ehrler, Jonna Engle, Caroline 

Engle, Shawn Erickson, Jared Figurski, Melissa Foley, Samantha Forde, Victor Galvan, Lauren Garske, 

Maya George, Bob Gladden, Luke Granger, Jim Grant, Nora Grant, Cricket Grice, Steve Hampton, Jessica 

Hayden-Spear, Cynthia Hays, Mike Hearne, Dave Hinrichs, Dave Hubbard, Brittany Huntington, Zach

Hymanson, George Johnson, Korie Johnson, Llad Johnson, Alison Kendall, Scott Kimura, Kristen Kusic,

Kevin Lafferty, Julie Lancer, John Lane, Shannon Lee, Haven Livingston, Dave Lohse, Maille Lyons, 

Laird MacDonald, Spencer MacNeil, Erin Maloney, James Marquez, Melinda Mayes, Jamie McConnel,

Amy McClean, Whitman Miller, Ben Miner, Mike Mitchell, Sean Morton, Jennifer O'Grady, Kathy

Pfeiffer, Petra Pless, Mark Readdie, Dan Reineman, Andrew Rice, Dan Richards, Mark Rigby, Oscar 

Rivas, Grey Sanders, Stephanie Sapper, Kim Shirley, Loretta Slusher, Jayson Smith, Linda Smith, David

Sneed, John Steinbeck, Diana Steller, Jeanine Stier, Tetsuya Tsukamoto, Jeff Tupen, Meera Venkatesan, 

Sara Warden, Natalie Wenner, Dick Wilhelmsen, and Megan Williams. 

The MMS intertidal team provided tremendous assistance in the field. This team consists of: Ann Bull, 

Mary Elaine Dunaway, Maurice Hill, Dave Panzer, and Fred Piltz. Other MMS staff assisting in the field

include: Herb Leedy, Mike McCrary, Mark Pierson, and Lynnette Vesco. 

We would like to thank Don and Miranda Canestro and the University of California Rancho

Marino/Kenneth Norris Reserve for providing shelter from the elements and access to the reef. We

gratefully acknowledge Cojo/Jalama Ranch and the Bixby Ranch Corporation for granting us access to 

Government Point, the Hearst Corporation for access to Pt. Sierra Nevada, BLM for access to Piedras 

Blancas, Hollister Ranch for access to Alegria, Carpinteria State Beach for access to our site there, and 

Vandenberg Air Force Base for access to Boat House, Stairs, Purisima and Occulto. We would like to 

thank David Pryor for providing access and support for our work at Crystal Cove State Park, and Harry 

Helling and Jon Lewengrub of the Orange County Marine Institute for providing assistance and support for 

our work at Dana Point. 

5
Technical Summary 

Study Title: Inventory of Rocky Intertidal Resources in San Luis Obispo, Santa 

Barbara, Ventura, Los Angeles, and Orange Counties. 

Report Title: Monitoring of Rocky Intertidal Resources Along the Central and Southern 

California Mainland. Comprehensive Report for San Luis Obispo, Santa Barbara, 

Ventura, Los Angeles, and Orange Counties (1992-2003) 

Contract Number: 1435-01-02-CA-85144

Sponsoring OCS Region: Pacific 

Applicable Planning Area: Central and Southern California 

Fiscal Years of Project Funding: FY1999, FY2000, FY2001, FY2002, FY2003 

Completion Date of the Report: November 2005 

Costs: FY1999 - $200,504, FY2000 - $157,000, FY2001 - $477,000, FY2002 - 

$493,000, FY2003 - $300,000 

Cumulative Project Cost: $1,627,504 

Principal Investigators: Peter T. Raimondi, Richard F. Ambrose, John M. Engle, 

Steven N. Murray 

Key Words: Intertidal, Monitoring, California, Marine Algae, Marine Invertebrates, 

Seasonal Trend, Temporal Trend, Baseline Data, Withering Syndrome, Abalone, 

Surfgrass, Mussel, Barnacle, Limpet, Seastar, Anemone, Rockweed, Turfweed, Whelk, 

Snail, Chiton.

Background and Objectives: 

This report presents the results of nine years of monitoring rocky intertidal resources at 

six sites in San Luis Obispo County, twelve years at nine sites in Santa Barbara County, 

ten years at five sites in Ventura and Los Angeles Counties, and eight years at four sites 

in Orange County. Data for ten years from two additional sites on Catalina Island are 

summarized in appendix A. These sites are part of MARINe (Multi-Agency Rocky 

Intertidal Network), a regional intertidal monitoring network sponsored by the Minerals 

Management Service (MMS), with additional funding and support from local and state 

governments, universities, and private organizations (see www.marine.gov). The 

MARINe group includes sites in San Diego, Northern California, Oregon, and the 

Channel Islands in addition to those presented in this report. This work developed out of 

concerns for the protection of rocky intertidal resources and the need to evaluate impacts 

on rocky intertidal resources following an oil spill. Surveys began in Spring 1992 in 

6
Santa Barbara County, Fall 1994 in Ventura and Los Angeles Counties, Fall 1995 in San 

Luis Obispo County, and Fall 1996 in Orange County and were conducted through Fall 

2003. This work will continue indefinitely provided funding remains available. Goals of 

the monitoring program include: 

• Assessing the temporal dynamics of target species across multiple sites spanning 

a broad geographic region. 

• Providing information to assess impacts of an oil spill or other anthropogenic 

activities in the context of natural changes in intertidal populations and 

communities. 

• Determining morphological aspects (e.g. color ratios) and key parameters 

describing population status (e.g. size structure) of selected target species. 

• Detecting and documenting invasions, changes in species ranges, disease spread, 

and other events important to developing an understanding of the structure and 

function of rocky intertidal populations and communities. 

Description:

Long-term monitoring sites were sampled using fixed plots in target community 

assemblages; this approach is similar to that used by the Channel Islands National Park 

and MMS-funded intertidal studies by Kinnetics and Littler. Fixed plots allow the 

dynamics of rocky intertidal species to be monitored with reasonable sampling effort. 

Targeting key species or assemblages allows the sampling effort to focus on ecologically 

important components of the assemblage, and provides greater statistical power to detect 

changes over space or time. 

Permanent plots for long-term monitoring were established at twenty-four sites, 

stretching from Point Sierra Nevada in the north to Dana Point in the south. Sites are 

located approximately 10-30 kilometers apart and span nearly 600 kilometers of 

California coastline. 

Monitoring surveys targeted thirteen key species or species groups: rockweed (Silvetia

compressa), rockweed (Hesperophycus harveyanus), turfweed (Endocladia muricata), 

red algae Mastocarpus papillatus and Mazzaella spp., anemones (Anthopleura 

elegantissima/sola), barnacles (Chthamalus spp. and Balanus glandula), goose barnacles 

(Pollicipes polymerus), mussels (Mytilus californianus), surfgrass (Phyllospadix 

scouleri/torreyi), seastars (Pisaster ochraceus and Asterina miniata), owl limpets (Lottia 

gigantea), and black abalone (Haliotis cracherodii). Not all target species were sampled 

at each site. Some species were sampled in the field by counts or point-contact 

procedures, but most sampling was accomplished by photographing quadrats. The 

resulting digital images could then be scored in the lab for percent cover of both the 

target species and other general taxa included in the plots using a point contact method. 

Significant Results: 

The following is a brief summary of individual species trends. Species dynamics are 

discussed in more detail in the Results section. 

7
Silvetia declined slightly in cover at San Luis Obispo County (SLO) sites but remained 

stable in most other areas. Silvetia cover tended to be slightly higher in fall than in spring 

at many sites. At both sites where Hesperophycus was sampled, cover decreased 

substantially over time. Cover of Endocladia varied seasonally at most sites and inversely 

with Silvetia cover, with higher cover in spring than in fall. Barnacles and Silvetia were 

common in most Endocladia plots, and fluctuations in cover of each of these three 

species were inversely correlated with cover of the other two. Mastocarpus experienced a 

general decline while Mazzaella cover remained relatively stable over time at both sites 

where these algae were sampled. Anthopleura cover was relatively stable over time at all 

sites where this species was sampled. Extreme fluctuations in sand cover occurred at 

three sites in the low-lying fixed plots used for monitoring Anthopleura abundance. 

Barnacle cover varied over time and among sites, which was not surprising for these 

relatively short-lived species. Barnacles declined substantially at Occulto, where plots 

experienced increases in algae and mussel cover, and at Stairs, where barnacle 

recruitment has not occurred since the 1997/98 El Niño. Pollicipes cover was constant 

over time at two sites, but showed both a decreasing over-time trend and a seasonal trend 

(with higher cover in fall) at Carpinteria. Mussel cover was relatively stable over time at 

nine of the nineteen sites where it is sampled. Four sites showed a decreasing trend in 

mussel cover over time. Recovery plots were largely dominated by bare rock and noncoralline crusts, but several other algal species and a few acorn barnacles were also 

present. 

Surfgrass cover was highly seasonal, with higher cover in fall than in spring at all sites 

except Cayucos, where transects are located in pools. Sand cover in surfgrass plots was 

an important factor at several sites, particularly at Coal Oil Pt., where plants were 

frequently partially covered and, on one occasion, completely buried by sand. Surfgrass 

cover along transects declined over time at Stairs and Carpinteria although recovery was 

observed at both sites. 

Seastar counts were variable from sample to sample at all sites but the color ratio of 

purple/brown to orange Pisaster (3:1) has remained relatively constant. On average, 

orange individuals tended to be larger than purple/brown individuals. Both Lottia

abundance and mean size were relatively constant over time at most sites. Dramatic 

declines in black abalone numbers continued at the Santa Barbara County sites and 

spread north to the San Luis Obispo County sites. These declines were coupled with a 

near-absence of abalone recruitment, indicating that recovery, if it occurs, will be slow. 

Although abalone numbers at Piedras Blancas and Pt. Sierra Nevada (the northernmost 

SLO abalone sites) had not decreased substantially by the end of this study period, these 

sites have since been re-visited, and the numbers of black abalone appear to be on the 

decline. 

A new addition to our sampling protocol was the counting of motile invertebrates in 

selected photoplots. Limpets and littorines were the most abundant motile invertebrates, 

present mostly in barnacle and Endocladia plots. The turban snail, Tegula funebralis, was 

common in Silvetia, Endocladia, and mussel plots. Mussel plots also contained the 

whelk, Nucella spp. and the chiton Nuttallina spp. Lepidochitona hartwegii was found 

nearly exclusively in Silvetia plots. 

8
Study Products: 

Presentations (1998-2004)

Ambrose, R.F. 1998. (panelist) Biodiversity along the coast, “What is Happening Now.” Conference on 

California’s Biodiversity Crisis: The Loss of Nature in an Urbanizing World, UCLA Institute of the 

Environment.

Ambrose, R.F. 2004. "Understanding Rocky Intertidal Communities Through Long-Term Monitoring: The 

MARINe Experience.” CEA-CREST Annual Environmental Science Conference, Pasadena, CA. 

Ambrose, R.F. 2005. “Protecting Rocky Intertidal Resources.” Santa Monica Bay Restoration 

Commission’s State of the Bay Conference, Los Angeles, CA. 

Becker, B.J. 2000. (poster) MARINE: Multi-Agency Rocky Intertidal Network, coordinated ecological 

monitoring on a regional scale. Biodiversity Council, Santa Barbara, CA. 

Blanchette, C.A., J. Kovach, C. Svedlund, and A. Wyndham (UCSB) & P. Raimondi, E. Maloney, K.

Kusic, A. Kendall, and M. Williams (UCSC). 2002. (poster) Biological patterns at the large scale: 

rocky intertidal community structure of the west coast. PISCO Public Symposium, California State 

University Monterey Bay, Monterey, CA. 

Blanchette, C.A., PT. Raimondi, D. Lohse, M. George, A. Kendall, K. Kusic, E. Maloney, M. Williams, 

and M. Wilson. 2002. Long-term large scale patterns in rocky intertidal communities: connecting 

pattern and process. California and the World Ocean 2002 Conference, Santa Barbara, CA. 

Blanchette, C.A., PT. Raimondi, M. Wilson, D. Lohse, A. Kendall, K. Kusic, H. Livingston, E. Maloney

and M. Williams. 2003. Beyond biogeography: large-scale patterns of distribution and abundance of 

intertidal marine algae along the US west coast. Phycological Society of America Annual Meetings, 

Salishan, OR. Abstract in J. Phycol. 39 (1):4. 

Bullard, A.M. and S.N. Murray. 2003. Changing macrophyte abundances and primary productivity of a 

southern California shore. Southern California Academy of Sciences, Northridge, CA. Abstract in So. 

Cal. Acad. Sci. Bull. 102 (2):26. 

Bullard, A.M. and S. N. Murray. 2003. (poster) Comparisons of macrophyte cover and community primary 

productivity on two southern California shores. Phycological Society of America Annual Meetings, 

Salishan, OR. Abstract in J. Phycol. 39 (1):6. 

Bullard, A.M. and S.N. Murray. 2003. Variations in community primary productivity on two southern

California rocky shores. Western Society of Naturalists, Long Beach, CA. 

Bullard, A.M. and S.N. Murray. 2004. Community Primary Productivity on Two Southern California 

Shores. Coloquio sobre Ficología, Acuacultura y Ecologia Marina, Universidad Autonoma de Baja

California Norte, Ensenada, Mexico. 

Bullard, A.M. and S.N. Murray. 2004. Shifts in Seaweed Abundances Can Greatly Affect Community 

Primary Productivity. Western Society of Naturalists 85th Annual Meeting, Rohnert Park, CA. 

Bullard, A.M. and S.N. Murray. 2004. Shifting Macrophyte Abundances and the Primary Productivity of

Southern California Shores. Phycological Society of America Annual Meetings, Williamsburg, VA. 

Bullard, A.M. and S.N. Murray. 2004. Net Primary Productivity of Southern California Rocky Intertidal 

Communities: Potential effects of shifts in macrophyte abundances. Northwest Algal Society 

Symposium, Bamfield, BC, Canada. 

Denis, T. and S.N. Murray. 2001. Among-site variation in the effects of trampling disturbance on Silvetia

compressa (O. Fucales) populations. Phycological Society of America , Estes Park, Colorado (abstract 

in J. Phycol. 37:16). 

Dunaway, M.E. 2002. (poster) MARINE: The Multi-Agency Rocky Intertidal Network. Southern

California Academy of Sciences, Claremont, CA. Abstract in So. Cal. Acad. Sci. Bull. 101 (2):17. 

9
Dunaway, M.E. 2002. (poster) Long-term marine monitoring - MARINE – a model regional program.

California and the World Ocean 2002 Conference, Santa Barbara, CA. 

Dunaway, M.E. 2002. (poster) MARINE – Multi-Agency Rocky Intertidal Network; a model regional longterm monitoring program. Western Society of Naturalists, Monterey, CA. 

Dunaway, M.E. 2004. MARINe – the Multi-Agency Rocky Intertidal Network; a long-term partnership that 

works. Seventh Biennial Workshop on Research within the Gulf of the Farallones, San Francisco, CA.

Dunaway, M.E. and D.V. Richards. 2003. MARINe – a successful partnership. Department of the Interior

Conference on the Environment, Phoenix, AZ. 

Dunaway, M.E. and D.V. Richards. 2003. (poster) MARINe: a model regional partnership for long-term

coastal monitoring. Joint Ventures: Partners in Stewardship Conference, Los Angeles, CA. 

Engle, J.M. 2001. Marine Life Monitoring in Southern California. USC/Catalina Conservancy Monitoring 

Symposium, Long Beach Aquarium, Long Beach, CA. 

Engle, J.M. 2002. WSN “Naturalist of the Year” presentation. Western Society of Naturalists, Monterey, 

CA. 

Gilbane, L. 2003. (poster) Using Carbon (13C) and Nitrogen (15N) stable isotope signatures to analyze food 

web inputs into mussels on southern California rocky shores. Western Society of Naturalists, Long

Beach, CA. 

Gilbane, L. and S. Murray. 2003. (poster) Analysis of carbon (13C) and nitrogen (15N) stable isotope

signatures of inputs into benthic food webs on southern California rocky shores. Southern California 

Academy of Sciences, Northridge, CA. Abstract in So. Cal. Acad. Sci. Bull. 102 (2):31-32

Gilbane, L. and S. N. Murray. 2004. Using Carbon (13C) and Nitrogen (15N) Stable Isotope Signatures to

Analyze Food Web Inputs into Mussels on Southern California Rocky Shores. Coloquio sobre 

Ficología, Acuacultura y Ecologia Marina, Universidad Autonoma de Baja California Norte, 

Ensenada, Mexico. 

Gilbane, L. and S. N. Murray. 2004. A Carbon (δ13C) and Nitrogen (δ15N) Stable Isotope Analysis of 

Macrophyte Contributions to Mussel (Mytilus californianus) Diets on Southern California Shores. 

Phycological Society of America, Annual Meetings, Williamsburg, VA 

Gilbane, L. and S. N. Murray. 2004. The role of macrophytes in the diets of the suspension feeding mussel

Mytilus californianus on southern California shores: A Carbon (δ13C) and Nitrogen (δ15N) Stable 

Isotope Analysis. Southern California Academy of Sciences Meetings, Long Beach, CA:

Gilbane, L. and S. N. Murray. 2004. Sources of primary production in suspension-feeding mussels (Mytilus 

californianus) from urban southern California shores. 4th International Conference on Applications of

Stable Isotope Techniques to Ecology, New Zealand 

Goodson, J. and Murray, S.N. 1999. Long-term changes in the abundance of rocky intertidal populations at 

Little Corona Del Mar, California: a synthesis with traditional and non-traditional data sources. 

Western Society of Naturalists, Monterey, CA. 

Henkel, S.K. and S. N. Murray. 2003. Reproduction, recruitment, and Morphological variation in lower

intertidal populations of the kelp Egregia menziesii (O. Laminariales). Northeast Algal Symposium,

Saratoga Springs, NY. 

Henkel, S.K. and S. N. Murray. 2003. Patterns of reproduction and morphological variation in southern

California populations of the lower intertidal kelp: Egregia menziesii (Turner) Areschoug. 

Phycological Society of America Annual Meetings, Salishan, OR. Abstract in J. Phycol. 39 (1):23. 

Kido, J.S. and S.N. Murray. 1998. Status of owl limpets (Lottia gigantea) in southern California habitats

influenced by collecting pressure. Western Society of Naturalists, San Diego, CA. 

Kido, J.S. and S.N. Murray. 1998. Human impacts on the size structure, size composition, growth, and

reproductive output in southern California populations of the intertidal protandrous limpet Lottia

gigantea (Gray). Southern California Academy of Sciences, Pomona, CA. 

10
Koehnke, J.M. and S.N. Murray. 1999. Seasonal and spatial patterns of reproduction and recruitment in a 

population of the intertidal rockweed Pelvetia compressa (Fucales). Phycological Society of America, 

St. Louis, MO (Abstract in J. Phycol. 35:17).

Koehnke, J.M. and S.N. Murray. 1999. Temporal and spatial variation in reproduction and recruitment of

Pelvetia compressa. Southern California Academy of Sciences, California State University, 

Dominguez Hills. 

Kusic, K.E., P.T. Raimondi, A. Kendall, D. Lohse, E. Maloney, M. Williams, and M. Wilson. 2003. 

Comparison of biodiversity patterns among rocky intertidal communities. 7th Biennial Workshop on

Research in the Gulf of the Farallones, San Francisco, CA. 

Kusic, K.E., P.T. Raimondi, C.A. Blanchette, A. Kendall, D. Lohse, E. Maloney, M. Williams, and M. 

Wilson. 2003. Comparison of biodiversity patterns among rocky intertidal communities. Sixth

California Islands Symposium, Ventura, CA. 

Lee, S.F. and M. Hill. 2004. Comparison of methods for obtaining low-altitude aerial images of rocky 

intertidal habitats. CEA-CREST Annual Environmental Science Conference. Pasadena, CA

Minchinton, T.E., P.T. Raimondi, M. Wilson, R.F. Ambrose, and J.M. Engle. 2000. (poster) Continued 

declines of black abalone due to withering syndrome along the coast of California. Thirteenth Annual 

Research Symposium of the U.C. Toxic Substances Research and Teaching Program. 

Murray, S.N. 1998. Human influences on the coastal ocean: Visitor impact on rocky shores: are marine 

protected areas really protected? Monterey Bay National Marine Sanctuary Symposium, Santa Cruz,

CA. 

Murray, S.N. 1998. Visitor impacts on rocky shores: considerations for designing effective marine reserves. 

California Cooperative Oceanic Fisheries Investigations, Pacific Grove, CA. 

Murray, S.N. 1998. Visitor impact on rocky shores: are marine protected areas really protected? Monterey

Bay National Marine Sanctuary Symposium: Human Influences on the Coastal Ocean, Santa Cruz, 

CA 

Murray, S.N. 1999. The Need for Long-term Data Sets and the Ability to Map and Detect Changes in the 

Abundance of Rocky Intertidal Populations in Southern California. Second CCD Monitoring

Workshop, Santa Catalina Island, CA. 

Murray, S.N. 2000. Visitor Impacts on Rocky Shores: Are Rocky Intertidal Reserves Really Reserves in 

Southern California? MacMillan Coastal Biodiversity Workshop, Bamfield Marine Station

Murray, S.N. 2001. Visitor impact on rocky shores: are southern California’s marine protected areas

protecting coastal resources? Southern California Academy of Sciences, Los Angeles, CA. 

Murray, S.N., J. Goodson, A. Gerrard, and T. Luas. 2001. Long-term changes in rocky intertidal seaweed

populations in urban southern California. Phycological Society of America, Estes Park, CO (abstract

in J. Phycol.: 37:35). 

Murray, S.N., J. Goodson, A. Gerrard, and T. Luas. 2001. Changes in southern California macrophyte

populations since Dawson’s rocky intertidal surveys in the late 1950s. Western Society of Naturalists, 

Ventura, CA. 

Murray, S.N. 2002. Are southern California’s changing coastal waters and nearshore biological

communities becoming more susceptible to invasion by exotic seaweeds? Southern California 

Academy of Sciences, Claremont, CA. Abstract in So. Cal. Acad. Sci. Bull. 101 (2):30.

Murray, S. N. 2003. Changing patterns in the abundances of rocky intertidal seaweed population in the 

Southern California Bight over a fifty-year period. Third European Phycological Congress, Belfast,

Northern Ireland. 

Murray, S. N. 2003. Visitor impacts on rocky shores. Are southern California’s marine protected areas 

protecting coastal resources? Seminar. Irish National University, Galway, Ireland. 

11
Murray, S. N. 2003. Changing environmental conditions and changes in the abundances of rocky intertidal 

seaweeds on southern California shores. Phycological Society of America Annual Meetings, Salishan, 

OR. Abstract in J. Phycol. 39 (1):43. 

Murray, S. N. 2004. Long-term Trends in the Abundances of Rocky Intertidal Seaweeds on Southern 

California Shores. Coloquio sobre Ficología, Acuacultura y Ecologia Marina, Universidad Autonoma

de Baja California Norte, Ensenada, Mexico. 

Murray, S. N. 2004. Changing Abundances of rocky intertidal seaweeds on southern California shores. 

Fifth Annual CEA-CREST Annual Conference, Pasadena, Callifornia. 

Murray, S. N. 2004. Do MPAs protect rocky intertidal communities from human impacts on southern

California shores? Seminar, Cal Poly Pomona. 

Murray, S. N. 2004. Visitor impacts on rocky shores: are southern California’s marine protected areas 

protecting coastal resources? Seminar, NOAA Offices, Long Beach 

Murray, S. N. 2004. Visitor impacts on rocky shores: are southern California’s marine protected areas 

protecting coastal resources? Seminar, California State University, Los Angeles.

Murray, S. N. 2004. Changing abundances of rocky intertidal seaweeds on southern California shores. 

Seminar. Friday Harbor Laboratories, University of Washington. 

Murray, S. N. 2005. Science, Policy, and Protecting our Coastal Oceans. Special Lecture. Center for Ocean 

Sciences Education Excellence – West, California State University, Northridge. 

Raimondi, P.T. 1999. (seminar) Assessment of Impacts to Rocky Intertidal Biota: Torch/Platform Irene

Pipeline Oil Spill, September 1997, Santa Barbara County, CA. 

Raimondi, P.T. 1999. (seminar) Aspects of monitoring. Ocean Sciences Department, University of

California, Santa Cruz, CA. 

Raimondi, P.T. 2001. (seminars) Assessing variability in intertidal community structure. California 

Polytechnic University San Luis Obispo, California State University Humboldt, Bodega Marine 

Laboratory, Oregon State University, and Duke University. 

Raimondi, P.T. 2002. Intertidal monitoring: its uses and abuses. PISCO Public Symposium, California 

State University Monterey Bay, Monterey, CA. 

Raimondi, P.T. 2002. Techniques of and lessons from intertidal monitoring. Research Activity Panel to

Monterey Bay National Marine Sanctuary.

Raimondi, P.T. 2003. (seminars) Unexpected dynamism in zonation and abundance revealed by long-term

monitoring on rocky shores. University of California Santa Cruz and Moore Foundation.

Raimondi, P.T. 2003. Unexpected dynamism in zonation and abundance revealed by long-term monitoring 

on rocky shores. Moss Landing Marine Laboratories. 

Raimondi. P.T. 2003. Ecological effects due to impingement and entrainment. State of California 

Desalinization Working Group.

Raimondi. P.T. 2003. Overview of PISCO. Gordon and Betty Moore Foundation.

Raimondi, P.T. 2003. Unexpected dynamism in zonation and abundance revealed by long-term monitoring 

on rocky shores. Marine Interest Group. 

Raimondi, P.T. 2003. Unexpected dynamism in zonation and abundance revealed by long-term monitoring 

on rocky shores. Monterey Bay National Marine Sanctuary.

Raimondi, P.T. 2004 Long Term monitoring of intertidal communities. National Park Service Monitoring 

Task Force, Oakland, CA

Raimondi, P.T. and M.C. Carr. 2000. Partnership for Interdisciplinary Studies of Coastal Oceans. PISCO at 

Santa Cruz. Larval Biology Meeting. University of California, Santa Cruz, CA. 

12
Raimondi, P.T. and M.C. Carr. 2001. Long-term patterns and processes in temperate rocky reef

communities: stasis is not the norm. Western Society of Naturalists, Ventura, CA. 

Raimondi, P.T., D. Lohse and C. Blanchette. 2003. Unexpected dynamism in zonation and abundance

revealed by long-term monitoring on rocky shores. Symposium on “Expanding Scales, Expanding 

Perspectives: New Insights into Marine Ecosystem Dynamics”, Ecological Society of America 88th

Annual Meeting, Savannah, GA. 

Raimondi, P.T, R. Sagarin, R. Ambrose, M. George, S. Lee, D. Lohse, C. M. Miner, S. Murray, and C. Roe 

2005. Color change and consistency in the sea star Pisaster ochraceus. Society for Integrative and 

Comparative Biology, San Diego, CA. 

Raimondi, P.T, R. Sagarin, R. Ambrose, M. George, S. Lee, D. Lohse, C. M. Miner, S. Murray, and C. Roe 

2004. Color change and consistency in the sea star Pisaster ochraceus. Western Society of Naturalists, 

Rohnert Park, CA. 

Readdie, M. 2000. (poster) Long-term change in intertidal zonation. Can succession drive vertical shifts in

species zones? Monterey Bay National Marine Sanctuary Symposium, Monterey, CA. 

Readdie, M. 2002. (poster) Long-term change in intertidal zonation. Can succession drive vertical shifts in

species zones?. PISCO Public Symposium, California State University Monterey Bay, Monterey, CA. 

Readdie, M. 2003. Shifting zones: how species upper limits can vary vertically on rocky shores, 6th

International Temperate Reef Symposium, Christchurch, NZ. 

Readdie, M. 2003. Shifting zones: how facilitation can cause species upper limits to vary vertically on

rocky shores, Western Society of Naturalists, Long Beach, CA. 

Roe, C. and P. Raimondi. 2001. Variability in the accumulation and persistence of tar in four intertidal 

zones. Western Society of Naturalists, Ventura, CA. 

Sagarin, R. 2003. The heat shock response is complex, but so is the intertidal. Western Society of 

Naturalists, Long Beach, CA. 

Sagarin, R. 2004. (seminar) Climate change, species change and the power of the naturalist. Scripps 

Institution of Oceanography, San Diego, CA. 

Sagarin R., R. Ambrose, B. Becker, J. Engle, S. Murray, P. Raimondi, D. Richards. 2004. Using monitoring

to study unpredictable, high impact events: effects of human collection of the intertidal limpet Lottia

gigantea. CEA-CREST Annual Environmental Science Conference. Pasadena, CA

Sagarin R., R. Ambrose, B. Becker, J. Engle, S. Murray, P. Raimondi, D. Richards. 2004. Using monitoring

to study unpredictable, high impact events: effects of human collection of the intertidal limpet Lottia

gigantea. Western Society of Malacologists. Annual Meeting. Ensenada, Mexico. 

Smith, J.R., R.F. Ambrose, and P. Fong. 2003. Long-term change in mussel (Mytilus californianus) 

communities along the coast of California. Western Society of Naturalists, Long Beach, CA. 

Smith, J.R., R.F. Ambrose, and P. Fong. 2003. (poster) Current condition and long-term change in the 

abundance and biodiversity of mussel bed communities of wave-exposed rocky intertidal zones of the 

Channel Islands. Sixth California Islands Symposium, Ventura, CA. 

Whiteside, K. and S. N. Murray. 2004. Spatial and temporal patterns of abundance in southern California 

populations of Caulacanthus ustulatus (Rhodophyta). Phycological Society of America, Annual 

Meetings, Williamsburg, VA 

Wilson, M. 2001. Is Sargassum muticum a benign invader of tidepools on the Pacific coast? Western

Society of Naturalists, Ventura, CA. 

Wilson, C.M., J.M. Altstatt, P.T. Raimondi, and T.E. Minchinton. 2002. Changes in intertidal community

structure following mass mortality of the black abalone, Haliotis cracherodii, and implications for 

abalone recovery. Western Society of Naturalists, Monterey, CA. 

13
Wilson, C.M., J.M. Altstatt, P.T. Raimondi, and T.E. Minchinton 2004. Changes in intertidal community

structure following mass mortality of the black abalone, Haliotis cracherodii, and implications for 

recovery. NOAA ESA Workshop on Status Review of Black Abalone, La Jolla, CA. 

Publications (1998-2004)

Ambrose, R.F. 2002. Sampling design. Pp. 67-90 in: Methods for performing monitoring, impact, and

ecological studies on rocky shores (S.N. Murray, R.F. Ambrose, and M.N. Dethier, eds.). OCS Study

MMS 01-070, U.S. Minerals Management Service, Pacific OCS Region. 

Ambrose, R.F. 2002. Transects, quadrats, and other sampling units. Pp. 91-115 in: Methods for performing

monitoring, impact, and ecological studies on rocky shores (S.N. Murray, R.F. Ambrose, and M.N.

Dethier, eds.). OCS Study MMS 01-070, U.S. Minerals Management Service, Pacific OCS Region. 

Chambers Group, Inc. 2000. Santa Barbara County Shoreline Inventory. U.S. Minerals Management

Service, Pacific OCS Region. 

Dunaway, M.E., R.A. Ambrose, J. Campbell, J.M. Engle, M. Hill, Z. Hymanson, and D.V. Richards. 1998. 

Establishing a Southern California rocky intertidal monitoring network. Pp. 1278-1294 in: California

and the World Ocean ’97 (O.T. Magoon, H. Converse, B. Baird, & M. Miller-Henson, eds). American

Society of Civil Engineers, Reston, Virginia. 

Forde, S. 2002. Modelling the effects of an oil spill on open populations of intertidal invertebrates. Journal 

of Applied Ecology 39:595-604. 

Forde, S.E. and P.T. Raimondi. 2004 An experimental test of the effects of variation in recruitment

intensity on intertidal community structure. Journal of Experimental Marine Biology and Ecology

301:1-14 

Henkel, S. K. and S. N. Murray. (under revision). Reproduction and morphological variation in southern

California populations of the lower intertidal kelp Egregia menziesii (O. Laminariales). Journal of

Phycology 

Kido, J.S. 2000. Variations in the structure of Lottia gigantea Sowerby (owl limpet) populations among and

within sites on southern California rocky shores. M.S. Thesis, California State University, Fullerton, 

California. 

Kido, J.S. and S.N. Murray. 2003. Variation in owl limpet Lottia gigantea population structures, growth

rates, and gonadal production on southern California rocky shores. Mar. Eco. Prog. Ser.257:111-124. 

Menge B. A., C.A. Blanchette, T.L. Freidenburg, S.D. Gaines, J. Lubchenco, D. Lohse, P. Raimondi. 2001.

Cross-scale linkages between bottom-up factors and interaction strength in rocky intertidal

communities. [Meeting] Ecological Society of America Annual Meeting Abstracts. [print] 86. 157. 

Miller, A.W. and R.F. Ambrose. 2000. Sampling patchy distributions: comparison of different sampling 

designs in rocky intertidal habitats. Mar. Ecol. Prog. Ser. 196:1-14.

Miner, C.M., J.M. Altstatt, P.T. Raimondi, T.E. Minchinton (in prep). Shifts in habitat structure following

mass mortality of a threatened species limits its prospects for recovery. 

Miner, C.M. (in prep). Is Sargassum muticum a benign invader of tidepools on the Pacific coast of North 

America? 

Moeller, J.M. and S.N. Murray. (under revision). Seasonal and spatial patterns of reproduction and

recruitment in a southern California population of the intertidal rockweed Silvetia compressa (O. 

Fucales). J. Phycol.

Murray, S.N. 1998. Effectiveness of marine life refuges on southern California shores. Pp. 1453-1465 in:

California and the World Ocean '97 (O.T. Magoon, H. Converse, B. Baird, & M. Miller-Henson, 

eds.). American Society of Civil Engineers, Reston, VA.

Murray, S.N. 1999. Are marine life refuges effective? Tidelines 19:6-7. 

14
Murray, S.N. 2002. Designing rocky intertidal monitoring and impact field studies: a brief overview. Pp. 1-

14 in: Methods for performing monitoring, impact, and ecological studies on rocky shores (S.N. 

Murray, R.F. Ambrose, and M.N. Dethier, eds.). OCS Study MMS 01-070, U.S. Minerals

Management Service, Pacific OCS Region. 

Murray, S.N. 2002. Quantifying abundance: density and cover. Pp. 116-147 in: Methods for performing

monitoring, impact, and ecological studies on rocky shores (S.N. Murray, R.F. Ambrose, and M.N.

Dethier, eds.). OCS Study MMS 01-070, U.S. Minerals Management Service, Pacific OCS Region. 

Murray, S.N. 2002. Quantifying abundance: biomass. Pp. 148-162 in: Methods for performing monitoring, 

impact, and ecological studies on rocky shores (S.N. Murray, R.F. Ambrose, and M.N. Dethier, eds.).

OCS Study MMS 01-070, U.S. Minerals Management Service, Pacific OCS Region. 

Murray, S.N. 2002. Individual-based parameters: age-determination, growth rates, size-structure, and 

reproduction. Pp. 116-147 in: Methods for performing monitoring, impact, and ecological studies on

rocky shores (S.N. Murray, R.F. Ambrose, and M.N. Dethier, eds.). OCS Study MMS 01-070, U.S. 

Minerals Management Service, Pacific OCS Region. 

Murray, S.N., T.G. Denis, J.S. Kido, and J.R. Smith. 1999. Frequency and potential impacts of human 

collecting in rocky intertidal habitats in southern California marine reserves. CalCOFI Rep. 40:100-

106. 

Murray, S.N., R.F. Ambrose, and M.N. Dethier. 2002. Methods for performing monitoring, impact, and 

ecological studies on rocky shores. OCS Study MMS 01-070, U.S. Minerals Management Service,

Pacific OCS Region. 

Murray, S. N., R. F. Ambrose, and M. N. Dethier. 2005. Monitoring Rocky Shores. University of

California Press, Berkeley (contracted and submitted for publication).

Raimondi, P.T. R. Sagarin, R. Ambrose, M. George, S. Lee, D. Lohse, C. M. Miner, S. Murray, C. Roe 

(submitted). Color change and consistency in the sea star Pisaster ochraceus. 

Raimondi, P.T., C.M. Wilson, R.F. Ambrose, J.M. Engle, T.E. Minchinton. 2002. Continued declines of

black abalone along the coast of California: are mass mortalities related to El Niño events? Marine

Ecology Progress Series 242:143-152. 

Roy, K., A.G. Collins, B.J. Becker, E. Begovic, and J.M. Engle. 2003. Anthropogenic impacts and 

historical decline in body size of rocky intertidal gastropods in southern California. Ecology Letters 

6:205-211. 

Sagarin, R. D., R. F. Ambrose, B. J. Becker, J. M. Engle, J. Kido, S. F. Lee, C. M. Miner, S. N. Murray, P. 

T. Raimondi, D. V. Richards, C. Roe (submitted). Effects of human foraging on the limpet Lottia 

gigantea across California rocky intertidal shores. 

Sapper, S.A. 1998. Variation in an intertidal subcanopy assemblage dominated by the rockweed Pelvetia

compressa (Phaeophyceae, Fucales). M.S. Thesis. California State University, Fullerton, California. 

Sapper, S.A. and S.N. Murray 2003. Variation in structure of the subcanopy assemblage associated with 

southern California populations of the intertidal rockweed Silvetia compressa (Fucales). Pacific 

Science 57(4):433-462. 

Sato, L.M. and S.N. Murray. (under revision). Variations in the abundances and structures of intertidal 

Tegula gallina and T. funebralis (Trochidae) populations on southern California rocky shores. J. Exp. 

Mar. Bio. Eco. 

Smith, J. R. and S. N. Murray. 2005? The effects of experimental bait collection and trampling on a 

southern California Mytilus californianus Conrad bed. Mar. Biol. (in press). 

Smith, J.R. and S. N. Murray. The effects of recreational fishers on Mytilus californianus Conrad beds on

southern California rocky coasts. Mar. Envir. Research (under review). 

15
Page intentionally left blank 

16
INTRODUCTION 

The central/southern California coast possesses an exceptional diversity of valuable rocky 

intertidal resources. Major factors contributing to the richness of coastal marine life in 

this region include its location along the boundary of two major biogeographic provinces 

(cold-temperate Oregonian and warm-temperate Californian), a high diversity of habitat 

types, and exposure to varying local oceanographic conditions. Oil and gas activities, 

especially the transport of oil along the California coast, raise the possibility of an oil 

spill or other impact to coastal resources. Population monitoring of coastal biota in 

central and southern California provides baseline information in case an event such as a 

spill were to damage these resources. This baseline information is essential for scientific 

studies investigating the short- and long-term effects of a spill and for natural resource 

damage assessment. In addition, well-designed monitoring studies yield important data 

on population dynamics on local and regional scales, which can be utilized for more 

effective resource management, as well as provide fundamental ecological knowledge 

about the dynamics of the systems. Moreover, quantitative data describing the dynamics 

of key intertidal populations over a broad, geographic scale are essential for robust testing 

of impacts of oil spills or other major disturbance events. 

Data from this study have thus far been used as the bases for advancing scientific 

understanding of rocky intertidal populations in the region as evidenced by more than 

125 presentations and publications. Of particular significance to MMS was the successful 

use of monitoring data in damage assessment, when a small oil spill occurred off the 

coastline of northern Santa Barbara County in 1997 (see Raimondi et al. 1999). 

Additionally, we have documented disease in black abalone and the resulting massive 

declines in population sizes at Santa Barbara and San Luis Obispo County sites. In 

Orange County, the red alga Caulacanthus ustulatus has become increasingly abundant. 

This species is thought to be a non-native invader and is rapidly becoming the dominant 

species in the mid-upper algal turf community. These are just a few examples of how 

long-term monitoring data obtained from the MARINe program are proving to be 

valuable to MMS and other regional constituents. 

REPORT ORGANIZATION 

This report is organized into four sections consisting of the main data summary section 

plus three appendices. The first section summarizes our findings from surveys done in 

San Luis Obispo, Santa Barbara, Ventura, Los Angeles, and Orange Counties from 1992-

2003 (surveys in some areas began after 1992). Appendix A is a summary of findings 

from surveys done at two sites on Catalina Island from 1994-2003. Appendix B contains 

natural history information about the target species sampled in this study. It is an updated 

version of the natural history section from a previous report (Raimondi et al. 1999), 

containing additional information from recent literature, and from our own field 

observations. Appendix C contains tables of summary data (mean percent covers and 

standard errors) for target and other common species in permanent plots at all sites. 

17
SAMPLING METHODS 

Only brief descriptions of key elements of the sampling program will be included here, 

including changes in procedures adopted since our last report (Raimondi et al. 1999). 

Specific methods for carrying out the MARINe monitoring program and site-specific 

information for the Santa Barbara County sites is given in the rocky intertidal monitoring 

handbook for Santa Barbara County (Engle et al. 1994). Sampling protocols can also be 

found on the MARINe website (www.marine.gov), and will be published in a revised 

intertidal monitoring handbook in 2005. Additional discussions of sampling issues crucial 

to the design of MARINe and other intertidal monitoring programs can be found in 

Murray et al. (2004). 

Criteria used in site selection include: 

• Areas previously surveyed or monitored that provide historical data

• Unsurveyed areas representing major data gaps

• Areas of concern with regard to human impacts, including potential oil spills

• Areas with relatively pristine habitats

• Areas which provide habitat for sensitive or rare intertidal species

• Areas with optimum conditions for long-term monitoring

Optimum conditions for monitoring include reasonable and safe site access, adequate 

bedrock surfaces for establishing permanent plots, sufficient abundance of key species, 

and minimal disturbance to sensitive resources (e.g., seabirds, marine mammals). 

Sampling of the 26 rocky intertidal sites in San Luis Obispo, Santa Barbara, Ventura, Los 

Angeles, and Orange Counties (Figure 1) follows the protocol described in Ambrose et 

al. (1992), which is modeled after methods used by the National Park Service intertidal 

monitoring program (Richards and Davis 1988), and focuses on target species or 

assemblages. Permanent photoplots were established in assemblages such as barnacles, 

mussels, anemones, turfweed, and rockweed. Cover of the major taxa within these 

photoplots was determined by point-contact analysis from photographs, except for 

barnacle plots in San Luis Obispo and Northern Santa Barbara Counties (see below). 

Permanent plots also were established for large motile species such as owl limpets, black 

abalone, and sea stars. Permanent line transects are used to estimate the cover of 

surfgrass. A series of photographic pans along with field notes are used to describe 

general conditions at the site and to document the distribution and abundance of 

organisms not found within the photoplots. 

Several changes have been made to our protocols since the 1998 report (Raimondi et al 

1999). First, in 2002 we switched from using slide film to taking digital images of our 

photoplot assemblages. This change significantly reduces the likelihood of getting 

unscorable images, because we are able to view images in the field. Now, instead of 

projecting a slide onto a grid to score percent cover of species in photoplots, we score 

plots on a computer monitor. We also use digital images for site pans, which have 

replaced the video footage previously taken at all sites. Starting in Spring 2001, we began 

scoring barnacle plots in the field at the eight northernmost sites. This allows us to 

distinguish Chthamalus spp. from Balanus glandula, and dead barnacles from live ones. 

At one site, Stairs, we added “recovery” plots, in which we are following natural 

18
recruitment and succession processes in areas where large sections of the layered rock 

were removed by the 1997/98 El Niño storms. Our seastar sampling protocol was 

modified slightly in 1996 (later at some sites), when we began measuring Pisaster 

ochraceus and recording color information. Pisaster colors can be quite variable, so for 

simplicity, we used two color categories: orange and other. An individual was classified 

as “orange” if it was bright orange, with little brown or purple pigment. Pisaster

classified as “other” included all purple and brown individuals, and everything in 

between. Finally, we began counting and measuring small motile invertebrates in the

photoplots at each site. Species targeted in these plots include Tegula funebralis, 

Acanthina spp., Nucella emarginata, N. canaliculata, Ocenebra circumtexta, 

Lepidochitona harwegii, Nuttallina spp., Mopalia spp., three species of Pagurus, 

Littorina spp., and various limpets. 

Another new development to the methods used by MARINe groups involves the way in 

which data are entered. In the past, each group kept separate spreadsheets, and although 

these were quite similar in structure, it was difficult to combine datasets due to 

differences in the use of species codes, or the way in which the data were organized. In 

2002-2003, the Southern California Coastal Water Research Project (SCCWRP), 

developed an Access database (Microsoft TM) for us, which contains all previous data, and 

is used for data entry by all MARINe groups. This has fully standardized our data entry 

methods, and makes it possible to easily extract data from the entire network of MARINe 

sites. In the near future, this database will also contain intertidal monitoring data from

sites north of San Luis Obispo County, which are funded by PISCO (Partnership for 

Interdisciplinary Studies of Coastal Oceans). 

19
multi-species site

black abalone only

(+ owl limpets at Rancho Marino)

N

Mussel Shoals

Old Stairs

Santa Barbara Co.

Crystal Cove

Shaw’s Cove

Treasure Is.

Dana Pt.

Orange Co.

San Luis Obispo Co.

Pt. Sierra Nevada

Piedras Blancas

Rancho Marino

Cayucos

Hazard’s

Shell Beach

Occulto

Purisima

Stairs

Boat House

Government Pt.

Alegria

Arroyo Hondo

Coal Oil Pt.

Carpinteria

Ventura Co.

Paradise Cove

White’s Point

Point Fermin

Los Angeles Co.

Little Harbor

Bird

Rock

0 50 km

multi-species site

black abalone only

(+ owl limpets at Rancho Marino)

N

Mussel Shoals

Old Stairs

Santa Barbara Co.

Crystal Cove

Shaw’s Cove

Treasure Is.

Dana Pt.

Orange Co.

San Luis Obispo Co.

Pt. Sierra Nevada

Piedras Blancas

Rancho Marino

Cayucos

Hazard’s

Shell Beach

Occulto

Purisima

Stairs

Boat House

Government Pt.

Alegria

Arroyo Hondo

Coal Oil Pt.

Carpinteria

Ventura Co.

Paradise Cove

White’s Point

Point Fermin

Los Angeles Co.

Little Harbor

Bird

Rock

multi-species site

black abalone only

(+ owl limpets at Rancho Marino)

N

Mussel Shoals

Old Stairs

Santa Barbara Co.

Crystal Cove

Shaw’s Cove

Treasure Is.

Dana Pt.

Orange Co.

San Luis Obispo Co.

Pt. Sierra Nevada

Piedras Blancas

Rancho Marino

Cayucos

Hazard’s

Shell Beach

Occulto

Purisima

Stairs

Boat House

Government Pt.

Alegria

Arroyo Hondo

Coal Oil Pt.

Carpinteria

Ventura Co.

Paradise Cove

White’s Point

Point Fermin

Los Angeles Co.

Little Harbor

Bird

Rock

0 50 km

Figure 1. Location of rocky intertidal monitoring sites along the central and southern 

 California Coast. 

20
DESCRIPTION OF REGIONS 

San Luis Obispo County 

Rocky intertidal communities in San Luis Obispo County (SLO) are well known for their 

diverse and relatively pristine biota. The majority of the 150 kilometer-long coast is 

privately-owned and undeveloped. The natural beauty and coastal resources of SLO 

County make it a popular tourist destination, as evidenced by more than 10 state and 

county parks and beaches. Fifty-eight percent of the shore consists of rugged rocky reefs 

that are fully or partially exposed to prevailing oceanic swells. Situated at the southern 

end of the cold-temperate Oregonian province, SLO intertidal habitats contain a unique 

mix of species, with warm-temperate species more characteristic of the southern 

(Californian) biogeographic province absent or declining and cold-temperate forms 

increasing in abundance compared with counties to the south. For example, warm-water 

sea palms (Eisenia arborea), rockweed (Hesperophycus harveyanus), barnacles 

(Tetraclita rubescens, Chthamalus fissus), and horse mussels (Brachidontes adamsianus) 

are less common or absent, while cold-water sea palms (Postelsia palmaeformis), 

rockweed (Fucus distichus, Pelvetiopsis limitata), barnacles (Balanus glandula, 

Chthamalus dalli), and horse mussels (Septifer bifurcata) appear or increase in 

abundance in SLO County. 

The rich rocky shore communities of SLO County are vulnerable to oil spills or impacts 

from other oil and gas operations, primarily from major coastal tanker traffic, but also 

from terminal operations at Estero Bay, onshore pipeline breaks, and future oil 

exploration leases. In recent years there have been spills affecting marine resources from 

onshore operations at Avila (in 1992) and Guadalupe (in 1994), but data on impacts 

remain confidential. Natural oil seeps also exist, resulting in the presence of tar on many 

rocky shores in the region (Kinnetics 1992). Prior to the MARINe study, population 

dynamics of rocky coast flora and fauna in SLO County were largely unstudied except 

for impact surveys associated with the Diablo Canyon Nuclear Power Plant located north 

of Avila Beach (North et al. 1989, Pacific Gas and Electric 1988, 1994, Schiel et al. 

2004), and research on seasonal and successional variation in intertidal community 

structure conducted at two sites (Point Sierra Nevada and Diablo Canyon, Kinnetics 

1992). The ongoing Diablo Canyon surveys, initiated in the 1970’s, represent an 

excellent time series for this area. The seasonal and successional studies at Point Sierra 

Nevada and Diablo Canyon were conducted for the Minerals Management Service during 

1985-1991; MMS biologists continued to monitor mussel recovery plots at Point Sierra 

Nevada until 1998. 

Santa Barbara County 

The Santa Barbara County (SBC) coastline is an important biogeographical transition 

area for rocky intertidal organisms because the west-facing shore north of Point 

Conception is subject to largely different oceanographic influences than the south-facing 

shore down coast of the Point. Although there is considerable overlap, there are distinct 

differences between the organisms north and south of Point Conception (Murray and 

Littler 1981, Ambrose 1992, Blanchette et al. 2002). For example, seaweed communities 

north of Point Conception are characterized by cold-water species such as laminarialean 

brown algae and large, fleshy red algae, and by greater biomass, whereas communities 

21
south of the Point are characterized by warm water fucalean brown algae and shorter, 

more densely branched red algae (Abbott and Hollenberg 1976, Murray and Bray 1993). 

Concerns about impact of oil spills in SBC stem from transport by offshore tanker and 

onshore pipeline, production platforms, and terminal operations. Natural oil seeps are 

prominent features, especially at Point Conception, Coal Oil Point, and Carpinteria State 

Beach. Previous studies in SBC include work by Littler and colleagues at Coal Oil Point 

and Government Point (Littler 1979) and Kinnetics at Government Point and some sites 

north of Point Conception (Kinnetics 1985); other studies are summarized in Chambers 

(1993). 

Los Angeles and Ventura Counties 

Occurring within the southern California bight, the oceanographic conditions along the 

Los Angeles and Ventura County (LA/VEN) coast are influenced by the presence of 

Point Conception and the Channel Islands offshore. South of Point Conception, the 

California current proceeds offshore and the orientation of the northern Channel Islands 

serves to shelter the bight from the prevailing northwesterly winds and swells, especially 

those that develop during fierce pacific winter storms. This results in relatively benign 

oceanographic conditions most of the time, though periodic southern storms can have a 

devastating impact on south facing stretches of the coast. A large gyre that exists within 

the bight creates sea surface temperatures that are warmer, on average, than coastal 

sections to the north and (to a certain extent) to the south. Sea conditions are especially 

warm on Santa Catalina and San Clemente Islands, in the center of the gyre, which are 

frequently inhabited by subtropical species normally found far to the south. 

In southern California, sandy habitats comprise a much greater proportion of the 

shoreline than they do in central and northern California. This is especially true in the

Santa Monica Bay where rocky intertidal habitats are uncommon and patchy. Most of the 

monitoring sites are located on isolated rocky habitats that are flanked by wide stretches 

of beach, except for those sites on the Palos Verdes Penninsula, which has more 

continuous rocky intertidal habitat. In addition, several of the sites are subject to widely 

fluctuating sand levels and many of the plots experience frequent periods of burial and 

scour. This, along with abundant sunshine and the predominance of warmer coastal air 

temperatures create harsh conditions for species that are intolerant to desiccation. Thus, 

the rocky intertidal communities are largely devoid of larger foliose algae such as the

fleshy reds that are so common to the north. These are replaced by abundant turf forming 

species such as Corallina vancouveriensis, Gelidium coulteri, Endocladia muricata, and 

numerous species of filamentous algae. Macroalgal diversity may be just as high, or 

higher than at the more northern sites, but algal biomass is substantially lower. 

The LA/VEN coastline is heavily urbanized and subject to multiple anthropogenic 

influences. At monitoring sites that are closer to urban centers, the direct influence of 

people on the rocky intertidal community is substantial. White’s Point may be one of the 

most heavily impacted sites on the California coast because public use is high, and 

regulation and enforcement are low to non-existent. Point Fermin also has high public 

use, but is a marine reserve with increased public and legal enforcement over human 

actions. However, a large number of school groups visit this site and like White’s Point, 

the impacts of human trampling are likely very high. At other LA/VEN sites, public 

22
access and/or direct use of the rocky intertidal resources are lower. The Port of Long 

Beach is a major shipping hub, and numerous oil refineries and oil extraction platforms 

exist locally. The threat of an oil spill from a transiting tanker ship is substantial, though 

no major spills have occurred along the LA/VEN coast in many years. 

Orange County 

The Orange County (OC) coastline is dominated by sandy beaches, particularly in the 

north, and rocky shores account for only a small percentage of the shore habitat. Much of 

the OC coastline south of Newport Bay consists of a composite of rocky headlands and 

pocket beaches backed by eroded bluffs and is semi-protected from predominant wave 

patterns by offshore islands (Hickey 1993). It is along this stretch of coastline where our 

MARINe sites are located. Because of the proximity of rocky habitats to sandy beaches 

and the movement of sand with longshore currents, most OC rocky shores are 

characterized by periodic sand deposition and scour. Although the region encompasses 

thirteen Marine Protected Areas with varying degrees of protection, OC shores are 

readily accessible to the public and are heavily used for recreational purposes throughout 

the year. Evidence indicates that OC rocky shores are strongly influenced by human 

activities, including the unlawful collecting of shore organisms in Marine Life Refuges 

and Ecological Reserves (Murray 1998, Murray et al. 1999). All OC MARINe sites 

except Treasure Island have had Marine Life Refuge status since 1969-71; the Laguna 

Beach Marine Life Refuge was extended to include the Treasure Island site in January 

1995. 

Prior to the MARINe project and related ecological studies initiated by Murray and 

colleagues over the past several years, little scientific research had been performed on OC 

rocky intertidal populations and communities. Historical knowledge is best developed for 

seaweeds based on surveys performed in the late 1950’s through the 1960’s by Dawson 

(1959, 1965), Widdowson (1971), Nicholson and Cimberg (1971), and Thom and 

Widdowson (1978). Previous quantitative studies in OC include research by Littler and 

colleagues at Dana Point and Little Corona del Mar (Littler 1979). 

The four OC sites are located in a region near production platforms and offshore tanker 

traffic servicing Los Angeles/Long Beach Harbors. Because of common southerly 

flowing nearshore currents, oil spills in coastal waters near LA/LB Harbors or from

production platforms off Seal Beach near the Los Angeles and Orange County border 

will likely impact OC rocky intertidal populations and communities located at one or

more of our MARINe sites. 

Temporal Coverage of Report 

Rocky intertidal monitoring sites were established in Santa Barbara County (SBC) in 

Spring 1992 (Ambrose et al. 1992). Sites in Los Angeles (LA) and Ventura (VEN) 

Counties were established in Fall 1994 (Engle et al. 1994b, 1995). San Luis Obispo 

County (SLO) sites were established in Fall 1995 and Orange County (OC) sites in Fall 

1996 (Engle et al. 1998). This combination of surveys has resulted in twelve years of data 

for SBC sites, ten years for LA and VEN sites, nine for SLO sites, and eight for OC sites. 

Survey results from Spring 1992 through Spring 1995 were reported in Ambrose et al. 

(1995) and were updated through Spring 1998 in Raimondi et al (1999). This report 

updates the monitoring results from Fall 1998 through Fall 2003. Trends discussed in 

23
24

this report apply to the entire monitoring period unless otherwise noted. A summary of 

the species monitored and methods used at each site is given in Table 1. 
25

Table 1. Summary of key species monitored at all sites, including survey methods and number of replicate pl

SPECIES PSN PB RM CAY HAZ SB OCC PUR ST BH GP AL AH COP CAR MS OS PC WP PF 

Silvetia 5

PP 

5

PP 

5 

PP 

5 

PP 

5

PP 

5 

PP 

5 

PP 

5

PP 

Hesperophycus 5 

PP 

5

PP 

Endocladia 5 5 

PP PP 

5 

PP 

5 

PP 

5

PP 

5 

PP 

5 

PP 

5

PP 

5 

PP 

5 

PP 

Mastocarpus 5 

PP 

5

PP 

Mazzaella 5 

PP 

5

PP 

Phyllospadix 3

PT 

3

PT 

3 

PT 

3 

PT 

3

PT 

3

PT 

3 

PT 

2 

PT 

3 

PT 

3 

PT 

3 

PT 

3

PT 

3

PT 

Anthopleura 5 

PP 

5

PP 

5

PP 

5 

PP 

5 

PP 

5 

PP 

Barnacles 5

PP 

5

PP 

5 

PP 

5 

PP 

5 

PP 

5

PP 

5 

PP 

5 

PP 

5 

PP 

5 

PP 

5

PP 

5 

PP 

5 

PP 

5 

PP 

5 

PP 

5 

PP 

Pollicipes 5 

PP 

5 

PP 

5

PP 

Mussels 5

PP 

5

PP 

5 

PP 

5 

PP 

5 

PP 

5

PP 

5 

PP 

5 

PP 

5 

PP 

5 

PP 

5

PP 

5 

PP 

5 

PP 

5 

PP 

5 

PP 

“Recovery” 7

PP 

Lottia 5 

CP 

5 

CP 

5 

RP 

5

CP 

6 

CP 

5 

CP 

5 

CP 

Haliotis 5 

IP 

4 

IP 

3 

IP 

3 

IP 

3

IP 

4 

BT 

3 

IP 

3 

IP 

Seastars 3

IP 

3

IP 

3 

IP 

3 

IP 

1 

IP 

3

IP 

3 

IP 

3 

IP 

3

BT 

3

BT 

Total Species Per Site 9 1 2 9 8 7 4 1 9 8 9 6 4 2 6 4 4 4 3 3 

Key to survey techniques: PP=Photoplot IP=Irregular Plot PT=Point-intercept Transect CP=Circular Plot RP=Rectangular Plot BT=Band TranseKey to site codes: PSN=Pt. Sierra Nevada PB=Piedras Blancas RM=Rancho Marino CAY=Cayucos HAZ=Hazards SB=Shell Beach OCC=Occu

BH=Boat House GP=Government Pt. AL=Alegria AH=Arroyo Hondo COP=Coal Oil Pt. CAR=Carpinteria MS=Mussel Shoals OS=Old Stairs 

WP=White’s Pt. PF=Pt. Fermin CC=Crystal Cove SC=Shaw’s Cove TI=Treasure Island DP=Dana Pt.
RESULTS 

Below are brief summaries of the major trends in target species abundances to 

accompany the figures that follow. Species that are shown in the figures were chosen

because they were most abundant overall, however, additional species important to 

particular sites are also mentioned in the results. Values for means and standard errors are 

given in Appendix C. 

Silvetia compressa ssp. compressa (Rockweed, formally called Pelvetia compressa and 

P. fastigiata)

The distribution of the rockweed Silvetia compressa is interesting because there is a large 

gap between Pt. Conception (near Government Pt.), and Los Angeles where the alga is 

either absent or too rare to target with photoplots (Figures 2a-c). The reasons for this gap 

in the distribution are unclear. Gradual declines in Silvetia cover were observed at all of 

the San Luis Obispo (SLO) sites over time, while cover at most other sites remained 

relatively stable and high (around 80%). One exception was Stairs, where the 1997/98 El 

Niño storms reduced cover of the alga and it has not yet recovered. 

Many sites showed a seasonal pattern in Silvetia cover, with lower values in spring vs. 

fall samples. This lower cover in spring may be due to a combination of factors including 

seasonal growth cycles, physical removal by winter storms and desiccation from extreme 

low tides that occur in the middle of the day in the spring. In southern California, Silvetia

grows rapidly in the summer resulting in peak cover in late summer/fall. It then 

reproduces November-February and receptacles are shed or deteriorate leaving lower 

cover in spring. This pattern may also hold for central California sites, although to our 

knowledge, no growth measurements have been done in this region. In the fall, Silvetia

was often described in the field notes as “dense in cover and very healthy”, whereas 

spring observations indicate that the alga was “dried out, ragged, and sparse.”

Silvetia is an important species to monitor not only because it, like other rockweeds, may 

be particularly sensitive to oiling (see appendix B), but also because it is adversely 

affected by human trampling (Murray and Denis 1997). This may be a factor influencing 

Silvetia abundance at urban southern California sites, where the alga appears to have 

undergone a significant decline since the 1950s (Murray, unpublished data). 

Hesperophycus californicus (Rockweed) 

Decline of the upper-shore rockweed, Hesperophycus harveyanus, was striking at the two 

SLO sites where it is monitored (Figure 3). Although cover of Endocladia muricata and 

barnacles increased slightly over time, Hesperophycus plots consisted mostly of open 

space (bare rock) after the rockweed declined. Although some reduction in 

Hesperophycus abundance occurred site-wide at both SLO sites, declines were more 

severe within permanent plots. Thus the plots did not necessarily reflect the sites overall. 

26
Endocladia muricata (Turfweed) 

The turfweed, Endocladia muricata, has a similar distribution to Silvetia compressa,

with a large gap between areas where it is abundant from Pt. Conception and Pt. Mugu 

(near Old Stairs, Figures 4a-c). Endocladia cover varied seasonally at many sites where it 

is monitored, with up to 40% greater cover in spring than in fall. At some sites where 

Endocladia cover declined, a subsequent increase in barnacle cover occurred (e.g. 

Hazards, Shell Beach, Old Stairs, and Shaw’s Cove). Endocladia facilitates recruitment 

of Silvetia, and three photoplots have been nearly taken over by this rockweed at Boat 

House, with Endocladia showing a strong reduction in a fourth plot. This decline in cover 

of Endocladia in permanent plots at Boat House does not reflect the abundance at the site 

overall, where the turf weed is still qualitatively described as “common to abundant”. At 

Occulto, Mytilus has steadily recruited into the Endocladia plots and is now a dominant 

part of what used to be a turf weed dominated community. Presence of Mytilus does not 

necessarily exclude Endocladia, as the alga can grow on top of mussels. Indeed, 

Endocladia abundance at the site overall is described as “common to abundant”. 

Mastocarpus papillatus

The red alga, Mastocarpus papillatus, is sampled at just two SLO sites, and is generally 

mixed with Endocladia, rather than occurring in pure stands (Figure 5). These two 

species appeared to have an inverse relationship in cover at both Pt. Sierra Nevada and 

Shell Beach. Mastocarpus experienced a general decline in cover in permanent plots over 

time at both sites, although in site-wide field notes the alga was qualitatively described at 

“common” at both sites. 

Mazzaella spp. (formally called Iridaea spp.) 

Mean cover of Mazzaella remained relatively constant over time at both SLO sites where 

it is sampled, although periodic fluctuations occured (Figure 6). Declines in Mazzaella

cover often were associated with increases in cover of articulated corallines, suggesting 

that coralline algae persisted as a stable understory below Mazzaella, and reduced canopy 

cover of Mazzaella simply exposed more corallines. The dominant species in the “other 

red algae” category quantified along with Mazzaella spp. included Chondracanthus 

canaliculatus (formerly Gigartina canaliculata), and Callithamnion pikeanum. These and 

the other red algae included in this category appeared to show a slight seasonal trend, 

with higher cover in fall than in spring. 

Anthopleura elegantissima/sola (Green Anemone, formally lumped as Anthopleura

elegantissima) 

Anthopleura spp. plots were quite variable in anemone cover, although cover at 

individual sites was fairly constant over time (Figures 7a-b). The primary reason for the 

variation in anemone cover was that plots at some sites (Boat House, Coal Oil Pt., and 

Carpinteria) contained the medium-sized solitary Anthopleura sola, while remaining sites 

(Alegria, Mussel Shoals, and Old Stairs) contained the small, densely aggregated, clonal 

27
Anthopleura elegantissima. Red algae, mainly consisting of small filamentous turfforming species that trap sand to form a dense “sand-turf” mixture, were abundant in 

anemone plots at Boat House, Coal Oil Pt., and Carpinteria. At Coal Oil Pt. and 

Carpinteria, red algal cover varied inversely with sand cover. Anemone plots at these 

sites are located on large, flat reefs where sand levels are constantly changing and red 

algae were likely buried and unburied by sand. 

Chthamalus fissus/dalli and Balanus glandula (White Barnacles) 

Barnacle cover varied substantially over time and among sites, which is not surprising 

since the monitored species of barnacles are relatively short-lived (Figures 8a-e). At three 

of the eight sites where individual barnacle species were distinguished (Hazards, Occulto, 

and Government Pt.), a mixture of Chthamalus and Balanus was present. The remaining 

five sites (Pt. Sierra Nevada, Cayucos, Shell Beach, Stairs, and Boat House) contained 

nearly pure stands of Chthamalus. 

At many sites, barnacle cover experienced periodic declines followed by recovery 

resulting from recruitment events. Some exceptions to this include five sites where 

barnacle abundance continued to decline after possible ENSO-induced declines (Pt. 

Sierra Nevada, Cayucos, and Stairs), or values stabilized at levels lower than pre-El Niño 

values (Hazards and Government Pt.). At Stairs, barnacles have nearly disappeared 

within the plots, and are uncommon at the sites overall, presumably due to mortality of 

adults and a near-complete lack of recruitment to the area. At Boat House the turfweed, 

Endocladia, was abundant in the barnacle plots for several years, but eventually the 

barnacles recovered and once again became the dominant space-occupiers. Barnacles 

within plots at Occulto have declined with little outlook for recovery, due to 

encroachment by mussels and several algal species, including Endocladia, Silvetia and

Cladophora, in what once were barnacle-dominated plots. Barnacles recruiting into this

site tended to settle above the barnacle plots, in an area that is at a higher tidal elevation 

than barnacles were commonly found when monitoring began at Occulto thirteen years 

ago. Thus, although barnacles in the plots have declined substantially, they are still 

common at the site overall. Barnacle cover at Arroyo Hondo declined substantially 

during the 1997/98 ENSO period, presumably due to impact from large swells and 

scouring, and then fell to near-zero in Fall 1998. Ephemeral algae were common in the 

plots in the seasons following this decline, but barnacles eventually recruited back in 

pulses, with all plots recovering by Fall 2000. 

One interesting note is that we have found “bent” Chthamalus at several sites, a condition 

previously only seen in the Gulf of California species Chthamalus anisopoma. We have 

been collecting bent individuals in order to identify them to species (requires examination 

of their calcareous plates) and thus far have found all collected individuals to be the 

southern species C. fissus.

Pollicipes polymerus (Goose Barnacle)

Goose barnacles were sampled at Government Pt., Alegria, and Carpinteria (Figure 9).

Cover was fairly constant over time at Government Pt. and Alegria. At Carpinteria 

28
however, Pollicipes showed a seasonal trend with lower cover in spring than in fall; 

Pollicipes cover also showed a decreasing trend over time at this site. At all sites, there 

was a slight increase in mussel cover within the Pollicipes plots over time. This trend was 

most pronounced at Carpinteria. 

Mytilus californianus (California Mussel)

Mussels were sampled through Fall 2003 at every site except Coal Oil Pt. (Figures 10ae). Mussel plots recently have been established at Coal Oil Pt. but had only been sampled 

once (Fall 2003) at the time of this report, and thus were not included. Mussel cover was 

relatively stable over time at seven of the nineteen sites (Pt. Sierra Nevada, Hazards,

Occulto, Government Pt., Mussel Shoals, Paradise Cove, and White’s Pt.). Four sites 

showed a decreasing trend in mussel cover over time. Mussels declined severely at Shell 

Beach, Stairs, and Pt. Fermin, and to a lesser degree at Alegria. The decline in mussel 

abundance was less severe, but still present at the site-wide scale at all sites except 

Alegria. At both Shell Beach and Alegria mussels appear to be recovering, while at Stairs 

cover has steadily declined since the initial sharp decline that occurred during the 

1997/98 El Niño storms. Mussels at Pt. Fermin also have declined substantially since 

being initially sampled in Fall 1999. Mussels at this site are relatively small, patchy in 

distribution, and tend to be restricted to fissure lines along horizontal rock surfaces. The 

owl limpet, Lottia gigantea, increased in abundance over time in the open spaces between 

mussel patches in the plots, and may have contributed to the continued decline of mussel 

cover by “bulldozing” newly recruited mussels off the rock. 

A decline in mussel cover was followed by recovery at nine sites. At seven sites, 

(Cayucos, Boat House, Arroyo Hondo, Crystal Cove, Shaw’s Cove, Treasure Island, and 

Dana Point), recovery was at or near initial levels by Fall 2003. Recovery was not 

complete at Old Stairs, and cover fluctuated substantially over time at Carpinteria. While 

the dynamics in mussel cover did not appear to be correlated among neighboring sites in 

most regions, mussels in plots at Orange County sites showed similar patterns of 

fluctuation (Figure 10e). Mussel cover declined at all four sites in Spring 1998, probably 

as a result of El Niño storms and perhaps stress due to warmer water temperatures. 

Mussels at these sites experienced varying degrees of recovery over the next five-year 

period, and then all sites exhibited a pulse of increased cover, likely due to a recruitment 

event, and possibly increased growth rates of established mussels, in Fall 2003. 

At sites where Pollicipes cover generally exceeded 20% (Cayucos, Hazards, Shell Beach, 

Stairs, Boat House, Arroyo Hondo, Old Stairs, Pt. Fermin, Shaw’s Cove, Treasure Island, 

Dana Pt.) decreases in mussel cover were associated with increases in Pollicipes cover. 

These two species have similar habitat requirements, and likely compete for space in 

some areas. 

Recovery Plots 

Recovery plots were established at Stairs in Spring 1998 after powerful El Niño storms

removed upper layers of the sedimentary reef in some areas, exposing the underlying 

29
clean, bare rock. To monitor natural recovery of these bare areas, we established seven 50 

x 75 cm photoplots in which we could monitor recruitment and persistence of species. 

Data from these plots were graphed differently than other plot types in order to show the 

diversity of species that have settled into these previously bare areas over the six-year 

period that they have been monitored (Figure 11). The first species to settle in the 

recovery plots were Ulva/Enteromorpha in Spring 1998. Ulva and Enteromorpha are 

short-lived “weedy” species, which are good colonizers of freshly disturbed substrata. 

Bare (unoccupied) rock declined in cover over time, but was still abundant in Fall 2003, 

and thus Ulva/Enteromorpha continued to move into and out of recovery plots 

throughout the six-year period. Non-coralline crusts settled into the plots in Fall 1998, 

along with Cladophora spp. Non-coralline crusts consist mostly of Ralfsia spp., with 

some Petrospongium rugosum and a small amount of the crustose form of Mastocarpus 

(formerly called “Petrocelis”). While the amount of non-coralline crusts increased 

slightly over time, Cladophora spp. became quite rare. Endocladia and other red algae, 

which included species such as Cryptosiphonia woodii, Cumagloia andersonii, 

Ceramium spp., and Polysiphonia spp., were present in small amounts beginning in Fall 

2000. Barnacles recruited into the plots in Fall 1999 and grew larger during successive 

years, resulting in an increase in percent cover. No species has yet dominated these plots, 

and a large amount of bare rock is still present, so it is likely that full their return to a 

stable state will take several more years. 

Phyllospadix scouleri/torreyi (Surfgrass) 

Phyllospadix showed a strong seasonal pattern, with higher cover in the fall than in the 

spring for all sites except Cayucos (Figures 12a-d). This seasonal pattern was likely due 

in part to the effects of winter storms, which abrade and shorten surfgrass blades, and 

thus reduce cover. In the southern sites, where sand is present in higher abundance as 

compared to northern sites, lower surfgrass cover in the spring was often associated with 

higher sand cover. Thus, the seasonal pattern in surfgrass cover may also be a result of 

sand movement, leading to partial burial of surfgrass. The lack of a seasonal pattern in 

surfgrass cover at Cayucos is likely due to the unique location of the surfgrass transects at 

this site. Unlike other sites, where surfgrass transects were established in areas that drain 

during low tide, transects at Cayucos are located in large pools, which reduces the 

amount of stress experienced by the plants due to air exposure, and perhaps also abrasion. 

Although many sites experienced extreme seasonal fluctuations in surfgrass cover, only 

two sites, Stairs and Carpinteria, showed substantial lasting reductions. Transects at Stairs 

were decimated by the 1997/98 El Niño storms, which ripped out whole sections of reef, 

as well as a large numbers of surfgrass plants. The decline at Carpinteria was associated 

with a sharp increase in Egregia cover. Surfgrass cover has been slowly recovering at 

both sites, and in Fall 2003 was at about 50% of 1992 cover values. 

Pisaster ochraceus & Asterina miniata (Seastars) 

Species of seastars counted in plots included Pisaster ochraceus, Asterina miniata, 

Pisaster giganteus, and Pycnopodia helianthoides. Pisaster ochraceus was the only 

30
species found at every site where seastars were sampled. At Boat House, Asterina 

miniata was also consistently present. Pisaster ochraceus numbers have fluctuated at 

most sites over time, but the color ratio of “other” (colors other than orange) to orange 

has remained fairly consistent at 3:1 for most sites (Figures 13a-e). There appears to have 

been slight increases in P. ochraceus populations at Pt. Sierra Nevada and Cayucos 

(Figure 13a), and also at Old Stairs (Figure 13d). Numbers of Asterina in the plots at Boat

House have declined slightly over time (Figure 13b). 

The fluctuations in seastar numbers over time likely reflect movement into and out of our 

permanent plots in response to factors such as water temperature, tidal height and food 

availability, and may not indicate true increases or decreases in site populations. 

Although we do not have much data for P. ochraceus populations at Orange County sites 

prior to the 1997/98 ENSO event, seastar numbers were low during this period and then 

spiked when water temperatures cooled in 1999. It is thought that P. ochraceus may 

move into deeper, subtidal areas during periods of elevated water temperatures, which

would be reflected in lower counts for our intertidal plots. At Occulto, seastar counts may 

be influenced by sampling conditions because much of the plot consists of steep reef 

edges that can be difficult to sample when the swell is large. At Arroyo Hondo and Old 

Stairs, seastars were sampled within narrow 2m x 5m swaths, which did not appear to 

accurately represent the site-wide abundance dynamics. Larger, irregular plots were set 

up at these sites in Spring 2004 in an attempt to address this issue. Despite potentially 

being too small, the original “swath” plots did capture major fluctuations in P. ochraceus

numbers at the sites overall. Qualitative observations note population expansions of P. 

ochraceus in 2001, which are reflected in the trend lines for plots at these sites. 

At most sites, orange P. ochraceus are slightly larger on average than individuals of other 

colors (Figures 14a-b). Indeed, we found very few small orange individuals, and we 

hypothesize that some purple or brown individuals may change to orange as they grow

larger. Figures 15a-c display size structure for the two color groups. From these figures it 

is evident that at most sites, the highest proportion of small individuals fall in the “other” 

category. Exceptions are at Shell Beach, Carpinteria, Mussel Shoals, and Old Stairs. 

Seastar numbers are low at all of these sites, and thus color ratio and size structure 

patterns would be expected to be more variable than at sites with large numbers of 

seastars because just a few individuals can skew the pattern. Sites with consistently high 

proportions of juvenile P. ochraceus. included Hazards, Stairs, and Boat House (Figures 

15a-b). 

Lottia gigantea (Owl Limpet) 

Numbers of the owl limpet, Lottia gigantea, were relatively stable at most sites over time

(Figures 16a-c). Three exceptions were Cayucos, where L. gigantea numbers declined 

slightly over time, and Government Pt. and Alegria, where numbers increased over time. 

At Crystal Cove, numbers of L. gigantea declined between Fall 1999 and Fall 2000 so a 

sixth plot was added in Spring 2001, resulting in an increase in numbers at this site. At 

Cayucos L. gigantea gradually declined from Spring 1996-Spring 1998. A recruitment 

event in Fall 1998, indicated by a decrease in the mean shell length (Figure 17a) and a 

positively skewed population distribution (Figure 18a) resulted in an increase in limpet 

31
numbers at Cayucos; however, L. gigantea numbers in the plots have slowly declined 

since Fall 1998. Some of this decline may be a result of habitat loss within the plots due 

to encroachment by surrounding mussels. At Government Pt., an increase in mussel cover 

within the L. gigantea plots appears to have had a different effect, with increased 

numbers of small owl limpets occurring on and amongst the mussels. This trend is 

reflected by an increase in the mean number of L. gigantea (Figure 16b), and a positively 

skewed population distribution over the past 5 1⁄2 years. Numbers of L. gigantea at 

Alegria also have increased over time, but at this site the increase appears to be due to a 

combination of recruitment and an influx of moderate-sized individuals from areas 

outside the plots because population size structure plots are not positively skewed. At Old 

Stairs and Paradise Cove there was a large recruitment event in Spring 2000 that resulted 

in substantially higher numbers of L. gigantea than previously recorded. Gradual declines 

in recent years at these sites and also at Mussel Shoals can be attributed to habitat loss

due to encroachment by mussels and Pollicipes. The reverse trend occurred at Shaw’s 

Cove between Fall 1996-Spring 1998, where a decline in mussel cover within L. gigantea

plots opened up more bare rock habitat for the limpets. A decrease in the average L. 

gigantea size was also noted during this period, suggesting that smaller limpets either 

recruited to the plots or migrated in from surrounding mussel shells. Then, in Fall 1998 

mussel cover began to increase, and L. gigantea numbers gradually declined. 

Large L. gigantea were consistently common at Stairs, Boat House, and Alegria (Figure 

18a-b). All of these sites have restricted human access, a condition that correlates with 

the presence of large L. gigantea likely because the limpets are harvested for food 

(Sagarin et al. in prep). However, these are not the only sites that are protected from the 

public, so other factors must influence L. gigantea size structure. Rancho Marino, for 

example, has extremely limited public access, but owl limpets are generally quite small 

(Figure 18a). In addition, sites that have high levels of human use (e.g. Crystal Cove and 

Shaw’s Cove) have very different population size structures (Figure 18c). Thus, it seems 

likely that a combination of factors, including human use, recruitment levels, and habitat 

structure are important parameters for determining population size structure in L. 

gigantea. 

Haliotis cracherodii (Black Abalone) 

At the sites presented in this report, black abalone typically occurred within rocky 

crevices. These crevices are more susceptible to wave damage than the surrounding flat 

reef areas and thus the number or location of plots in which black abalone were counted 

changed at several sites due to destruction of plots during large wave events. At Piedras 

Blancas, most of plot 1 broke away in Spring 2000, and a fourth plot was added as a 

replacement. We continued to count abalone in the undamaged sections of plot 1, but the 

newly added plot enabled us to get a more accurate representation of the abalone 

population at Piedras Blancas. Similarly, a fourth plot was added at Stairs in Spring 1998, 

after El Niño storms completely destroyed plot 1 and most of plot 3. At Pt. Sierra 

Nevada, we noted substantial black abalone numbers in two of our seastar plots, and 

began counting these as two additional abalone plots in Spring 1997. The addition of 

these plots at Pt. Sierra Nevada explains the jump in the number of abalone counted in 

Spring 1997 (Figure 19a). 

32
The fatal condition termed “withering syndrome” has caused drastic declines in black 

abalone populations as far north as Cayucos (Figures 19a-b). Evidence of withering 

syndrome was seen at Rancho Marino, Piedras Blancas, and Pt. Sierra Nevada although 

numbers have not yet declined substantially. The northward movement of withering 

syndrome has prompted PISCO (Partnership for Interdisciplinary Studies of Coastal 

Oceans) researchers at UC Santa Cruz to set up additional black abalone sites (in 

collaboration with the Monterey Bay Sanctuary) north of Pt. Sierra Nevada in Monterey, 

Santa Cruz, and San Mateo Counties. 

Recovery of black abalone populations to pre-withering syndrome levels is unlikely in 

the near term because recruitment is thought to be localized and the remaining 

individuals at these sites are probably too sparsely distributed to allow for successful 

spawning. This idea is substantiated by the lack of juvenile black abalone at all sites that 

have experienced WS-induced declines (Figures 19a-b, Figures 21a-b). Black abalone 

recruitment and survival may be further limited by habitat changes that occur after the 

animals disappear from an area, which we have observed in a separate study (see 

discussion for details). 

Motile Invertebrates 

A new addition to our monitoring program allowed us to capture abundance and size 

estimates for some of the common small motile invertebrates at our sites. Motile 

invertebrates were counted, and a subset of species measured, within select photoplot 

types. Abundances for the two or three most abundant motile species present in mussel, 

barnacle, Endocladia, and Silvetia plots are shown in Figures 22-25. Counts also were 

done in Hesperophycus, Mastocarpus, and Balanus plots, but only data for the four most 

common plot types are presented in this report. Motile invertebrate counts are currently 

done at all sites included in this report, but counts did not begin at Orange County sites 

until Fall 2003 and thus are not presented. 

Nearly all Silvetia plots contained high numbers of Tegula funebralis and limpets (Figure 

22). Lepidochitona hartwegii was consistently present in low levels at four sites, and was 

abundant at Pt. Fermin (Figure 22). This chiton is frequently associated with Silvetia, 

which it uses for protection from desiccation. 

Limpets and littorines were common in Endocladia plots at most sites (Figure 23), 

although littorines were most abundant in barnacle plots (see below). Interestingly, peaks 

in limpet or littorine abundance did not appear to be correlated with changes in percent 

cover of sessile species (Figures 4a-c), although we may not have a long enough time 

series for motile species to determine such temporal trends. Tegula funebralis was 

common at three sites, Hazards, Shell Beach, and Boat House. 

Limpets and littorines were the only motile organisms consistently abundant in barnacle 

plots (Figure 24). Littorines were typically present in very high numbers, often upwards 

of 1000 individuals per plot, while limpets tended to be an order of magnitude less 

common. Only one site, Stairs, had fewer littorines than limpets in the barnacle plots. 

Tegula funebralis was the most common motile species in mussel plots at seven of the 

seventeen sites (Pt. Sierra Nevada, Hazards, Shell Beach, Stairs, Alegria, Arroyo Hondo, 

33
and Paradise Cove, Figure 25). The mean number of T. funebralis per plot varied greatly 

from site to site, with highest turban snail densities occurring at Shell Beach and Alegria, 

and zero or near zero values at Cayucos, Occulto, Old Stairs, White’s Pt., and Pt. Fermin. 

Nucella spp. (mostly N. emarginata/ostrina) also was common in mussel plots, with 

highest mean abundance at Mussel Shoals and Old Stairs (Figure 25). Nucella spp. is a 

predator on Mytilus spp., and lives amongst the often tightly packed mussels, so counts 

for these whelks may be underestimates of the true population sizes. The third species 

commonly found in mussel plots was Nuttallina spp. These chitons were present at most 

sites, but were abundant only at Pt. Fermin (Figure 25). 

Large Nucella (>25mm) were consistently common in mussel plots at Pt. Sierra Nevada, 

Occulto, Government Pt., Alegria, and Arroyo Hondo (Figures 26a-b). The highest 

proportions of small size classes (<10mm) were present at Pt. Sierra Nevada and Alegria. 

Tegula sizes are presented for both mussel and Silvetia plots (Figures 27a-c). In general, 

size distributions were similar between the two plot types at sites where both were 

monitored, although small individuals (<10mm) tended to be more abundant in mussel 

plots than in Silvetia plots. This trend was especially apparent at Pt. Sierra Nevada, where 

mussel plots are located directly below Silvetia plots on the reef. A high proportion of 

small Tegula were present in mussel plots at Pt. Sierra Nevada, Shell Beach, Alegria, 

Arroyo Hondo, Coal Oil Pt., Carpinteria, Mussel Shoals, and Paradise Cove. Only four 

sites had high proportions of small individuals in Silvetia plots: Hazards, Shell Beach,

Stairs, and Boat House. High proportions of large Tegula (>20mm) were present in 

mussel plots at Stairs, Alegria, Arroyo Hondo, and Mussel Shoals. Silvetia plots 

contained high proportions of large snails at Stairs, Government Pt., and Pt. Fermin. 

DISCUSSION 

The MARINe consortium, encompassing sites in San Diego, Northern California, 

Oregon, and the Channel Islands, as well as those presented in this report, is the longestrunning, large scale rocky intertidal monitoring program on the west coast of North 

America. The data resulting from this collaborative monitoring program allow us to 

determine temporal and spatial community dynamics over both short and long temporal 

scales. This information is essential for assessing impact due to natural (e.g. ENSO event) 

or human induced (e.g. oil spill) disturbance (see Raimondi et al. 1999 for examples of 

both) as well as understanding biological community response to important physical 

phenomena, such as inter-decadal changes in sea surface temperature (McGowan et al.

1998). 

Assessing Long-term Impacts of the 1997/98 ENSO Event 

The 1997/98 ENSO event was one of the most powerful on record. The combination of 

warmer water temperatures and powerful storms associated with this oceanographic 

phenomenon resulted in significant declines in many species targeted by the MARINe 

monitoring program. These effects were presented in detail in our previous report 

(Raimondi et al. 1999) for the eleven northernmost MARINe sites (for which we had 

sufficient data prior to the ENSO event), but we revisit them here to examine whether the 

ENSO event may have had a lasting impact on the abundance of some target species. In 

34
the 1999 report we determined that five of the six most ubiquitous target species 

(barnacles, mussels, Endocladia, Silvetia, and Phyllospadix) appeared to be negatively 

impacted by the ENSO event. While declines in Endocladia did not persist for the long 

term, the ENSO event may have had a lasting impact on the four other species or species 

complexes in some areas. 

Perhaps the most widespread lasting effects were to barnacle and mussel communities. At 

five sites, barnacle declines that began in 1997/98 either continued through Fall 2003 (Pt. 

Sierra Nevada, Cayucos, and Stairs), or values stabilized at levels lower than pre-El Niño 

values (Hazards and Government Pt.), suggesting a long-term effect of the ENSO event 

on barnacle populations in these areas. For mussel communities, lasting negative impacts 

from the ENSO event may have occurred at Shell Beach and Stairs. In addition, although 

Orange County sites were not included in the original analysis of ENSO effects 

(Raimondi et al. 1999), mussel communities at three of the sites in this region 

experienced depressions in abundance following the ENSO event that persisted for 

several years. 

Another species that suffered extensive and lasting impacts from the ENSO event is the 

black abalone, Haliotis cracherodii. Declines in the population sizes of this species due 

to withering syndrome (see below) appear to be accelerated by the warm water associated 

with ENSO conditions. During warm water periods the disease moves up the coast more 

rapidly, and the rate of population decline increases once die-offs begin at a site 

(Raimondi et al. 2002). 

Species for which long-term effects of the ENSO event are not so clear include 

Phyllospadix and Silvetia. At many sites, abundances of both of these species were in 

decline prior to the 1997/98 ENSO event, although declines may have intensified during 

this period (e.g. surfgrass at Carpinteria, Mussel Shoals, and Paradise Cove). One 

exception is Stairs, where abundances of both species were severely depressed, and had 

not recovered by Fall 2003. 

In addition to documenting impacts to target species in established plots, the El Niño 

induced storm damage at Stairs created natural clearings on the reef by removing large 

sections of rock, and provided us with a unique opportunity to document community 

succession in the mid-intertidal. Nearly all succession studies are done in artificially 

cleared patches, which are small relative to the reef size and often recover via 

encroachment by surrounding species. Newly exposed sections of reef at Stairs were 

large (several square meters in size), and were likely to “recover” via colonization by 

propagules. Thus, the areas had the potential to develop into communities quite different 

from surrounding, undisturbed areas. We have been monitoring photoplots in these 

cleared areas for six years, and they continue to be dominated by bare rock and weedy, 

opportunistic species; thus, it will likely be several more years before the plots reach a 

more stable equilibrium state, where longer-lived mid-zone species such as Silvetia and 

mussels, dominate plot cover. 

The MARINe monitoring project now encompasses enough data, temporally and 

spatially, that we can begin to understand how major phenomena, such as the 1997/98 

ENSO event, impact rocky intertidal communities in both the short and long term. It is

clear that the lasting effects of a disturbance event can be site specific and depend on the 

35
severity of the initial disturbance as well as local physical conditions that influence 

recruitment, growth and survivorship of organisms. 

Documenting Disease and Invasion 

While arguably one of the greatest benefits of long term monitoring is documenting 

community response to known natural or human induced phenomena, such as the 

1997/98 ENSO event, additional value comes from having researchers consistently 

present in the field so that we may opportunistically capture events that would otherwise 

be missed. The most important example of this for the MARINe intertidal monitoring is 

the ongoing decline of black abalone populations due to the disease called withering 

syndrome. Withering syndrome (WS) is caused by the bacterium Candidatus

Xenohaliotis californiensis, which attacks the lining of the digestive track and results in 

reduced body mass, weakness, and eventual withering of the abalone’s foot until it can no 

longer cling to the substratum (Friedman et al. 2000). It was first discovered on the 

Channel Islands, but was not expected to progress north along the mainland due to 

prevailing southward currents and cooler temperatures north of Pt. Conception (Lafferty 

and Kuris 1993, Richards and Davis 1993). In 1993, one year after the first long-term

monitoring sites were established on the mainland, we documented a severe decline of 

black abalone at Government Pt., near Pt. Conception, and have seen subsequent declines 

in populations as far north as Piedras Blancas (Altstatt et al 1996, Raimondi et al. 2002). 

Currently, we are the only group tracking the northward progression of this devastating 

disease on a large scale. Recovery is hindered by a near lack of recruitment into areas 

affected by WS, and perhaps also by habitat changes occurring to the crevice 

communities where black abalone were once abundant (Miner et al. in prep.) Declines 

have been so severe across all regions in southern California that the species is now a 

candidate for protection under the USA Endangered Species Act. 

An additional benefit of long-term monitoring work is that researchers become familiar 

with the composition and abundance of species at a site, and are thus more likely to 

notice if a new species invades, or if particular species increase or decrease in abundance. 

General site notes are an integral part of our monitoring work, and researchers working at 

sites from San Diego to northern Oregon have an identical list of species for which 

relative abundance (including absence) and condition (e.g. reproductive, bleached) is 

recorded. This standard species list allows us to consistently make general observations 

about species that are not targeted in our plots. One species included on this list is the turf 

alga, Caulacanthus ustulatus. Caulacanthus ustulatus is believed to be a non-native 

invasive alga that was first documented in southern California in 1999 (Murray personal 

observation). The alga also occurs in Washington and British Columbia where it was first 

recorded in the 1960’s and 1980’s respectively. In these northern regions there is some

debate over whether it is truly non-native or simply a cryptic native that was previously 

overlooked (Gabrielson pers. com.). Genetic analysis aligns southern California 

specimens with an Asian population introduced to France but the particular markers 

utilized cannot definitively determine whether the southern California material also 

represents an introduction. Alternatively, the presence of C. ustulatus in southern 

California might represent a recent and substantial range extension along the coast, 

because thorough floristic studies by Hollenberg and Dawson in the 1950’s and 1960’s 

36
and extensive floristic work by Littler and colleagues in the 1970’s did not record the 

presence of C. ustulatus. 

The CSUF group has documented a significant increase in C. ustulatus cover at several 

southern California sites south of Santa Monica Bay over the last few years, including 

abundant growths at Dana Pt., Shaw’s Cove and Crystal Cove (Whiteside and Murray, 

unpublished data). This species now grows extensively in upper-mid shore habitat 

occupied previously by other red algal species such as Gelidium spp. and Endocladia 

muricata, where it now exceeds the cover of these other taxa at these study sites. 

Caulacanthus ustulatus also grows on mussel shells and Silvetia holdfasts and fronds 

suggesting a potential interaction with these key species. Observations such as this are 

important in light of recent evidence suggesting that species ranges may be shifting due 

to global warming (Barry et al. 1995, Sagarin et al. 1999) and also that introductions of 

non-native species into the marine environment are rapidly increasing (Carlton 1993). 

Additional Work Resulting from Monitoring Observations 

Observations resulting from the MARINe long-term monitoring surveys serve an 

important function as the basis for more in-depth surveys, or experimental research, in 

which the processes that cause community change are more closely examined. One study, 

done by Mark Readdie, a PhD student at UCSC, was inspired by the observation at 

several sites that over time, barnacle plots became inundated with Endocladia, and plots 

that initially contained high cover of Endocladia became partially or wholly covered by

Silvetia. Further investigation showed that the barnacle zone at these sites had shifted 

upward, into areas that were previously bare rock. It has long been assumed that the 

upper limits of species’ zones are set by physical factors such as temperature and 

emersion time, and are thus stable, while the lower limits are set by biological 

interactions (e.g. competition and predation). However, the upward shifts of species 

zones documented in this study suggest that facilitation, a biological factor, is important 

for establishing species’ upper limits. In this case, barnacles are facilitating the upward 

movement of Endocladia, and Endocladia is providing suitable habitat for Silvetia, above 

the zone where it previously occurred. 

Another study resulting from observations by MARINe researchers focused on changes 

that occurred in crevice communities after black abalone disappeared. Researchers 

working at sites on the Channel Islands and on the mainland noticed that after black 

abalone populations had been severely depressed by withering syndrome, the biological 

communities in the crevices in which they had lived began to change, with increasing 

cover of encrusting invertebrates. A detailed study comparing abalone crevice 

communities pre- and post-withering syndrome indicated that these areas were indeed 

changing from being dominated by bare rock and crustose algae, to having increased 

cover of encrusting invertebrates (Miner et al. in prep.) This change from “good” abalone 

habitat, to habitat less favorable for abalone recruitment and survival may hinder black 

abalone recovery. 

New Additions to Protocol 

Several new additions have been made to our sampling procedures since the last report 

(see Methods). Two of these, field scoring of barnacle plots, and counts of motile 

invertebrates in photoplots, were added in the hope that they would increase our 

37
understanding of observed patterns of community change in the photoplots. Field scoring 

of barnacles allows us to distinguish between Chthamalus spp. and Balanus glandula, 

which have very different life spans and recruitment dynamics. In addition, we can 

separate live barnacles from dead, and can potentially link cover of empty barnacle tests 

to the presence of other species that use these structures for protection from desiccation 

or predation (e.g. Endocladia, Silvetia, littorines). Counts of motile invertebrates may 

help to explain the presence or absence of certain sessile species in the photoplots, 

although these relationships have yet to be rigorously examined. 

Another new addition to our sampling protocol is recording size and color information 

for Pisaster ochraceus, rather than just counts of seastar numbers. Measuring P. 

ochraceus gives us an indication of population size structure at our sites and enables us to 

determine which sites receive the highest levels of recruitment, and contain the largest 

individuals. P. ochraceus color data has proved to be interesting and we have identified a 

consistent ratio of approximately 3 purple/brown stars for every orange star in larger size 

classes at most sites. This ratio changes in smaller size classes, where orange P. 

ochraceus appear to be less common. 

Conclusions 

Long term monitoring of MARINe sites allows for assessment of change within intertidal 

communities on many levels. We have documented naturally occurring seasonal 

variation, as well as longer-term fluctuations in the abundance of species. This record of 

species dynamics within sites has enabled us to assess damage and document recovery 

following human induced impacts (e.g. oil spill), and natural disturbances (e.g. El Niño 

storms). One of the most interesting findings of this study is that within-site species 

dynamics can be very different among neighboring sites, but overall community structure 

within regional areas tends to be quite similar. In other words, neighboring sites tend to 

be more similar to one another than more distant sites, even though individual species 

dynamics may be quite different. In the future, we plan to investigate the large-scale 

processes that likely drive regional grouping patterns and link them to observed intertidal 

community dynamics. 

38
rock

Silvetia compressa

Point Sierra Nevada

Figure 2a. Species abundances in Silvetia

plots. S=spring samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Shell Beach

Hazards

Cayucos

Percent Cov

e

r

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

rock

Silvetia compressa

Point Sierra Nevada

Figure 2a. Species abundances in Silvetia

plots. S=spring samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Shell Beach

Hazards

Cayucos

Percent Cov

e

r

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

39
rock

Silvetia compressa

Stairs

Figure 2b. Species abundances in Silvetia

plots. Boat House was not sampled in F95 

and Gov’t. Pt. was not sampled in F99.

S=spring samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Point Fermin

Government Point

Boat House

Percent Cov

e

r

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

rock

Silvetia compressa

Stairs

Figure 2b. Species abundances in Silvetia

plots. Boat House was not sampled in F95 

and Gov’t. Pt. was not sampled in F99.

S=spring samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Point Fermin

Government Point

Boat House

Percent Cov

e

r

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

40
rock

Silvetia compressa

Crystal Cove

Figure 2c. Species abundances in Silvetia

plots. S=spring samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Dana Point

Treasure Island

Shaw’s Cove

Percent Cov

e

r

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

rock

Silvetia compressa

Crystal Cove

Figure 2c. Species abundances in Silvetia

plots. S=spring samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Dana Point

Treasure Island

Shaw’s Cove

Percent Cov

e

r

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

41
Point Sierra Nevada

Figure 3. Species abundances in 

Hesperophycus plots. S=spring samples,

F=fall samples.

Cayucos

Percent Cov

e

r

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

rock

Chthamalus + Balanus

Endocladia muricata

Hesperophycus

0

20

40

60

80

100

0

20

40

60

80

100

Point Sierra Nevada

Figure 3. Species abundances in 

Hesperophycus plots. S=spring samples,

F=fall samples.

Cayucos

Percent Cov

e

r

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

rock

Chthamalus + Balanus

Endocladia muricata

Hesperophycus

rock

Chthamalus + Balanus

Endocladia muricata

Hesperophycus

rock

Chthamalus + Balanus

Endocladia muricata

Hesperophycus

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

42
Cayucos

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Occulto

Shell Beach

Hazards

Percent Cov

e

r

Figure 4a. Species abundances in 

Endocladia plots. Occulto plot 4 was not 

sampled F99. S=spring samples, F=fall 

samples. rock

Chthamalus & Balanus

Silvetia compressa

Endocladia muricata

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

Cayucos

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Occulto

Shell Beach

Hazards

Percent Cov

e

r

Figure 4a. Species abundances in 

Endocladia plots. Occulto plot 4 was not 

sampled F99. S=spring samples, F=fall 

samples. rock

Chthamalus & Balanus

Silvetia compressa

Endocladia muricata

rock

Chthamalus & Balanus

Silvetia compressa

Endocladia muricata

rock

Chthamalus & Balanus

Silvetia compressa

Endocladia muricata

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

43
Stairs

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Government Point

Boat House

Percent Cov

e

r

Figure 4b. Species abundances in 

Endocladia plots. Gov’t. Pt. was not sampled

in F99. S=spring samples, F=fall samples.

rock

Chthamalus & Balanus

Silvetia compressa

Endocladia muricata

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

Stairs

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Government Point

Boat House

Percent Cov

e

r

Figure 4b. Species abundances in 

Endocladia plots. Gov’t. Pt. was not sampled

in F99. S=spring samples, F=fall samples.

rock

Chthamalus & Balanus

Silvetia compressa

Endocladia muricata

rock

Chthamalus & Balanus

Silvetia compressa

Endocladia muricata

rock

Chthamalus & Balanus

Silvetia compressa

Endocladia muricata

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

44
Old Stairs

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Shaw’s Cove

White’s Point

Paradise Cove

Percent Cov

e

r

Figure 4c. Species abundances in 

Endocladia plots. S=spring samples, F=fall

samples.

rock

Chthamalus & Balanus

Silvetia compressa

Endocladia muricata

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

Old Stairs

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Shaw’s Cove

White’s Point

Paradise Cove

Percent Cov

e

r

Figure 4c. Species abundances in 

Endocladia plots. S=spring samples, F=fall

samples.

rock

Chthamalus & Balanus

Silvetia compressa

Endocladia muricata

rock

Chthamalus & Balanus

Silvetia compressa

Endocladia muricata

rock

Chthamalus & Balanus

Silvetia compressa

Endocladia muricata

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

45
Point Sierra Nevada

Figure 5. Species abundances in 

Mastocarpus plots. S=spring samples, F=fall

samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Shell Beach

Percent Cov

e

r

0

20

40

60

80

100

0

20

40

60

80

100

rock

Endocladia muricata

Mastocarpus papillatus

Point Sierra Nevada

Figure 5. Species abundances in 

Mastocarpus plots. S=spring samples, F=fall

samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Shell Beach

Percent Cov

e

r

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

rock

Endocladia muricata

Mastocarpus papillatus

rock

Endocladia muricata

Mastocarpus papillatus

46
Point Sierra Nevada

Hazards

Percent Cov

e

r

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Figure 6. Species abundances in Mazzaella

plots. S=spring samples, F=fall samples.

other red algae

articulated corallines

Mazzaella spp.

0

20

40

60

80

100

0

20

40

60

80

100

Point Sierra Nevada

Hazards

Percent Cov

e

r

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Figure 6. Species abundances in Mazzaella

plots. S=spring samples, F=fall samples.

other red algae

articulated corallines

Mazzaella spp.

other red algae

articulated corallines

Mazzaella spp.

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

47
Boat House

Figure 7a. Species abundances in 

Anthopleura plots. Carpinteria plot 2 was not

sampled F98. S=spring samples, F=fall 

samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Carpinteria

Coal Oil Point

Alegria

Percent Cov

e

r

rock

sand

red algae

Anthopleura spp.

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

Boat House

Figure 7a. Species abundances in 

Anthopleura plots. Carpinteria plot 2 was not

sampled F98. S=spring samples, F=fall 

samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Carpinteria

Coal Oil Point

Alegria

Percent Cov

e

r

rock

sand

red algae

Anthopleura spp.

rock

sand

red algae

Anthopleura spp.

rock

sand

red algae

Anthopleura spp.

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

48
Mussel Shoals

Old Stairs

Percent Cov

e

r

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Figure 7b. Species abundances in 

Anthopleura plots. Old Stairs plot 5 was not

sampled S97. S=spring samples, F=fall 

samples. rock

sand

red algae

Anthopleura spp.

0

20

40

60

80

100

0

20

40

60

80

100

Mussel Shoals

Old Stairs

Percent Cov

e

r

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Figure 7b. Species abundances in 

Anthopleura plots. Old Stairs plot 5 was not

sampled S97. S=spring samples, F=fall 

samples. rock

sand

red algae

Anthopleura spp.

rock

sand

red algae

Anthopleura spp.

rock

sand

red algae

Anthopleura spp.

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

49
0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

Point Sierra Nevada

Figure 8a. Species abundances in barnacle 

plots. Note that barnacle species were

separated starting in spring 2001. Barnacles

were 100% Chthamalus at sites where 

individual species are not visible. Hazards 

plot 5 was not sampled F95. S=spring 

samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Shell Beach

Hazards

Cayucos

Percent Cov

e

r

Balanus glandula

Chthamalus fissus/dalli

Endocladia muricata

Balanus + Chthamalus

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

Point Sierra Nevada

Figure 8a. Species abundances in barnacle 

plots. Note that barnacle species were

separated starting in spring 2001. Barnacles

were 100% Chthamalus at sites where 

individual species are not visible. Hazards 

plot 5 was not sampled F95. S=spring 

samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Shell Beach

Hazards

Cayucos

Percent Cov

e

r

Balanus glandula

Chthamalus fissus/dalli

Endocladia muricata

Balanus + Chthamalus

Balanus glandula

Chthamalus fissus/dalli

Endocladia muricata

Balanus + Chthamalus

Balanus glandula

Chthamalus fissus/dalli

Endocladia muricata

Balanus + Chthamalus

50
Occulto

Figure 8b. Species abundances in barnacle 

plots. Note that barnacle species were

separated starting in spring 2001. Barnacles

were 100% Chthamalus at sites where 

individual species are not visible. Occulto 

plot 4 was not sampled F98. Gov’t. Pt. was

not sampled F99. S=spring samples, F=fall

samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Government Point

Boat House

Stairs

Percent Cov

e

r

Balanus glandula

Chthamalus fissus/dalli

Endocladia muricata

Balanus + Chthamalus

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

Occulto

Figure 8b. Species abundances in barnacle 

plots. Note that barnacle species were

separated starting in spring 2001. Barnacles

were 100% Chthamalus at sites where 

individual species are not visible. Occulto 

plot 4 was not sampled F98. Gov’t. Pt. was

not sampled F99. S=spring samples, F=fall

samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Government Point

Boat House

Stairs

Percent Cov

e

r

Balanus glandula

Chthamalus fissus/dalli

Endocladia muricata

Balanus + Chthamalus

Balanus glandula

Chthamalus fissus/dalli

Endocladia muricata

Balanus + Chthamalus

Balanus glandula

Chthamalus fissus/dalli

Endocladia muricata

Balanus + Chthamalus

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

51
Alegria

Figure 8c. Species abundances in barnacle 

plots. Note that barnacle species were not

separated for these sites. Alegria plot 2 and 

Mussel Shoals plot 5 were not sampled F98.

Arroyo Hondo plot 1 was not sampled S00.

S=spring samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Mussel Shoals

Carpinteria

Arroyo Hondo

Percent Cov

e

r

Endocladia muricata

Balanus + Chthamalus

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

Alegria

Figure 8c. Species abundances in barnacle 

plots. Note that barnacle species were not

separated for these sites. Alegria plot 2 and 

Mussel Shoals plot 5 were not sampled F98.

Arroyo Hondo plot 1 was not sampled S00.

S=spring samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Mussel Shoals

Carpinteria

Arroyo Hondo

Percent Cov

e

r

Endocladia muricata

Balanus + Chthamalus

Endocladia muricata

Balanus + Chthamalus

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

52
Old Stairs

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Point Fermin

White’s Point

Paradise Cove

Percent Cov

e

r

Endocladia muricata

Figure 8d. Species abundances in barnacle Balanus + Chthamalus

plots. Note that barnacle species were not

separated for these sites. White’s Pt. plot 2 

was not sampled F96. S=spring samples, 

F=fall samples.

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

Old Stairs

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Point Fermin

White’s Point

Paradise Cove

Percent Cov

e

r

Endocladia muricata

Balanus + Chthamalus

Endocladia muricata

Figure 8d. Species abundances in barnacle Balanus + Chthamalus

plots. Note that barnacle species were not

separated for these sites. White’s Pt. plot 2 

was not sampled F96. S=spring samples, 

F=fall samples.

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

53
Crystal Cove

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Dana Point

Treasure Island

Shaw’s Cove

Percent Cov

e

r

Endocladia muricata

Figure 8e. Species abundances in barnacle Balanus + Chthamalus

plots. Note that barnacle species were not

separated for these sites. S=spring samples,

F=fall samples.

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

Crystal Cove

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Dana Point

Treasure Island

Shaw’s Cove

Percent Cov

e

r

Endocladia muricata

Balanus + Chthamalus

Endocladia muricata

Figure 8e. Species abundances in barnacle Balanus + Chthamalus

plots. Note that barnacle species were not

separated for these sites. S=spring samples,

F=fall samples.

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

54
Government Point

Figure 9. Species abundances in Pollicipes

plots. Gov’t. Pt. was not sampled in F99. 

S=spring samples, F=fall samples.

Carpinteria

Alegria

Percent Cov

e

r

rock

Mytilus californianus

Pollicipes polymerus

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

Government Point

Figure 9. Species abundances in Pollicipes

plots. Gov’t. Pt. was not sampled in F99. 

S=spring samples, F=fall samples.

Carpinteria

Alegria

Percent Cov

e

r

rock

Mytilus californianus

Pollicipes polymerus

rock

Mytilus californianus

Pollicipes polymerus

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

55
Point Sierra Nevada

Figure 10a. Species abundances in Mytilus

plots. S=spring samples, F=fall samples.

Shell Beach

Hazards

Cayucos

Percent Cov

e

r

rock

Pollicipes polymerus

Mytilus californianus

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

Point Sierra Nevada

Figure 10a. Species abundances in Mytilus

plots. S=spring samples, F=fall samples.

Shell Beach

Hazards

Cayucos

Percent Cov

e

r

rock

Pollicipes polymerus

Mytilus californianus

rock

Pollicipes polymerus

Mytilus californianus

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

56
Occulto

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Government Point

Boat House

Stairs

Percent Cov

e

r

Figure 10b. Species abundances in Mytilus

plots. Occulto plot 5 was not sampled F98.

Gov’t. Pt. was not sampled in F99. S=spring

samples, F=fall samples.

rock

Pollicipes polymerus

Mytilus californianus

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

Occulto

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Government Point

Boat House

Stairs

Percent Cov

e

r

Figure 10b. Species abundances in Mytilus

plots. Occulto plot 5 was not sampled F98.

Gov’t. Pt. was not sampled in F99. S=spring

samples, F=fall samples.

rock

Pollicipes polymerus

Mytilus californianus

rock

Pollicipes polymerus

Mytilus californianus

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

57
Alegria

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Mussel Shoals

Carpinteria

Arroyo Hondo

Percent Cov

e

r

Figure 10c. Species abundances in Mytilus

plots. The following plots were not sampled 

at Carpinteria: 3 F92, 1 S93, all F01, all F02.

S=spring samples, F=fall samples.

rock

Pollicipes polymerus

Mytilus californianus

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

Alegria

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Mussel Shoals

Carpinteria

Arroyo Hondo

Percent Cov

e

r

Figure 10c. Species abundances in Mytilus

plots. The following plots were not sampled 

at Carpinteria: 3 F92, 1 S93, all F01, all F02.

S=spring samples, F=fall samples.

rock

Pollicipes polymerus

Mytilus californianus

rock

Pollicipes polymerus

Mytilus californianus

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

58
Old Stairs

White’s Point

Paradise Cove

Figure 10d. Species abundances in Mytilus

plots. S=spring samples, F=fall samples.

rock

Pollicipes polymerus

Mytilus californianus

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Point Fermin

Percent Cov

e

r

Old Stairs

White’s Point

Paradise Cove

Figure 10d. Species abundances in Mytilus

plots. S=spring samples, F=fall samples.

rock

Pollicipes polymerus

Mytilus californianus

rock

Pollicipes polymerus

Mytilus californianus

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Point Fermin

Percent Cov

e

r

59
Crystal Cove

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Dana Point

Treasure Island

Shaw’s Cove

Percent Cov

e

r

Figure 10e. Species abundances in Mytilus

plots. Crystal Cove plots 1 & 2 were not

sampled S00. S=spring samples, F=fall 

samples.

rock

Pollicipes polymerus

Mytilus californianus

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100 Crystal Cove

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Dana Point

Treasure Island

Shaw’s Cove

Percent Cov

e

r

Figure 10e. Species abundances in Mytilus

plots. Crystal Cove plots 1 & 2 were not

sampled S00. S=spring samples, F=fall 

samples.

rock

Pollicipes polymerus

Mytilus californianus

rock

Pollicipes polymerus

Mytilus californianus

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

60
S98 F98 S99 F99 S00 F00 S01 F01 S02 F02 S03 F03

0

10

20

30

40

50

60

70

80

90

100

Percent Cov

er

Chthamalus/Balanus

Other Red Algae

Endocladia

Cladophora

Silvetia

Ulva/Enteromorpha

Non-coralline Crusts

Rock

Figure 11. Species abundances in Recovery plots at Stairs. S=spring 

samples, F=fall samples.

S98 F98 S99 F99 S00 F00 S01 F01 S02 F02 S03 F03

0

10

20

30

40

50

60

70

80

90

100

Percent Cov

er

S98 F98 S99 F99 S00 F00 S01 F01 S02 F02 S03 F03

0

10

20

30

40

50

60

70

80

90

100

Percent Cov

er

Chthamalus/Balanus

Other Red Algae

Endocladia

Cladophora

Silvetia

Ulva/Enteromorpha

Non-coralline Crusts

Rock

Chthamalus/Balanus

Other Red Algae

Endocladia

Cladophora

Silvetia

Ulva/Enteromorpha

Non-coralline Crusts

Rock

Figure 11. Species abundances in Recovery plots at Stairs. S=spring 

samples, F=fall samples.

61
Point Sierra Nevada

Figure 12a. Species abundances in 

Phyllospadix transects. Hazards was not 

sampled in S02, and only transect 3 was

sampled in F02 and S03. Shell Beach 

transect 3 was not sampled F01. S=spring

samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Shell Beach

Hazards

Cayucos

Percent Cov

e

r

sand

other algae

Egregia menziesii

Phyllospadix spp.

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

Point Sierra Nevada

Figure 12a. Species abundances in 

Phyllospadix transects. Hazards was not 

sampled in S02, and only transect 3 was

sampled in F02 and S03. Shell Beach 

transect 3 was not sampled F01. S=spring

samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Shell Beach

Hazards

Cayucos

Percent Cov

e

r

sand

other algae

Egregia menziesii

Phyllospadix spp.

sand

other algae

Egregia menziesii

Phyllospadix spp.

sand

other algae

Egregia menziesii

Phyllospadix spp.

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

62
Stairs

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Arroyo Hondo

Alegria

Government Point

Percent Cov

e

r

Figure 12b. Species abundances in 

Phyllospadix transects. Stairs transect 1 was

not sampled F98. Gov’t Pt. was not sampled 

F99. S=spring samples, F=fall samples. sand

other algae

Egregia menziesii

Phyllospadix spp.

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

Stairs

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Arroyo Hondo

Alegria

Government Point

Percent Cov

e

r

Figure 12b. Species abundances in 

Phyllospadix transects. Stairs transect 1 was

not sampled F98. Gov’t Pt. was not sampled 

F99. S=spring samples, F=fall samples. sand

other algae

Egregia menziesii

Phyllospadix spp.

sand

other algae

Egregia menziesii

Phyllospadix spp.

sand

other algae

Egregia menziesii

Phyllospadix spp.

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

63
Coal Oil Point

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Paradise Cove

Mussel Shoals

Carpinteria

Percent Cov

e

r

Figure 12c. Species abundances in 

Phyllospadix transects. S=spring samples,

F=fall samples.

sand

other algae

Egregia menziesii

Phyllospadix spp.

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

Coal Oil Point

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Paradise Cove

Mussel Shoals

Carpinteria

Percent Cov

e

r

Figure 12c. Species abundances in 

Phyllospadix transects. S=spring samples,

F=fall samples.

sand

other algae

Egregia menziesii

Phyllospadix spp.

sand

other algae

Egregia menziesii

Phyllospadix spp.

sand

other algae

Egregia menziesii

Phyllospadix spp.

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

64
Point Fermin

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Crystal Cove

Percent Cov

e

r

Figure 12d. Species abundances in 

Phyllospadix transects. S=spring samples,

F=fall samples.

sand

other algae

Egregia menziesii

Phyllospadix spp.

0

20

40

60

80

100

0

20

40

60

80

100

Point Fermin

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Crystal Cove

Percent Cov

e

r

Figure 12d. Species abundances in 

Phyllospadix transects. S=spring samples,

F=fall samples.

sand

other algae

Egregia menziesii

Phyllospadix spp.

sand

other algae

Egregia menziesii

Phyllospadix spp.

sand

other algae

Egregia menziesii

Phyllospadix spp.

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

65
Shell Beach

Hazards

Cayucos

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Point Sierra Nevada

Figure 13a. Seastar abundances and 

Pisaster ochraceus colors. Note differences 

in scale. S=spring samples, F=fall samples.

Mean Number Per Plot

other P. ochraceus

orange P. ochraceus

Total Pisaster ochraceus

0

10

20

30

0

2

4

6

8

10

0

10

20

30

0

20

40

60

Shell Beach

Hazards

Cayucos

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Point Sierra Nevada

Figure 13a. Seastar abundances and 

Pisaster ochraceus colors. Note differences 

in scale. S=spring samples, F=fall samples.

Mean Number Per Plot

other P. ochraceus

orange P. ochraceus

Total Pisaster ochraceus

0

10

20

30

0

2

4

6

8

10

0

2

4

6

8

10

0

10

20

30

0

20

40

60

66
Occulto

Figure 13b. Seastar abundances and 

Pisaster ochraceus colors. Note differences 

in scale. Gov’t. Pt. plot #1 was not sampled 

S92 or S01 and no plots were sampled S93 

or F99. S=spring samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Government Point

Boat House

Stairs

Asterina miniata

other P. ochraceus

orange P. ochraceus

Total Pisaster ochraceus

Mean Number Per Plot

0

50

100

150

200

250

0

10

20

30

40

50

0

10

20

30

40

50

0

20

40

60

80

100

120

Occulto

Figure 13b. Seastar abundances and 

Pisaster ochraceus colors. Note differences 

in scale. Gov’t. Pt. plot #1 was not sampled 

S92 or S01 and no plots were sampled S93 

or F99. S=spring samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Government Point

Boat House

Stairs

Asterina miniata

other P. ochraceus

orange P. ochraceus

Total Pisaster ochraceus

Asterina miniata

other P. ochraceus

orange P. ochraceus

Total Pisaster ochraceus

Mean Number Per Plot

0

50

100

150

200

250

0

50

100

150

200

250

0

10

20

30

40

50

0

10

20

30

40

50

0

10

20

30

40

50

0

10

20

30

40

50

0

20

40

60

80

100

120

0

20

40

60

80

100

120

67
Alegria

Figure 13c. Seastar abundances and 

Pisaster ochraceus colors. Note differences 

in scale. Carpinteria plot 3 was not sampled 

in F94 and no plots were sampled in S95, 

F95, F96, F98, F99, F01, F02 or F03.

S=spring samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Mussel Shoals

Carpinteria

Arroyo Hondo

other P. ochraceus

orange P. ochraceus

Total Pisaster ochraceus

Mean Number Per Plot

0

20

40

60

80

0

10

20

30

0

10

20

30

0

10

20

30

Alegria

Figure 13c. Seastar abundances and 

Pisaster ochraceus colors. Note differences 

in scale. Carpinteria plot 3 was not sampled 

in F94 and no plots were sampled in S95, 

F95, F96, F98, F99, F01, F02 or F03.

S=spring samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Mussel Shoals

Carpinteria

Arroyo Hondo

other P. ochraceus

orange P. ochraceus

Total Pisaster ochraceus

Mean Number Per Plot

0

20

40

60

80

0

10

20

30

0

10

20

30

0

10

20

30

68
Old Stairs

Figure 13d. Seastar abundances and 

Pisaster ochraceus colors. Old Stairs was

not sampled in F00 or S01. Crystal Cove

was not sampled in S00. Note differences in 

scale. S=spring samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Crystal Cove

Point Fermin

Paradise Cove

other P. ochraceus

orange P. ochraceus

Total Pisaster ochraceus

Mean Number Per Plot

0

20

40

60

80

100

0

10

20

30

0

20

40

60

80

100

0

10

20

30

Old Stairs

Figure 13d. Seastar abundances and 

Pisaster ochraceus colors. Old Stairs was

not sampled in F00 or S01. Crystal Cove

was not sampled in S00. Note differences in 

scale. S=spring samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Crystal Cove

Point Fermin

Paradise Cove

other P. ochraceus

orange P. ochraceus

Total Pisaster ochraceus

Mean Number Per Plot

0

20

40

60

80

100

0

20

40

60

80

100

0

10

20

30

0

20

40

60

80

100

0

20

40

60

80

100

0

10

20

30

69
Shaw’s Cove

Figure 13e. Seastar abundances and 

Pisaster ochraceus colors. Note differences 

in scale. S=spring samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Treasure Island

other P. ochraceus

orange P. ochraceus

Total Pisaster ochraceus

Mean Number Per Plot

Dana Point

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

5

10

15

20

Shaw’s Cove

Figure 13e. Seastar abundances and 

Pisaster ochraceus colors. Note differences 

in scale. S=spring samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Treasure Island

other P. ochraceus

orange P. ochraceus

Total Pisaster ochraceus

Mean Number Per Plot

Dana Point

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

5

10

15

20

70
0

20

40

60

80

100

120

140

160

0

20

40

60

80

100

120

140

160

0

20

40

60

80

100

120

140

160

Figure 14a. Pisaster ochraceus mean size. Carpinteria was 

not sampled in F01, F02, or F03. S=spring samples, F=fall 

samples.

Other

Orange

F00

S01

F01

S02

F02

S03

F03

F00

S01

F01

S02

F02

S03

F03

F00

S01

F01

S02

F02

S03

F03

Point Sierra

Nevada

Shell

Beach

Cayucos Hazards

Occulto

Boat House Government Point

Stairs

Arroyo Hondo

Alegria

Carpinteria

Mean Pisaster Size (mm)

Mussel 

Shoals

0

20

40

60

80

100

120

140

160

0

20

40

60

80

100

120

140

160

0

20

40

60

80

100

120

140

160

0

20

40

60

80

100

120

140

160

0

20

40

60

80

100

120

140

160

0

20

40

60

80

100

120

140

160

0

20

40

60

80

100

120

140

160

0

20

40

60

80

100

120

140

160

0

20

40

60

80

100

120

140

160

0

20

40

60

80

100

120

140

160

Figure 14a. Pisaster ochraceus mean size. Carpinteria was 

not sampled in F01, F02, or F03. S=spring samples, F=fall 

samples.

Other

Orange

Other

Orange

Other

Orange

F00

S01

F01

S02

F02

S03

F03

F00

S01

F01

S02

F02

S03

F03

F00

S01

F01

S02

F02

S03

F03

F00

S01

F01

S02

F02

S03

F03

F00

S01

F01

S02

F02

S03

F03

F00

S01

F01

S02

F02

S03

F03

Point Sierra

Nevada

Shell

Beach

Cayucos Hazards

Occulto

Boat House Government Point

Stairs

Arroyo Hondo

Alegria

Carpinteria

Mean Pisaster Size (mm)

Mussel 

Shoals

0

20

40

60

80

100

120

140

160

0

20

40

60

80

100

120

140

160

71
72

0

20

40

60

80

100

120

140

160

Figure 14b. Pisaster ochraceus mean size. S=spring

samples, F=fall samples. Other

Orange

F00

S01

F01

S02

F02

S03

F03

F00

S01

F01

S02

F02

S03

F03

F00

S01

F01

S02

F02

S03

F03

Old Stairs Paradise Cove Point Fermin

Mean Size (mm)

0

20

40

60

80

100

120

140

160

0

20

40

60

80

100

120

140

160

0

20

40

60

80

100

120

140

160

Figure 14b. Pisaster ochraceus mean size. S=spring

samples, F=fall samples. Other

Orange

Other

Orange

Other

Orange

F00

S01

F01

S02

F02

S03

F03

F00

S01

F01

S02

F02

S03

F03

F00

S01

F01

S02

F02

S03

F03

F00

S01

F01

S02

F02

S03

F03

F00

S01

F01

S02

F02

S03

F03

F00

S01

F01

S02

F02

S03

F03

Old Stairs Paradise Cove Point Fermin

Mean Size (mm)
Figure 15a. Pisaster ochraceus size distributions by color. Figures show percent 

of total (y-axes vary by plot) and do not reflect abundance. 

COLOR 

10 80 150 220 10 80 150 220 10 80 150 220 10 80 150 220 1

Pt. Sierra

Nevada

Cayucos Hazards Shell Beach 

FA00

SP01

FA01

SP02

FA02

SP03

FA03

Size (mm) Size (mm) Size (mm) Size (mm) 

Figure 15a. Pisaster ochraceus size distributions by color. Figures show percent of total (y-axes vary by plot) and do not reflect abundance. 

COLOR COLOR 

10 80 150 220 10 80 150 220 10 80 150 220 10 80 150 220 1

Pt. Sierra

Nevada

Cayucos Hazards Shell Beach 

FA00

SP01

FA01

SP02

FA02

SP03

FA03FA00

SP01

FA01

SP02

FA02

SP03

FA03

Size (mm) Size (mm) Size (mm) Size (mm) 

73
10 80 150 220 10 80 150 220 10 80 150 220 10 80 150 220 10 

COLOR 

Stairs Boat House Government Pt. Alegria Ar

FA00

SP01

FA01

SP02

FA02

SP03

FA03

Size (mm) Size (mm) Size (mm) Size (mm) 

Figure 15b. Pisaster ochraceus size distributions by color. Figures show percentof total (y-axes vary by plot) and do not reflect abundance.

10 80 150 220 10 80 150 220 10 80 150 220 10 80 150 220 10 

COLOR COLOR 

Stairs Boat House Government Pt. Alegria Ar

FA00

SP01

FA01

SP02

FA02

SP03

FA03FA00

SP01

FA01

SP02

FA02

SP03

FA03

Size (mm) Size (mm) Size (mm) Size (mm) 

Figure 15b. Pisaster ochraceus size distributions by color. Figures show percentof total (y-axes vary by plot) and do not reflect abundance.

74
10 80 150 220 10 

10 80 150 220

10 80 150 220 10 80 150 220

C

Carpinteria Mussel Shoals Old Stairs Paradise Cove 

FA00

SP01

FA01

SP02

FA02

SP03

FA03

Size (mm) Size (mm) Size (mm) Size (mm) 

Figure 15c. Pisaster ochraceus size distributions by color. Figures show percent of total (y-axesvary by plot) and do not reflect abundance. Carpinteria was not sampled FA01, FA02, or FA03.

10 80 150 220 10 

10 80 150 220

10 80 150 220 10 80 150 220

CC

Carpinteria Mussel Shoals Old Stairs Paradise Cove 

FA00

SP01

FA01

SP02

FA02

SP03

FA03FA00

SP01

FA01

SP02

FA02

SP03

FA03

Size (mm) Size (mm) Size (mm) Size (mm) 

Figure 15c. Pisaster ochraceus size distributions by color. Figures show percent of total (y-axesvary by plot) and do not reflect abundance. Carpinteria was not sampled FA01, FA02, or FA03.

75
Rancho Marino

Figure 16a. Lottia gigantea abundances. Note differences in scale. S=spring 

samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Stairs

Hazards

Cayucos

Mean Number Per Plot

Boat House

0

20

40

60

0

25

50

75

100

125

0

10

20

30

40

0

20

40

60

0

20

40

60

80

Rancho Marino

Figure 16a. Lottia gigantea abundances. Note differences in scale. S=spring 

samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Stairs

Hazards

Cayucos

Mean Number Per Plot

Boat House

0

20

40

60

0

25

50

75

100

125

0

25

50

75

100

125

0

10

20

30

40

0

20

40

60

0

20

40

60

80

76
Mean Number Per Plot

Government Point

Mussel Shoals

Carpinteria

Alegria

Old Stairs

Figure 16b. Lottia gigantea abundances. Gov’t. Pt. was not sampled in F99. Note 

differences in scale. S=spring samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

0

20

40

60

80

Mean Number Per Plot

Government Point

Mussel Shoals

Carpinteria

Alegria

Old Stairs

Figure 16b. Lottia gigantea abundances. Gov’t. Pt. was not sampled in F99. Note 

differences in scale. S=spring samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

0

20

40

60

80

77
Figure 16c. Lottia gigantea abundances. One additional plot was added at 

Crystal Cove in S01. See text for details. Note differences in scale. S=spring

samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Mean Number Per Plot

Paradise Cove

Shaw’s Cove

Crystal Cove

Point Fermin

Dana Point

0

40

80

120

160

200

240

0

10

20

30

40

50

0

25

50

75

100

125

0

40

80

120

160

200

240

0

20

40

60

Figure 16c. Lottia gigantea abundances. One additional plot was added at 

Crystal Cove in S01. See text for details. Note differences in scale. S=spring

samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Mean Number Per Plot

Paradise Cove

Shaw’s Cove

Crystal Cove

Point Fermin

Dana Point

0

40

80

120

160

200

240

0

40

80

120

160

200

240

0

10

20

30

40

50

0

10

20

30

40

50

0

25

50

75

100

125

0

25

50

75

100

125

0

40

80

120

160

200

240

0

40

80

120

160

200

240

0

20

40

60

78
Rancho Marino

Hazards Stairs

Cayucos

Boat House Government Point

Alegria Carpinteria

Figure 17a. Lottia gigantea mean size. Only spring samples are labeled.

Unlabeled bars represent fall samples. Gov’t. Pt. was not sampled in FA99.

Mean Lottia Size (mm)

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

Rancho Marino

Hazards Stairs

Cayucos

Boat House Government Point

Alegria Carpinteria

Figure 17a. Lottia gigantea mean size. Only spring samples are labeled.

Unlabeled bars represent fall samples. Gov’t. Pt. was not sampled in FA99.

Mean Lottia Size (mm)

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

79
Mussel Shoals

Paradise Cove Point Fermin

Old Stairs

Crystal Cove Shaw’s Cove

Dana Point

Figure 17b. Lottia gigantea mean size. Only spring samples are labeled.

Unlabeled bars represent fall samples.

Mean Lottia Size (mm)

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

Mussel Shoals

Paradise Cove Point Fermin

Old Stairs

Crystal Cove Shaw’s Cove

Dana Point

Figure 17b. Lottia gigantea mean size. Only spring samples are labeled.

Unlabeled bars represent fall samples.

Mean Lottia Size (mm)

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

0

10

20

30

40

50

60

80
Rancho

Marino

Cayucos Hazards Stairs Boat House

SP92

FA92

SP93

FA93

SP94

FA94

SP95

FA95

SP96

FA96

SP97

FA97

SP98

FA98

SP99

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

Figure 18a. Lottia gigantea size distributions. SP=spring sample, FA=fall sample.

Rancho

Marino

Cayucos Hazards Stairs Boat House

SP92

FA92

SP93

FA93

SP94

FA94

SP95

FA95

SP96

FA96

SP97

FA97

SP98

FA98

SP99

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

SP92

FA92

SP93

FA93

SP94

FA94

SP95

FA95

SP96

FA96

SP97

FA97

SP98

FA98

SP99

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

Figure 18a. Lottia gigantea size distributions. SP=spring sample, FA=fall sample.

81
0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

Government

Pt.

Alegria Carpinteria Mussel

Shoals

Old Stairs

Figure 18b. Lottia gigantea size distributions. Government Pt. was not sampled FA99

SP92

FA92

SP93

FA93

SP94

FA94

SP95

FA95

SP96

FA96

SP97

FA97

SP98

FA98

SP99

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

Government

Pt.

Alegria Carpinteria Mussel

Shoals

Old Stairs

Figure 18b. Lottia gigantea size distributions. Government Pt. was not sampled FA99

SP92

FA92

SP93

FA93

SP94

FA94

SP95

FA95

SP96

FA96

SP97

FA97

SP98

FA98

SP99

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

SP92

FA92

SP93

FA93

SP94

FA94

SP95

FA95

SP96

FA96

SP97

FA97

SP98

FA98

SP99

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

82
0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

Paradise

Cove

Pt. Fermin Crystal

Cove

Shaw’s

Cove

Dana Pt.

Figure 18c. Lottia gigantea size distributions. SP=spring sample, FA=fall sample.

SP92

FA92

SP93

FA93

SP94

FA94

SP95

FA95

SP96

FA96

SP97

FA97

SP98

FA98

SP99

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

0 45 90

Size (mm)

Paradise

Cove

Pt. Fermin Crystal

Cove

Shaw’s

Cove

Dana Pt.

Figure 18c. Lottia gigantea size distributions. SP=spring sample, FA=fall sample.

SP92

FA92

SP93

FA93

SP94

FA94

SP95

FA95

SP96

FA96

SP97

FA97

SP98

FA98

SP99

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

SP92

FA92

SP93

FA93

SP94

FA94

SP95

FA95

SP96

FA96

SP97

FA97

SP98

FA98

SP99

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

83
Cayucos

Rancho Marino

Piedras Blancas

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Point Sierra Nevada

Figure 19a. Black abalone abundances. Note differences in scale. Two additional 

plots were added at PSN in S97. One plot was added at Piedras in S00. See text

for details. Plot 4 was not sampled at PSN F97. S=spring samples, F=fall samples.

Total Number (all plots combined)

0

100

200

300

400

500

0

50

100

150

200

250

0

10

20

30

40

50

60

70

0

5

10

15

20

Juveniles (<50mm)

Adults (>50mm)

Total

Cayucos

Rancho Marino

Piedras Blancas

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Point Sierra Nevada

Figure 19a. Black abalone abundances. Note differences in scale. Two additional 

plots were added at PSN in S97. One plot was added at Piedras in S00. See text

for details. Plot 4 was not sampled at PSN F97. S=spring samples, F=fall samples.

Total Number (all plots combined)

0

100

200

300

400

500

0

100

200

300

400

500

0

50

100

150

200

250

0

50

100

150

200

250

0

10

20

30

40

50

60

70

0

10

20

30

40

50

60

70

0

5

10

15

20

Juveniles (<50mm)

Adults (>50mm)

Total

Juveniles (<50mm)

Adults (>50mm)

Total

84
Government Point

Boat House

Stairs

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Purisima

Figure 19b. Black abalone abundances. Note differences in scale. One additional 

plot was added at Stairs in S98. See text for details. Purisima was not sampled 

F94, and Gov’t. Pt. was not sampled S93 or F99. Plot 1 was not sampled at Gov’t.

Pt. S92. S=spring samples, F=fall samples.

0

100

200

300

400

500

600

0

100

200

300

400

500

600

0

100

200

300

400

500

600

0

10

20

30

40

50

60

70

80

90

Juveniles (<50mm)

Adults (>50mm)

Total

Total Number (all plots combined)

Government Point

Boat House

Stairs

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Purisima

Figure 19b. Black abalone abundances. Note differences in scale. One additional 

plot was added at Stairs in S98. See text for details. Purisima was not sampled 

F94, and Gov’t. Pt. was not sampled S93 or F99. Plot 1 was not sampled at Gov’t.

Pt. S92. S=spring samples, F=fall samples.

0

100

200

300

400

500

600

0

100

200

300

400

500

600

0

100

200

300

400

500

600

0

100

200

300

400

500

600

0

100

200

300

400

500

600

0

100

200

300

400

500

600

0

10

20

30

40

50

60

70

80

90

0

10

20

30

40

50

60

70

80

90

Juveniles (<50mm)

Adults (>50mm)

Total

Juveniles (<50mm)

Adults (>50mm)

Total

Total Number (all plots combined)

85
Mean Abalone Size (mm) 60

80

100

120

140

60

80

100

120

140

120

140

60

80

100

60

80

100

120

140

60

80

100

120

140

60

80

100

120

140

140

60

80

100

120

60

80

100

120

140

Point Sierra Nevada

Rancho Marino Cayucos

Piedras

Purisima Stairs

Boat House Government Point

Figure 20. Black abalone mean size. Note that y-axis starts at 60 mm. Only

spring samples are labeled. Unlabeled bars represent fall samples. Purisima was

not sampled FA94, and Gov’t. Pt. was not sampled SP93 or FA99.

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

Mean Abalone Size (mm) 60

80

100

120

140

60

80

100

120

140

60

80

100

120

140

60

80

100

120

140

120

140

60

80

100

120

140

60

80

100

60

80

100

120

140

60

80

100

120

140

60

80

100

120

140

60

80

100

120

140

60

80

100

120

140

60

80

100

120

140

140

60

80

100

120

60

80

100

120

60

80

100

120

140

60

80

100

120

140

Point Sierra Nevada

Rancho Marino Cayucos

Piedras

Purisima Stairs

Boat House Government Point

Figure 20. Black abalone mean size. Note that y-axis starts at 60 mm. Only

spring samples are labeled. Unlabeled bars represent fall samples. Purisima was

not sampled FA94, and Gov’t. Pt. was not sampled SP93 or FA99.

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

SP 1993

SP 1994

SP 1996

SP 1997

SP 1998

SP 1999

SP 1992

SP 2000

SP 2001

SP 2002

SP 2003

SP 1995

86
Pt. Sierra

Nevada

Piedras 

Blancas

Rancho

Marino

Cayucos

Figure 21a. Black abalone size distributions. Figures show proportion of total (y-axes 

vary by plot) and do not reflect abundance. See text for changes in sampling over time.

FA95

SP96

FA96

SP97

FA97

SP98

FA98

SP99

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

Size (mm)

0

50

100

150

200

Size (mm)

0

50

100

150

200

Size (mm)

0

50

100

150

200

Size (mm)

0

50

100

150

200

Pt. Sierra

Nevada

Piedras 

Blancas

Rancho

Marino

Cayucos

Figure 21a. Black abalone size distributions. Figures show proportion of total (y-axes 

vary by plot) and do not reflect abundance. See text for changes in sampling over time.

FA95

SP96

FA96

SP97

FA97

SP98

FA98

SP99

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

Size (mm)

0

50

100

150

200

Size (mm)

0

50

100

150

200

Size (mm)

0

50

100

150

200

Size (mm)

0

50

100

150

200

Size (mm)

0

50

100

150

200

Size (mm)

0

50

100

150

200

Size (mm)

0

50

100

150

200

Size (mm)

0

50

100

150

200

87
Purisima Stairs Boat House Government Pt.

SP92

FA92

SP93

FA93

SP94

FA94

SP95

FA95

SP96

FA96

SP97

FA97

SP98

FA98

SP99

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

Figure 21b. Black abalone size distributions. Figures show proportion of total (y-axes vary

by plot) and do not reflect abundance. See text for changes in sampling over time.

Size (mm)

0

50

100

150

200

Size (mm)

0

50

100

150

200

Size (mm)

0

50

100

150

200

Size (mm)

0

50

100

150

200

Purisima Stairs Boat House Government Pt.

SP92

FA92

SP93

FA93

SP94

FA94

SP95

FA95

SP96

FA96

SP97

FA97

SP98

FA98

SP99

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

SP92

FA92

SP93

FA93

SP94

FA94

SP95

FA95

SP96

FA96

SP97

FA97

SP98

FA98

SP99

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

Figure 21b. Black abalone size distributions. Figures show proportion of total (y-axes vary

by plot) and do not reflect abundance. See text for changes in sampling over time.

Size (mm)

0

50

100

150

200

Size (mm)

0

50

100

150

200

Size (mm)

0

50

100

150

200

Size (mm)

0

50

100

150

200

Size (mm)

0

50

100

150

200

Size (mm)

0

50

100

150

200

Size (mm)

0

50

100

150

200

Size (mm)

0

50

100

150

200

88
Lepidochitona hartwegii

limpets

Tegula funebralis

0

20

40

60

80

100

0

100

200

300

0

20

40

60

80

0

50

100

150

200

0

100

200

300

0

50

100

150

0

50

100

150

0

10

20

30

Pt. Sierra Nevada

Cayucos

Hazards

Shell Beach

Stairs

Boat House

Government Pt.

Pt. Fermin

Figure 22. Common motile invertebrates in Silvetia plots. Note differences in 

scale.

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

Mean Nu

m

ber Per Plot

Lepidochitona hartwegii

limpets

Tegula funebralis

Lepidochitona hartwegii

limpets

Tegula funebralis

0

20

40

60

80

100

0

20

40

60

80

100

0

100

200

300

0

100

200

300

0

20

40

60

80

0

50

100

150

200

0

50

100

150

200

0

100

200

300

0

100

200

300

0

50

100

150

0

50

100

150

0

50

100

150

0

50

100

150

0

10

20

30

Pt. Sierra Nevada

Cayucos

Hazards

Shell Beach

Stairs

Boat House

Government Pt.

Pt. Fermin

Figure 22. Common motile invertebrates in Silvetia plots. Note differences in 

scale.

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

Mean Nu

m

ber Per Plot

89
0

100

200

300

400

0

100

200

300

400

500

0

100

200

300

400

500

600

0

100

200

300

0

100

200

300

0

50

100

150

0

50

100

150

200

0

100

200

300

400

500

0

50

100

150

0

100

200

300

400

500

Tegula funebralis

littorines

limpets

Figure 23. Common motile invertebrates in 

Endocladia plots. Note differences in scale.

Cayucos

Hazards

Shell Beach

Occulto

Stairs

Boat House

Government Pt.

Old Stairs

Paradise Cove

White’s Pt.

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

Mean Nu

mber P

er Plot

0

100

200

300

400

0

100

200

300

400

500

0

100

200

300

400

500

0

100

200

300

400

500

600

0

100

200

300

400

500

600

0

100

200

300

0

100

200

300

0

50

100

150

0

50

100

150

200

0

100

200

300

400

500

0

100

200

300

400

500

0

50

100

150

0

100

200

300

400

500

0

100

200

300

400

500

Tegula funebralis

littorines

limpets

Tegula funebralis

littorines

limpets

Figure 23. Common motile invertebrates in 

Endocladia plots. Note differences in scale.

Cayucos

Hazards

Shell Beach

Occulto

Stairs

Boat House

Government Pt.

Old Stairs

Paradise Cove

White’s Pt.

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

Mean Nu

mber P

er Plot

90
91

1

10

100

1

10

100

1

10

100

1000

1

10

100

1

10

100

1

10

100

1

10

100

1

10

100

1000

1

10

100

1000

1

10

100

1000

1

10

100

1000

1

10

100

1000

1

10

100

1000

1

10

100

1

10

100

1000

littorines

limpets

Figure 24. Common motile invertebrates in barnacle plots. Note log scale and variation

in scale among plots.

* Indicates 0 limpets were found

Mean Nu

mber Per Plot

Pt. Sierra

Nevada

Cayucos

Hazards

Shell Beach

Occulto

Stairs

Boat House

Government Pt.

Alegria

Arroyo Hondo

Carpinteria

Mussel Shoals

Old Stairs

Paradise Cove

White’s Pt.

Pt. Fermin

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

*

1

10

100

1000

1

10

100

1

10

100

1

10

100

1000

1

10

100

1

10

100

1

10

100

1

10

100

1

10

100

1000

1

10

100

1000

1

10

100

1000

1

10

100

1000

1

10

100

1000

1

10

100

1000

1

10

100

1000

1

10

100

1000

1

10

100

1

10

100

1000

1

10

100

1000

littorines

limpets

littorines

limpets

Figure 24. Common motile invertebrates in barnacle plots. Note log scale and variation

in scale among plots.

* Indicates 0 limpets were found

Mean Nu

mber Per Plot

Pt. Sierra

Nevada

Cayucos

Hazards

Shell Beach

Occulto

Stairs

Boat House

Government Pt.

Alegria

Arroyo Hondo

Carpinteria

Mussel Shoals

Old Stairs

Paradise Cove

White’s Pt.

Pt. Fermin

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

*

1

10

100

1000
92

Figure 25. Common motile invertebrates in Mytilus plots. Note differences in scale. 

0

5

10

15

20

25

0

1

2

3

4

5

0

2

4

6

8

10

12

0

10

20

30

0

10

20

30

0

5

10

15

0

1

2

3

4

5

6

7

0

10

20

30

0

20

40

60

80

100

120

0

2

4

6

8

10

12

0

5

10

15

Nuttallina spp.

Nucella spp.

Tegula funebralis

0

2

4

6

8

0

5

10

15

20

0

50

100

150

0

5

10

15

20

0

5

10

15

20

0

5

10

15

20

Mean Num

ber P

er Plot

Pt. Sierra Nevada

Cayucos

Hazards

Shell Beach

Occulto

Stairs

Boat House

Government Pt.

Alegria

Arroyo Hondo

Coal Oil Pt.

Carpinteria

Mussel Shoals

Old Stairs

Paradise Cove

White’s Pt.

Pt. Fermin

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

Figure 25. Common motile invertebrates in Mytilus plots. Note differences in scale. 

0

5

10

15

20

25

0

1

2

3

4

5

0

2

4

6

8

10

12

0

10

20

30

0

10

20

30

0

5

10

15

20

25

0

5

10

15

20

25

0

1

2

3

4

5

0

1

2

3

4

5

0

2

4

6

8

10

12

0

2

4

6

8

10

12

0

10

20

30

0

10

20

30

0

5

10

15

0

1

2

3

4

5

6

7

0

1

2

3

4

5

6

7

0

10

20

30

0

20

40

60

80

100

120

0

20

40

60

80

100

120

0

2

4

6

8

10

12

0

2

4

6

8

10

12

0

5

10

15

Nuttallina spp.

Nucella spp.

Tegula funebralis

Nuttallina spp.

Nucella spp.

Tegula funebralis

0

2

4

6

8

0

5

10

15

20

0

50

100

150

0

5

10

15

20

0

5

10

15

20

0

5

10

15

20

0

2

4

6

8

0

5

10

15

20

0

50

100

150

0

50

100

150

0

5

10

15

20

0

5

10

15

20

0

5

10

15

20

Mean Num

ber P

er Plot

Pt. Sierra Nevada

Cayucos

Hazards

Shell Beach

Occulto

Stairs

Boat House

Government Pt.

Alegria

Arroyo Hondo

Coal Oil Pt.

Carpinteria

Mussel Shoals

Old Stairs

Paradise Cove

White’s Pt.

Pt. Fermin

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03

FA99

SP00

FA00

SP01

FA01

SP02

FA02

SP03

FA03
SP00

FA00SP01FA01SP02FA02SP03FA03

0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 

Size (mm)

Pt. Sierra

Nevada

Cayucos Occulto Stairs Boat House 

Figure 26a. Nucella emarginata/ostrina size distributions in mussel plots. Figures show proportion of toplot) and do not reflect abundance. Sites not shown had insufficient numbers of Nucella for plotting siz

Hazards

SP00FA00SP01FA01SP02FA02SP03FA03SP00FA00SP01FA01SP02FA02SP03FA03

0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 

Size (mm)

Pt. Sierra

Nevada

Cayucos Occulto Stairs Boat House 

Figure 26a. Nucella emarginata/ostrina size distributions in mussel plots. Figures show proportion of toplot) and do not reflect abundance. Sites not shown had insufficient numbers of Nucella for plotting siz

Hazards

93
0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 0 10 2

Size (mm)

Alegria Arroyo Hondo Coal Oil

Point

Carpinteria Mussel

Shoals

Old Stairs Para

Co

SP00FA00SP01FA01SP02FA02SP03FA03

Figure 26b. Nucella emarginata/ostrina size distributions in mussel plots. Figures show proportion of plot) and do not reflect abundance. Sites not shown had insufficient numbers of Nucella for plotting s

0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 0 10 2

Size (mm)

0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 0 10 20 30 40 0 10 2

Size (mm)

Alegria Arroyo Hondo Coal Oil

Point

Carpinteria Mussel

Shoals

Old Stairs Para

Co

SP00FA00SP01FA01SP02FA02SP03FA03SP00FA00SP01FA01SP02FA02SP03FA03

Figure 26b. Nucella emarginata/ostrina size distributions in mussel plots. Figures show proportion of plot) and do not reflect abundance. Sites not shown had insufficient numbers of Nucella for plotting s

94
Figure 27a. Tegula funebralis size distributions in mussel and Silvetia plots. Figures show proportion o

by plot) and do not reflect abundance. Sites not shown had insufficient numbers of Tegula for plotting s

SP00FA00SP01FA01SP02FA02SP03FA03

Pt. Sierra Nevada Cayucos

Mussel Silvetia

Hazards

Mussel Mussel Silvetia

She

Mussel 

0 10 20 30 0 10 20 30 0 10 20 30 0 10 20 30 0 10 20 30 0 10 20 

Size (mm)

Figure 27a. Tegula funebralis size distributions in mussel and Silvetia plots. Figures show proportion o

by plot) and do not reflect abundance. Sites not shown had insufficient numbers of Tegula for plotting s

SP00FA00SP01FA01SP02FA02SP03FA03SP00FA00SP01FA01SP02FA02SP03FA03

Pt. Sierra Nevada Cayucos

Mussel Silvetia

Hazards

Mussel Mussel Silvetia

She

Mussel 

0 10 20 30 0 10 20 30 0 10 20 30 0 10 20 30 0 10 20 30 0 10 20 

Size (mm)

95
Figure 27b. Tegula funebralis size distributions in mussel and Silvetia plots. Figures show proportion o

by plot) and do not reflect abundance. Sites not shown had insufficient numbers of Tegula for plotting s

SP00FA00SP01FA01SP02FA02SP03FA03

0 10 20 30 0 10 20 30 0 10 20 30 0 10 20 30 0 10 20 30 0 

Stairs

Mussel Silvetia

Boat House

Mussel Silvetia

Government P

Mussel S

Size (mm)

Figure 27b. Tegula funebralis size distributions in mussel and Silvetia plots. Figures show proportion o

by plot) and do not reflect abundance. Sites not shown had insufficient numbers of Tegula for plotting s

SP00FA00SP01FA01SP02FA02SP03FA03SP00FA00SP01FA01SP02FA02SP03FA03

0 10 20 30 0 10 20 30 0 10 20 30 0 10 20 30 0 10 20 30 0 

Stairs

Mussel Silvetia

Boat House

Mussel Silvetia

Government P

Mussel S

Size (mm)

96
0 10 20 30 

Figure 27c. Tegula funebralis size distributions in mussel and Silvetia plots. Figures show proportion o

by plot) and do not reflect abundance. Sites not shown had insufficient numbers of Tegula for plotting 

SP00FA00SP01FA01SP02FA02SP03FA03

0 10 20 30 0 10 20 30 0 10 20 30 0 10 20 30 0 10 20 

Size (mm)

Alegria Arroyo Hondo

Mussel

Coal Oil Pt.

Mussel Mussel

Carpinteria

Mussel

Mussel Shoals

Mussel

Paradise Co

Mussel

0 10 20 30 

Figure 27c. Tegula funebralis size distributions in mussel and Silvetia plots. Figures show proportion o

by plot) and do not reflect abundance. Sites not shown had insufficient numbers of Tegula for plotting 

SP00FA00SP01FA01SP02FA02SP03FA03SP00FA00SP01FA01SP02FA02SP03FA03

0 10 20 30 0 10 20 30 0 10 20 30 0 10 20 30 0 10 20 

Size (mm)

Alegria Arroyo Hondo

Mussel

Coal Oil Pt.

Mussel Mussel

Carpinteria

Mussel

Mussel Shoals

Mussel

Paradise Co

Mussel

97
LITERATURE CITED 

Abbott, I.A. and G.J. Hollenberg. 1976. Marine algae of California. Stanford University 

Press, Stanford, California. 827 pp. 

Altstatt, J. A., R. F. Ambrose, J. M. Engle, P. L. Haaker, K. D. Lafferty and P. T. 

Raimondi. 1996. Recent declines of black abalone Haliotis cracherodii on the 

mainland coast of central California. Mar. Ecol. Prog. Ser. 142: 185-192. 

Ambrose, R.F., P.T. Raimondi and J.M. Engle. 1992. Final Study Plan for Inventory of 

Intertidal Resources in Santa Barbara County. Report to the Minerals Management 

Service, Pacific OCS Region. January 1992. 

Ambrose, R.F., J.M. Engle, P.T. Raimondi, M. Wilson and J. Altstatt 1995. Rocky 

Intertidal and Subtidal Resources: Santa Barbara County Mainland. Final Report to 

the Pacific Outer Continental Shelf Region of the Minerals Management Service. 

OCS Study MMS 95-0067 

Barry, J.P., C.H. Baxter, R.D. Sagarin and S.E. Gilman. 1995. Climate-related, long-term

faunal change in a California rocky intertidal community. Science 267:672-675. 

Blanchette, C.A., B.G Miner, S.D. Gaines. 2002. Geographic variability in form, size and 

survival of Egregia menziesii around Point Conception, California. Marine EcologyProgress Series. 239:69-82 

Carlton, J. T., and J. B. Geller. 1993. Ecological roulette: The global transport of 

nonindigenous marine organisms. Science 261:78-82. 

Chambers Group, Inc. 1993. Santa Barbara County shoreline inventory database. 

Prepared for the Energy Division, Resource Management Department, County of 

Santa Barbara, California. 

Dawson, E. Y. 1959. A primary report on the benthic marine flora of southern 

California. In: An oceanographic and biological survey of the continental shelf area 

of southern California. Publs. Calif. State Wat. Poll. Contr. Bd., Volume 20: 169-

264. 

Dawson, E. Y. 1965. Intertidal algae. In: An oceanographic and biological survey of 

the southern California mainland shelf. Publs. Calif. State Wat. Qual. Contr. Bd. 

Volume 27: 220-231, 351-438. 

Engle, J.M., J.M. Altstatt, P.T. Raimondi and R.F. Ambrose. 1994. Rocky Intertidal 

Monitoring Handbook for Inventory of Intertidal Resources in Santa Barbara 

County. Report to the Minerals Management Service, Pacific OCS Region. 

February 1994. 

Engle, J.M., L. Gorodezky, K.D. Lafferty, R.F. Ambrose, and P.T. Raimondi. 1994b. 

First year study plan for inventory of coastal ecological resources of the northern 

98
Channel Islands and Ventura/Los Angeles Counties. California Coastal Commission. 

31p. 

Engle, J.M., K.D. Lafferty, J.E. Dugan, D.L. Martin, N. Mode, R.F. Ambrose, and P.T. 

Raimondi. 1995. Second year study plan for inventory of coastal ecological 

resources of the northern Channel Islands and Ventura/Los Angeles Counties. 

California Coastal Commission. 46p. 

Engle, J.M., R.F. Ambrose, P.T. Raimondi, S.N. Murray, M. Wilson, and S. Sapper. 

1998a. Rocky intertidal resources in San Luis Obispo, Santa Barbara, and Orange 

Counties. 1997 Annual Report. OCS Study, MMS 98-0011, U.S. Minerals 

Management Service, Pacific OCS Region. 73p. + appendices 

Friedman, C. S., K. B. Andree, K. A. Beauchamp, J. D. Moore, T. T. Robbins, J. D. 

Shields, and R. P. Hedrick. 2000. 'Candidatus Xenohaliotis californiensis', a newly 

described pathogen of abalone, Haliotis spp., along the west coast of North America. 

International Journal of Systematic and Evolutionary Microbiology 50:847-855. 

Hickey, B. M. 1993. Physical oceanography. Pages 19-70 in M. D. Dailey, D. J. Reish, 

and J. W. Anderson, eds. Ecology of the Southern California Bight. A synthesis and 

interpretation. University of California Press, Berkeley. 

Kinnetics Laboratories, Inc. 1985. Field Survey Plan for successional and seasonal 

variation of the central and northern California rocky intertidal communities as 

related to natural and man-induced disturbances. Report to the Minerals Management 

Service. 

Kinnetics Laboratories, Inc. 1992. Study of the rocky intertidal communities of Central 

and Northern California. Report to the Minerals Management Service. OCS Study 

MMS 91-0089. 

Lafferty, K.D.and A.M. Kuris. 1993. Mass mortality of abalone Haliotis cracherodii on 

the California Channel Islands: tests of epidemiological hypotheses. Mar. Ecol. Prog. 

Ser. 96: 239-248. 

Littler, M.M., ed. 1979. The distribution, abundance, and community structure of rocky 

intertidal and tidepool biotas in the Southern California Bight. Southern California 

Baseline Study. Final Report, Vol. II, Rep. 1.0. U.S. Dept. of Interior, Bureau of 

Land Management, Washington, D.C. 

McGowan, J. A., D. R. Cayan, and L. M. Dorman. 1998. Climate-ocean variability and 

ecosystem response in the northeast Pacific. Science 281: 210-217. 

Murray, S.N. and M.M. Littler. 1981. Biogeographical analysis of intertidal macrophyte 

floras of southern California. J. Biogeography 8: 339-351. 

99
Murray, S.N., and R.N. Bray. 1993. Benthic macrophytes. Pages 304-368 in M.D. Dailey, 

D.J. Reish, and J.W. Anderson, eds. Ecology of the Southern California Bight: A 

synthesis and interpretation. University of California Press, Berkeley. 

Murray, S. N. 1998. Effectiveness of marine life refuges on southern California shores. 

Pp. 1453-1465, In: O. T. Magoon, H. Converse, B. Baird, and M. Miller-Henson, 

eds. California and the world ocean ’97. Taking a look at California’s ocean 

resources: an agenda for the future. American Society of Civil Engineers, Reston, 

Virginia. 

Murray, S. N., T. G., Denis, J. S. Kido, and J. R. Smith. 1999. Human visitation and the 

frequency and potential effects of collecting on rocky intertidal populations in 

southern California marine reserves. Calif. Coop. Oceanic Fish. Invest. Rept. 40:100-

106. 

Nicholson, N. L. and R. L. Cimberg. 1971. The Santa Barbara oil spills of 1969. A postspill survey of the rocky intertidal. Pp. 325-399, In: D. S. Straughan, comp. 

Biological and oceanographical survey of the Santa Barbara Channel oil spill 1969-

1970. Vol. 1. Allan Hancock Foundation, Univ. of Southern California, Los 

Angeles. 

North, W.J., E.K. Anderson, and F.A. Chapman. 1989. Wheeler J. North ecological 

studies at Diablo Canyon Power Plant. Final Report, 1967-1987. Pacific Gas and 

Electric Company, San Francisco, CA. Pacific Gas and Electric Company 1988. 

Thermal effects monitoring program. Diablo Canyon Power Plant. Final Report. 

May 1988 

Pacific Gas and Electric Company 1994. Thermal effects monitoring program. Diablo 

Canyon Power Plant. 1993 Annual Report. March 1994 

Raimondi, P.T., R.F. Ambrose, J.M. Engle, S.N. Murray, and M. Wilson. 1999. 

Monitoring of rocky intertidal resources along the central and southern California 

mainland. 3-Year Report for San Luis Obispo, Santa Barbara, and Orange Counties 

(Fall 1995-Spring 1998). OCS Study, MMS 99-0032, U.S. Minerals Management 

Service, Pacific OCS Region. 

Raimondi, P.T., C.M. Wilson, R.F. Ambrose, J.M. Engle, and T.E. Minchinton. 2002. 

Continued declines of black abalone along the coast of California: are mass 

mortalities related to El Nino events? Mar. Ecol. Prog. Ser. 242:143-152 

Richards, D.V. and G.E. Davis. 1988. Rocky intertidal communities monitoring 

handbook, Channel Islands National Park, California. National Park Service, 

Ventura, CA. 15 p. 

Richards, D.V. and G.E. Davis. 1993. Early warnings of modern population collapse in 

black abalone Haliotis cracherodii, Leach, 1814 at the California Channel Islands. J. 

Shellfish Res. 12(2):189-194. 

100
Sagarin, R., J. Barry, S. Gilman, and C. Baxter. 1999. Climate related changes in an 

intertidal community over short and long time scales. Ecological Monographs 69: 

465-490 

Schiel DR, J.R. Steinbeck, and M.S. Foster 2004. Ten years of induced ocean warming 

causes comprehensive changes in marine benthic communities. Ecology 85(7): 833-

1839 

Thom, R. M. and T. B. Widdowson. 1978. A resurvey of E. Yale Dawson’s 42 

intertidal algal transects on the southern California mainland after 15 years. Bull. 

South. Calif. Acad. Sci. 77: 1-13. 

Widdowson, T. B. 1971. Changes in the intertidal algal flora of the Los Angeles area 

since the survey by E. Yale Dawson in 1956-1959. Bull. South. Calif. Acad. Sci., 70: 2-

16. 

101
Appendix A: Catalina Island Sites 

DESCRIPTION OF REGION 

Santa Catalina Island, southeastern-most of the eight Channel Islands, lies 35 km

southwest of Los Angeles. The island coast is rugged, with steep volcanic or 

metamorphic cliffs backing narrow intertidal shores. Rocky intertidal habitats encompass 

85% of Catalina Island; of these, 58% are boulder habitats and 42% are narrow bedrock 

benches (Littler and Littler 1979). 15% of island shores are pocket sand or gravel 

beaches. Mainland-facing shores typically are sheltered from prevailing oceanic swells, 

while windward shores are exposed to open-ocean waves, except where headlands protect 

various coves. 

Intertidal communities at Catalina Island, as well as at San Clemente Island to the south, 

are composed of warm-temperate biota, with fewer cold-water species than the 

northwestern Channel Islands and mainland shores north of Point Conception (Dykzeul 

and Given 1979, Coyer and Engle 1981). Rocky intertidal monitoring surveys include the 

U.S Bureau of Land Management surveys conducted in the 1970’s (Littler 1980) and 

studies associated with the University of Southern California’s Wrigley Marine 

Laboratory (e.g., Given and Lees 1967). Target species monitoring has taken place at 

Bird Rock, a small, lee-side, volcanic islet near the marine lab, since 1982, when the site 

was established to test Channel Islands National Park Service long-term monitoring 

protocols (VTN Oregon 1983). In 1994, the Bird Rock monitoring protocol was modified 

to fully conform to the MARINe protocol and a second island monitoring site was 

established on the windward side at Little Harbor. This site is located on a narrow, 

metamorphic promontory between two coves, with partial protection from prevailing 

swells afforded by headlands and subtidal reefs. 

SAMPLING METHODS 

Methods used for sampling the Catalina Island sites are identical to those used in other 

areas (see Sampling Methods p.16) 

RESULTS 

Below are summaries of the major trends in target species abundances at the two Catalina 

Island sites. Values for means and standard errors are given in Appendix C. 

Silvetia compressa subsp. deliquescens (Rockweed, formally called Pelvetia compressa 

and P. fastigiata. The Channel Islands subspecies of Silvetia is differentiated from the 

mainland form by narrower thalli that are more regularly and more densely branched).

Silvetia cover remained relatively high at both Catalina Island sites over time (Fig. A1). 

Cover at Little Harbor increased slightly from Fall 1994 to Spring 2003 (from 65% to 

86%), while Bird Rock cover declined slightly during the same period (from 95% to 

77%). Cover at both sites declined during 1997-1998, coincident with El Niño conditions. 

Rockweed at Little Harbor has since exhibited greater recovery than that at Bird Rock. 

Both sites showed a slight trend for higher Silvetia cover in fall compared to spring. 

 A1 
Chthamalus fissus/dalli and Balanus glandula (Barnacles, nearly all white barnacles at 

the Catalina sites were Chthamalus).

Barnacle cover fluctuated over time at Little Harbor, ending slightly lower in Spring 2003 

(61%) compared to Fall 1994 (81%) (Fig. A2). At Bird Rock barnacle cover was 

relatively stable (53-66% cover range) until Spring 1999, followed by nearly continuous 

declines to a low of 28% cover by Spring 2003. 

Pollicipes polymerus (Goose Barnacle)

Although Pollicipes cover was fairly stable over time at Bird Rock, there was a clear 

inverse relationship between cover of goose barnacles and mussels in these plots (Fig. 

A3). Declines in mussel cover were associated with increases in Pollicipes cover and 

increases in mussels were correlated with goose barnacle declines. 

Pollicipes was not targeted for monitoring at Little Harbor. 

Mytilus californianus (Mussel)

Mussel cover was highest at Little Harbor and Bird Rock between Fall 1995 and Spring 

1997 (Fig. A4). Declines at both sites began in Fall 1997 and stabilized at around 20% 

cover over the following years. The peak period of mussel cover was the same as for 

mussels in the goose barnacle plots at Bird Rock (see above); however, subsequent 

mussel declines were less in these higher zone Pollicipes plots.

DISCUSSION 

Declines of rockweed and mussels at Little Harbor and Bird Rock during 1997-1998 

coincided with a severe El Niño oceanographic anomaly in which storms were more 

frequent, water temperatures increased substantially, and seawater nutrients declined 

(McPhaden 1999). These trends were apparent despite the separation (opposite sides of 

Catalina Island) and variation in exposure and substrate between the two sites (Little 

Harbor: windward exposure, metamorphic rock; Bird Rock: leeward exposure, volcanic 

rock). A similar decline occurred in the acorn barnacles at Little Harbor, but not at Bird 

Rock. Goose barnacles increased slightly during the 1997-1998 El Niño period. Perhaps 

mussels are better competitors for space in the higher zone goose barnacle/mussel plots at 

Bird Rock, but goose barnacles are more resistant to El Niño environmental changes. 

The target species trends at the two Santa Catalina Island sites provide good 

representation for population dynamics patterns under warm temperate conditions.

Comparison of these long-term datasets with those of other regional sites in the MultiAgency Rocky Intertidal Network will provide valuable insight into the effects of the 

1997-1998 El Niño, as well as other latitudinal and island/mainland oceanographic 

environmental differences. The data also highlight the importance of long-term periodic 

surveys, illustrating that some species dynamics may be predictable on a regional scale, 

but complex interactions unique at local scales operate as well. Therefore, the prudent 

plan for determining the effects of unforeseen human impacts such as oil spills is to 

maintain a program of periodic intertidal community dynamics assessments at 

representative regional sites. 

 A2 
LITERATURE CITED 

Coyer, J. and J. Engle. 1981. Santa Catalina Island: Subarea II. California Marine Waters 

Areas of Special Biological Significance Reconnaissance Survey Report. California 

State Water Resources Control Board Water Quality Monitoring Report No. 81-10. 

Dykzeul, J. and R. Given. 1979. Santa Catalina Island: Subareas I-IV. California Marine 

Waters Areas of Special Biological Significance Reconnaissance Survey Report. 

California State Water Resources Control Board Water Quality Monitoring Report 

No. 79-6. 

Given, R. and D. Lees. 1967. Santa Catalina Island biological survey. Allan Hancock 

Foundation Survey Report No. 1, University of Southern California, Los Angeles. 

Littler, M. 1980. Overview of the rocky intertidal systems of southern California. Pp. 

265-306 In (D. Power, ed.) The California Islands: Proceedings of a 

Multidisciplinary Symposium. Santa Barbara Natural History Museum, Santa 

Barbara. 

Littler, M. and D. Littler. 1979. Rocky intertidal island survey. Vol. II, Report 5.0 in 

Southern California Intertidal Survey Year III. Bureau of Land Management, Los 

Angeles. 

McPhaden, M.J. 1999. Genesis and evolution of the 1997-98 El Niño. Science 283:950-

954. 

VTN Oregon, Inc. 1983. Visitor impact and recovery on Channel Islands tidepools: final 

report. National Park Service, Ventura. 

 A3 
rock

Silvetia compressa

Little Harbor

Figure A1. Species abundances in Silvetia

plots. S=spring samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Bird Rock

Percent Cov

e

r

0

20

40

60

80

100

0

20

40

60

80

100

rock

Silvetia compressa

Little Harbor

Figure A1. Species abundances in Silvetia

plots. S=spring samples, F=fall samples.

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Bird Rock

Percent Cov

e

r

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

 A4 
Little Harbor

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Bird Rock

Percent Cov

e

r

Endocladia muricata

Figure A2. Species abundances in barnacle Balanus + Chthamalus

plots. S=spring samples, F=fall samples.

0

20

40

60

80

100

0

20

40

60

80

100 Little Harbor

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F 

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Bird Rock

Percent Cov

e

r

Endocladia muricata

Balanus + Chthamalus

Endocladia muricata

Figure A2. Species abundances in barnacle Balanus + Chthamalus

plots. S=spring samples, F=fall samples.

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

 A5 
Little Harbor

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Bird Rock

Percent Cov

e

r

0

20

40

60

80

100

0

20

40

60

80

100

Figure A4. Species abundances in Mytilus

plots. Little Harbor was not sampled F01.

S=spring samples, F=fall samples. rock

Pollicipes polymerus

Mytilus californianus

Figure A3. Species abundances in Pollicipes

plots. Little Harbor was not sampled in F01.

S=spring samples, F=fall samples. rock

Mytilus californianus

Pollicipes polymerus

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Percent Cov

e

r

0

20

40

60

80

100 Bird Rock

Little Harbor

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Bird Rock

Percent Cov

e

r

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

0

20

40

60

80

100

Figure A4. Species abundances in Mytilus

plots. Little Harbor was not sampled F01.

S=spring samples, F=fall samples. rock

Pollicipes polymerus

Mytilus californianus

rock

Pollicipes polymerus

Mytilus californianus

Figure A3. Species abundances in Pollicipes

plots. Little Harbor was not sampled in F01.

S=spring samples, F=fall samples. rock

Mytilus californianus

Pollicipes polymerus

rock

Mytilus californianus

Pollicipes polymerus

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

S F S F S F S F S F S F S F S F S F S F S F S F

1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003

Percent Cov

e

r

0

20

40

60

80

100

0

20

40

60

80

100 Bird Rock

 A6 
Appendix B: Natural History of Target Species 

In this section, we summarize the natural history of the target species used in this study. 

These brief discussions provide context for the selection of these species by including 

information on life history, ecological importance, and sensitivity to anthropogenic 

activities. Additional information and photographs can be found on the MARINe website 

(http://www.marine.gov/species.htm). 

Silvetia compressa ssp. compressa (formerly Pelvetia compressa and P. fastigiata)

The rockweed Silvetia compressa ssp. compressa, a conspicuous fucoid alga, can be 

locally abundant in dense patches in upper mid-tidal regions of southern California rocky 

shores that are partially protected from open surf. The typical mainland form is an olive 

green or yellowish brown plant about 30 cm long, composed of thick, narrow, 

dichotomous branches. A finer-branched, lighter-colored form (S. compressa f. gracilis) 

is more typical of the Channel Islands (Abbott and Hollenberg 1976; Silva et al. 2004). 

Silvetia plants are tough, resilient, and long-lived; however, recruitment is irregular, 

survivorship low, and individuals slow-growing (Gunnill 1980). 

Silvetia is a dominant perennial whose thick clumps provide shelter and protection from

desiccation for many animals that otherwise could not exist so high up on the shore (Hill 

1980; Gunnill 1983; Ricketts et al. 1985; Sapper and Murray 2003). Rockweeds are 

vulnerable to trampling impacts (Murray and Denis 1997) and oil spills because of their 

location fairly high on the shore. Specific sensitivity of Silvetia to oiling is unclear, but 

other fucoids are known to be adversely affected (see Foster et al. 1988). Recovery from

impacts could take several years or more (Hill 1980; Vesco & Gillard 1980; Engle 

unpub.). 

Hesperophycus californicus (formerly H. harveyanus)

Hesperophycus is a common fucoid alga along the central coast of California, found in 

the upper-mid tidal regions sometimes mixed with Silvetia, Pelvetiopsis, or Fucus. Fucus

is replaced by Hesperophycus south of Pt. Conception. Plants are typically about 30 cm

long and greenish-olive to yellowish-brown with dichotomously branched blades. 

Hesperophycus can be distinguished from Fucus by the tiny tufts of white hairs, called 

cryptostomata, that grow in two parallel rows on either side of the midrib. Observations 

made during the Santa Barbara oil spill indicate that Hesperophycus is particularly 

susceptible to damage caused by oil pollution (Dawson and Foster 1982). 

Endocladia muricata 

Distinctive dark bands of the low-growing, red turfweed Endocladia muricata are 

characteristic of nearly all high rocky intertidal shores of the Pacific Coast north of Point 

Conception. Endocladia abundance fades in warmer waters to the south, being largely 

replaced in lower portions of its zone by other turfweeds (Gelidium spp.). 

B1
Endocladia forms dense 4-8 cm tall, dark red to blackish-brown perennial tufts made up 

of tiny spine-covered branchlets (Abbott & Hollenberg 1976). Together with spinybladed Mastocarpus papillatus, the Endocladia/Mastocarpus carpet traps sediment and 

seawater, thus providing a sheltered microhabitat for a host of small organisms, including 

other algae, worms, crustaceans, and mollusks. Glynn (1965) found over 90 species 

associated with Endocladia clumps in Monterey. Endocladia has been shown to facilitate 

recruitment of Silvetia compressa, possibly by providing propagules protection from

dislodgement, grazing, and/or desiccation (Johnson and Brawley 1998). Turfweed also 

can provide habitat for attachment of young mussels. Expanding mussel patches may 

displace Endocladia, but it can then grow on the mussel shells, creating a layered 

assemblage. 

Some Endocladia clumps appear donut- or crescent-shaped; this condition may be caused 

by storms tearing out center areas possibly weakened by accumulated anoxic sediment. 

At Diablo Canyon, Endocladia monitored quarterly for over 15 years tended to cycle 

between peak cover in summer and low abundance in winter (PG&E 1992). However, 

Kinnetics (1992) found few consistent seasonal patterns at 6 central California sites 

monitored in Spring and Fall for 6 years. 

Like Silvetia, Endocladia is hardy and quite resistant to desiccation, yet vulnerable to 

oiling from spills. Recovery from natural or human disturbances may vary from 1 to 

more than 6 years (see Kinnetics 1992). 

Mastocarpus papillatus 

Mastocarpus is an abundant dark reddish-black species which, along with Endocladia, 

forms a high-mid intertidal algal band that traps moisture and sediment, thus providing a 

refuge for a host of other intertidal organisms including snails, crustaceans, worms, and 

beach fly larvae (Dawson and Foster 1982). Plants rarely grow more than 15 cm tall and 

can be highly variable in form. Blade surfaces are covered with papillae that vary in 

density and size among growth forms (Carrington 1990). Papillae are typically 1-2 mm

in diameter, 1-3 mm long and are found on both sides of a thallus. No apparent 

correlation exists between density of papillae and any morphological trait of a thallus 

(Carrington 1990). Cystocarps, the female reproductive structures, are located on the 

papillae of female gametophytes (Zupan & West, 1988). 

Stress tests performed by Carrington (1990) revealed that mechanical failure most often 

occurred at the base of the stipe, leaving behind a small basal disc of tissue from which 

additional thalli could grow. Thus, perennial basal discs may give rise to many shorterlived thalli year after year. 

Mastocarpus exhibits heteromorphic alternation of generations. The blade form described 

above is the gametophytic stage. The sporophyte is a thick, black crust, which is 

commonly found in association with gametophytic blades, and was formerly classified in 

the genus Petrocelis. 

B2
Mazzaella spp. (formally Iridaea spp.)

Mazzaella forms a distinctive iridescent band in the mid to low intertidal of the central 

California rocky coast. Plants usually consist of several large (up to 1 m long), smooth, 

tapering blades that arise from a small crustose holdfast, although blade morphology 

varies with habitat and wave exposure (Mondragon and Mondragon 2003). Blades 

usually live less than one year, while basal crusts are longer lived (Dyck 1995). Frond 

regrowth from these basal crusts has been demonstrated in M. laminarioides, a species of 

Mazzaella common in southern Chile (Gomez 1991). Blade color can range from a 

greenish-olive to a deep purple. Several species of Mazzaella occur in SLO County, 

including M. splendens, M. flaccida and M. Heterocarpa; however, distinguishing these 

different species can be challenging, so they have been treated as a species group for this 

study. 

Mazzaella spp. have isomorphic sporangial and gametangial stages in life history. 

Seasonality in M. splendens abundance may be attributed to different strategies employed 

by the sporangial and gametangial stages (Dyck 1995). M. splendens gametophytes 

appear to exploit favorable spring and summer conditions, reaching high densities which 

help to compensate for greater loss in winter, while sporophytes maintain a generally 

lower, but somewhat more stable density throughout the year. 

The blades of Mazzaella are consumed by numerous grazers, including the snail Lacuna, 

the isopod Idotea, the chiton Katharina tunicata, the limpet Lottia pelta, and the sea 

urchin Strongylocentrotus purpuratus (Gaines 1985). 

Phyllospadix spp. 

Surfgrass (Phyllospadix spp.) is one of only two types of marine flowering plants on the 

West Coast. Unlike the eelgrass Zostera (often confused with surfgrass) that grows in 

quiet-water mud or sand habitats, surfgrass attaches by short roots to rock on surf-swept 

shores from the low intertidal down to 10-15 m depths. The 0.5-2 m tall, emerald green 

grass commonly occurs in dense perennial beds formed primarily by vegetative growth 

from spreading rhizomes. Two species (P. torreyi & P. scouleri) overlap in geographical 

distribution and morphological characteristics (see Dawson and Foster 1982). P. torreyi

generally has longer (1-2 m), narrower (1-2 mm) leaves, longer flower stems with several 

spadices, and occurs more in semi-protected habitats as well as at deeper depths. P. 

scouleri tends to have shorter (<50 cm), broader (2-4 mm) leaves, shorter flower stems 

with 1-2 spadices, and is found more often in wave-swept intertidal areas. 

Surfgrass meadows are highly productive ecosystems, providing structurally complex 

microhabitats for a rich variety of epiphytes, epibenthos, and infauna. Stewart and Myers 

(1980) identified 71 species of algae and 90 species of invertebrates associated with 

surfgrass habitats in San Diego. Some organisms, such as the red algae Smithora 

naiadum and Melobesia mediocris, are exclusive epiphytes on surfgrass (or eelgrass) 

(Abbott & Hollenberg 1976). Other species, such as the crabs Pugettia producta and 

Pachygrapsus crassipes are heavy consumers of Phyllospadix seeds (Holbrook et al. 

2000). Phyllospadix beds provide nursery habitat for various fishes and invertebrates, 

including the California spiny lobster Panulirus interruptus (Engle 1979). Green lobster 

B3
juveniles shelter in the thicket of leaves and forage on a variety of tiny gastropods and 

bivalves. 

Surfgrass cannot tolerate much heat or drying; the leaves will bleach quickly when 

midday low tides occur during hot, calm-water periods. Surfgrass can be particularly 

sensitive to sewage discharge (Littler and Murray 1975) and oil pollution (see Foster et 

al. 1988). Recovery can be relatively rapid if the rhizome systems remain functional, but 

might take many years if entire beds are lost, because recruitment is irregular and must be 

facilitated by the presence of perennial turf algae to which surfgrass seeds attach (Turner 

1983, 1985). Transplant projects undertaken to speed recovery of Phyllospadix beds 

destroyed by shoreline construction have been largely unsuccessful. 

Anthopleura elegantissima/sola (Anemone, formally lumped as Anthopleura 

elegantissima)

Previously thought to be a single species, Anthopleura elegantissima and Anthopleura 

sola were described as genetically, ecologically, and developmentally distinct by Pearse 

and Francis (2000). The two species are similar in appearance; however, A. sola grows 

larger (to 25 cm) and is solitary compared to the smaller (to 8 cm), aggregating A. 

elegantissima. Anthopleura elegantissima is able to persist practically indefinitely under 

normal conditions because genetically-identical individuals are periodically produced by 

longitudinal fission (Sebens 1982). A. sola does not divide and can be confused with A. 

xanthogrammica, a relative uncommon south of Point Conception. The green color of all 

of Anthopleura comes from symbiotic unicellular plants. 

Both A. sola and A. elegantissima are abundant throughout semi-protected rocky shores 

of the Pacific Coast. A. sola is common in tidepools and subtidally, and A. elegantissima

often occurs as small densely aggregated clones in middle intertidal zones, especially 

sand-influenced habitats (Morris et al. 1980). Extensive carpets of these clones may 

occur, but often go unrecognized under low tide conditions because the anemones 

contract to small sand or shell-covered blobs that provide protection from desiccation. 

Anemone mats create a moist microenvironment that allows the development of some

other species, such as coralline algae and sand tube worms (Phragmatopoma californica) 

at higher intertidal levels than they would normally occur (Taylor and Littler 1982). 

Adjacent anemone clones are separated by a narrow bare corridor caused by the 

withdrawal of non-clonemates following aggressive stinging encounters. 

Anthopleura species are quite resistant to disturbances from shifting sands. They not only 

withstand moderate sand abrasion, but also resist shallow sand burial by extending their 

columns to re-expose the tentacles and oral disk. If buried deeper, they can survive for at 

least 3 months by metabolizing body tissue (Sebens 1980). Anthopleura spp. are not 

known to be unusually sensitive to oiling. Recovery from major disturbances may take 1-

2 years or more (see Vesco & Gillard 1980). 

B4
Chthamalus fissus/dalli and Balanus glandula 

White acorn barnacles, Chthamalus fissus/dalli and Balanus glandula, typically dominate 

high intertidal zones along the West Coast. Chthamalus dalli and Balanus are most 

common in the colder waters north of Point Conception, but all three species overlap in 

southern California. Acorn barnacle species can be difficult to distinguish, especially in 

photographic monitoring. Tiny (to 8 mm) C. fissus and C. dalli require dissection and 

microscopic examination of scutal plates. Balanus glandula can be field identified in 

most cases by its larger size (to 22 mm), whiter color, and differing shell plate 

arrangements. Acorn barnacles spawn often, at variable times throughout the year (Hines 

1978), and settle in incredible densities (to 70,000/m2), forming distinct white bands 

along the upper intertidal that contain few other invertebrates except littorines and the 

hardiest limpets. Balanus can out compete Chthamalus by crowding or smothering, but 

Chthamalus can occupy higher tide levels than Balanus, because it is more resistant to 

desiccation. Slightly lower down, acorn barnacles mix in with the Endocladia

assemblage, and are common on mussel shells. Chthamalus species grow rapidly, but 

only survive a few months to a few years. Balanus can live longer (to 10 years), but its 

larger size and lower tidal position subject it to higher levels of mortality from predatory 

gastropods and ochre sea stars. 

Acorn barnacles (particularly Balanus spp.) facilitate recruitment of Endocladia and 

fucoid algae to the upper intertial by reducing grazing pressure by limpets (Farrell 1991). 

We have observed this facilitation at several sites, where barnacle plots become slowly 

inundated by Endocladia, Pelvetiopsis, and Silvetia. 

White acorn barnacles are highly vulnerable to smothering from oil spills because 

floating oil often sticks along the uppermost tidal levels. Significant, widespread barnacle 

impacts were reported after the 1969 Santa Barbara oil platform blow-out (Foster et 

al.(1971) and the 1971 collision of two tankers off San Francisco (Chan 1973). However, 

high recruitment rates may promote relatively rapid recovery of acorn barnacles; 

disturbance recovery times ranging from several months to several years have been 

reported (see Vesco & Gillard 1980). 

One interesting note is that we have found “bent” Chthamalus at several sites, a condition 

previously only seen in the Gulf of California species Chthamalus anisopoma. We have 

been collecting bent individuals in order to identify them to species (requires examination 

of their calcareous plates) and thus far have found them all to be the southern species C. 

fissus.

Pollicipes polymerus 

Goose barnacles, Pollicipes polymerus, are conspicuous in high to middle intertidal zones 

on surf-swept rocky shores all along the US Pacific Coast. Young goose barnacles settle 

preferentially among other Pollicipes, forming tight clusters on exposed outcrops, ridges, 

and walls, just above or intermixed with mussel beds. This distinctive black and white 

barnacle is firmly attached to the rock by a muscular (edible) stalk that holds the cirral 

net up to 8 cm high to filter-feed, primarily from wave backwash. Unlike white acorn 

barnacles, goose barnacles are slow-growing. Sexual maturity is reached in 

B5
approximately 5 years, and large adults may be 20 years old (Morris et al. 1980). 

Pollicipes is very resistant to desiccation and can tolerate all but the highest wave 

exposures. Mortality has been reported from oil spills (Foster et al. 1971; Chan 1973), 

and recovery could be slow. Populations have been reduced in accessible areas where 

goose barnacles are collected for food. 

Mytilus californianus 

The California mussel, Mytilus californianus, is abundant at middle to low levels of 

exposed rocky shores along the entire Pacific Coast. These 10-20 cm black/blue/gray 

mussels firmly attach to rocks or other mussels by tough byssal threads, forming dense 

patches or beds. The literature on Mytilus californianus is extensive, including key 

ecological studies on the effects of predation, grazing, and disturbance on succession and 

community structure (see for discussion Morris et al. 1980; Ricketts et al. 1985; 

Kinnetics 1992). Bay mussels (M. trossulus, or M. galloprovincialis) can co-occur with 

M. californianus, but are most common in sheltered habitats. Thick (≥ 20 cm) beds of 

California mussels trap water, sediment, and detritus that provide food and shelter for an 

incredible diversity of plants and animals, including cryptic forms inhabiting spaces 

between mussels as well as biota attached to mussel shells (Paine 1966; MacGinitie & 

MacGinitie 1968; Suchanek 1979; Kanter 1980, Lohse 1993). For example, MacGinitie 

& MacGinitie (1968) counted 625 mussels and 4,096 other invertebrates in a single 25 

cm2 clump, and Kanter (1980) identified 610 species of animals and 141 species of algae 

from mussel beds at the Channel Islands. Kinnetics (1992) documented locational 

differences in the composition and abundance of mussel bed species. Northern sites had 

densely packed, multi-layered beds, but the more open southern sites had higher species 

diversity. Mussels feed on suspended detritus and plankton. Young mussels settle 

preferentially into existing beds at irregular intervals, grow at variable rates depending on 

environmental conditions, and eventually reach ages of 8 years or more (see Morris et al. 

1980, Ricketts et al. 1985). Mussels can tolerate typical rigors of intertidal life quite 

successfully. However, desiccation likely limits the upper extent of mussel beds, storms

tear out various-sized mussel patches, and sea stars prey especially on lower zone 

mussels. Mytilus are adversely affected by oil spills (Chan 1973; Foster et al. 1971). 

Recovery from disturbance varies from fairly rapid (if clearings are small and surrounded 

by mussels that can move in) to periods greater than 10 years (if clearings are large and 

recruitment is necessary for recolonization)(see Vesco & Gillard 1980; Kinnetics 1992). 

Pisaster ochraceus 

The ochre seastar, Pisaster ochraceus, is found on middle and low tide levels of waveswept rocky coasts from Alaska to Baja California, but is much less common south of 

Point Conception. Its relatively large size (to 45 cm diameter), variety of colors (yellow, 

orange, purple, brown), and ability to withstand air exposure (at least 8 hours) attract 

considerable attention from visitors exploring the shore at low tide. The ochre seastar 

typically is associated with mussels, which constitute its chief food, but barnacles, 

limpets, snails, and chitons also may be taken (Morris et al. 1980). Predator-prey 

B6
interactions involving ochre seastars have been intensely studied, especially the role of P. 

ochraceus in determining the lower limit of northern mussel beds (Paine 1966, 1974: 

Dayton 1971). Like black abalone, ochre sea stars are relatively slow-growing, longlived, and apparently variable in recruitment success. They are tolerant of high surf, using 

their numerous tube feet to remain firmly in place, often in cracks and crevices. They 

have few predators, except for curious tidepool visitors. However, in southern California, 

P. ochraceus populations have been decimated by a widespread wasting disease caused 

by a warm-water bacterium of the genus Vibrio (Schroeter & Dixon pers. comm.). 

Sensitivity to oil spills is not well known; Chan (1973) saw no obvious effects from a San 

Francisco oil spill. Recovery time from any major population loss likely would be very 

long. 

Lottia gigantea 

The owl limpet, Lottia gigantea, is common in high and middle tide zones of exposed 

rocky shores from Washington south to Baja California. Adult Lottia are relatively easy 

to identify because of their large size (up to >100 mm), oval shape with low rounded 

profile, and color patterns of brown, white, and black on the often eroded shell. 

Accessory gills on the mantle increase surface area for aerial respiration during low tide 

periods. Owl limpet habitats extend from the barnacle and Endocladia zones down to the 

mussel beds. 

Lottia can either be territorial, maintaining and defending clearings of thick algal film, or 

nonterritorial, intruding on other limpets’ algal farms to graze (Shanks 2002). Territorial 

behavior is determined by previous agnostic experience, although large limpets tend to be 

territorial and smaller ones are more often intruders (Shanks 2002). Territorial 

individuals maintain feeding territories on relatively smooth rock surfaces which they 

keep free (by rasping and bulldozing) of most macroalgae and invertebrates, including 

turfweed, sea anemones, barnacles, mussels, and other limpets (Stimpson 1970). By 

removing most competitors for space and grazers, they promote the growth of algal films 

upon which they systematically graze. These “clearings” vary in appearance with Lottia

size and structural features of the substrate, creating a patchwork of differing 

microhabitats. Lottia tend to occupy one or more characteristic “home scars” within their 

territories. Here the shell margin conforms to the rock surface, making a tight seal to hold 

moisture during low tides. The limpets also may tuck into crevices and under mussels for 

protection from heat, desiccation, and high surf. 

Lottia grow slowly, taking up to 10-15 years to reach maximum size (Morris et al. 1980). 

As an ecological dominant, any change in Lottia populations greatly affects abundances 

of other species. Lindberg et al. (1998) have shown that if Lottia are removed from an 

area, cover of erect, fleshy algae increases, followed by increases in the number of small 

limpets. 

The limpets and their feeding territories are vulnerable to oiling, but oil impacts are 

unclear. For example, they were not obviously affected by the 1971 San Francisco oil 

spill (Chan 1973). Recovery from any major disturbance likely would be lengthy. Larger 

owl limpets are collected for food, tasting much like abalone. Mean Lottia shell length 

has been negatively correlated with collecting pressure in southern California (Kido & 

B7
Murray 2003; Roy et al. 2003). Because Lottia are protandrous hermaphrodites, the 

largest individuals are nearly always female (Ricketts et al. 1985), and collecting may 

impair reproductive capabilities within owl limpet populations. 

Haliotis cracherodii 

Black abalone (Haliotis cracherodii) inhabit mid-low intertidal levels down to shallow 

subtidal depths (to 6 m) from Oregon to Southern Baja California (Morris et al. 1980). 

They are readily identified by dark, bluish-black coloration, a smooth shell with 5-7 open 

respiratory holes, and relatively small size (5-20 cm as adults). Black abalone are 

relatively sedentary, and are typically found clustered in wet crevices, under boulders, or 

on the walls of surge channels along exposed shores. Juveniles graze on diatom films and 

coralline algae, while adults primarily eat drift algae, especially brown kelps. H. 

cracherodii compete with sea urchins and other crevice-dwellers for space and food. 

Before recent catastrophic declines (see below), abalone could occasionally be seen 

stacked on top of each other, reaching densities of more than 100/m2 (Douros 1987; 

Richards & Davis 1993). Black abalone are slow-growing and long-lived, with 

recruitment apparently being low and variable (Morris et al. 1980; VanBlaricom 1993). 

Growth rates depend on animal size, location, food availability, reproductive condition, 

and other factors. Absolute longevity has not been determined, but ages greater than 30 

years appear likely based on tagging and other population studies (e.g., VanBlaricom

1993). 

Although once an important human resource, both sport and commercial black abalone 

fisheries have been closed due to recent precarious declines. Mortality is associated with 

“withering syndrome” (WS), in which the foot shrinks and weakened individuals lose 

their grip on rock surfaces. H. cracherodii populations in Southern California suffered 

catastrophic declines in the mid-1980’s that resulted in near-complete disappearance of 

black abalone along mainland shores south of Point Conception, as well as on the 

Channel Islands (Lafferty & Kuris 1993; Richards & Davis 1993). In 1993 WS caused 

massive declines in mainland populations near Point Conception. Since then, WS has 

slowly spread up the coast, decimating some of the largest remaining black abalone 

populations (Altstatt et al. 1996; Raimondi et al. 2002). 

Other sources of mortality include: smothering by sand burial, dislodgment by storm

waves, and predation by octopus, sea stars, fishes, and sea otters (Morris et al. 1980; 

VanBlaricom 1993). Impacts from oil are little known, but North et al. (1965) reported 

black abalone mortality following a spill in Baja California. Because of low recruitment, 

slow growth, and already decimated reproductive populations, additional mortality from

oil spills would be devastating, and recovery prospects long-term at best. 

B8
LITERATURE CITED 

Abbott, I.A. and G.J. Hollenberg. 1976. Marine algae of California. Stanford University 

Press, Stanford, California. 827 pp. 

Altstatt, J.M., R.F. Ambrose, J.M. Engle, P.L. Haaker, K.D. Lafferty, and P.T. Raimondi. 

1996. Recent declines of black abalone Haliotis cracherodii on the mainland coast of 

central California. Mar. Ecol. Prog. Ser. 142:185-192. 

Carrington, E. 1990. Drag and dislodgment of an intertidal macroalga: consequences of 

morphological variation in Mastocarpus papillatus Kutzing. J. Exp. Mar. Biol. Ecol. 

139: 185-200. 

Chan, G.L. 1973. A study of the effects of the San Francisco oil spill on marine 

organisms. Pp. 741-782 in Proceedings of joint conference on prevention and control 

of oil spills. American Petroleum Institute, Washington, D.C. 

Dawson, E.Y. and M.S. Foster. 1982. Seashore plants of California. University of 

California Press, Los Angeles, California. 226 pp. 

Dayton, P.K. 1971. Competition, disturbance and community organization: the provision 

and subsequent utilization of space in a rocky intertidal community. Ecol. Monogr. 

41:351-389. 

Douros, W.J. 1987. Stacking behavior of an intertidal abalone: an adaptive response or a 

consequence of space limitation? J. Exp. Mar. Biol. Ecol. 108:1-14. 

Dyck, L.J. and R.E. De Wreede. 1995. Patterns of seasonal demographic change in the 

alternate isomorphic stages of Mazzaella splendens (Gigartinales, Rhodophyta). 

Phycologia 34: 390-395. 

Engle, J.M. 1979. Ecology and growth of juvenile California spiny lobster, Panulirus 

interruptus (Randall). Ph.D. Dissertation, University of Southern California. 298 pp. 

Farrell, T.M. 1991. Models and mechanisms of succession: an example from a rocky 

intertidal community. Ecological Monographs 61(1): 95-113 

Foster, M., M. Neushul, and R. Zingmark. 1971. The Santa Barbara oil spill. Part 2. 

Initial effects on intertidal and kelp bed organisms. Environ. Pollut. 2:115-134. 

Foster, M.S., A.P. DeVogelaere, C. Harrold. J.S. Pearse, and A.B. Thum. 1988. Causes of 

spatial and temporal patterns in rocky intertidal communities of Central and 

Northern California. Memoirs of the California Academy of Sciences Number 9, San 

Francisco, California. 45 pp. 

Gaines, S.D. 1985. Herbivory and between-habitat diversity: The differential 

effectiveness of defenses in a marine plant. Ecology 66: 473-485. 

B9
Glynn, P.W. 1965. Community composition, structure, and interrelationships in the 

marine intertidal Endocladia muricata - Balanus glandula association in Monterey 

Bay, California. Beaufortia 12:1-198. 

Gomez, I.M. and R.C. Westermeier. 1991. Frond regrowth from basal disc in Iridaea 

laminarioides (Rhodophyta, Gigartinales) at Mehuin, southern Chile. Mar. Ecol. 

Prog. Ser. 73: 83-91. 

Gunnill, F.C. 1980. Demography of the intertidal brown alga Pelvetia fastigiata in 

southern California, USA. Mar. Biol. 59:169-179. 

Gunnill, F.C. 1983. Seasonal variations in the invertebrate faunas of Pelvetia fastigiata

(Fucaceae): effects of plant size and distribution. Mar. Biol. 73:115-130. 

Hill, M.L. 1980. Structure, organization, and persistence of the Pelvetia fastigiata

(Phaeophyceae: Fucales) community on a rocky intertidal shoreline at Dana Point, 

Orange County, California. Master of Arts Thesis, California State University, 

Fullerton, California, 116 pp. 

Hines, A.H. 1978. Reproduction in three species of intertidal barnacles from central 

California. Biol. Bull. 154:262-281. 

Holbrook, S.J., D.C. Reed, K. Hansen, C.A. Blanchette. 2000. Spatial and temporal 

patterns of predation on seeds of the surfgrass Phyllospadix torreyi. Marine Biology 

136:739-747. 

Johnson, L.E., S.H. Brawley. 1998. Dispersal and recruitment of a canopy-forming 

intertidal alga: the relative roles of propagule availability and post-settlement 

processes. Oecologia. 117:517-526. 

Kanter, R.G. 1980. Biogeographic patterns in mussel community distribution from the 

southern California Bight. Pp. 341-355 in The California Islands: proceedings of a 

multidisciplinary symposium. D.M. Power, ed. Santa Barbara Museum of Natural 

History, Santa Barbara, California. 

Kido, J.S., S.N. Murray. 2003. Variation in owl limpet Lottia gigantea population 

structures, growth rates, and gonadal production on southern California rocky shores. 

Mar. Ecol. Prog. Ser. 257:111-124. 

Kinnetics Laboratories, Inc. 1992. Study of the rocky intertidal communities of Central 

and Northern California. Report to the Minerals Management Service. OCS Study 

MMS 91-0089. 

Lafferty, K.D.and A.M. Kuris. 1993. Mass mortality of abalone Haliotis cracherodii on 

the California Channel Islands: tests of epidemiological hypotheses. Mar. Ecol. 

Prog. Ser. 96: 239-248. 

B10 
Lindberg, D. R. , J. A. Estes and K. I. Warheit. 1998. Human influences on trophic 

cascades along rocky shores. Ecol. Appl. 8:880-890 

Littler, M.M. and S.N Murray. 1975. Impact of sewage on the distribution, abundance, and 

community structure of rocky intertidal macro-organisms. Mar. Biol. 30: 277-91. 

Lohse, D.P. 1993. The importance of secondary substratum in a rocky intertidal community. 

J. Exp. Mar. Biol. Ecol. 166:1-17 

MacGinitie, G.E. and N. MacGinitie. 1968. Natural history of marine animals. 2nd ed. 

McGraw Hill, New York. 523 pp. 

Mondragon, J and J. Mondragon. 2003. Seaweeds of the Pacific Coast. Common Marine 

Algae from Alsaka to Baja California. Sea Challengers, Monterey, CA. 97pp. 

Morris, R.H., D.P. Abbott, and E.C. Haderlie, eds. 1980. Intertidal invertebrates of 

California. Stanford University Press, Stanford, California. 690 pp. 

Murray, S.N. and T.G. Denis. 1997. Vulnerability of the rockweed Pelvetia compressa to 

anthropogenic disturbance on southern California rocky shores. Phycologia 36:75-

76. 

North, W.J., M. Neushul and K.A. Clendenning. 1965. Successive biological changes 

observed in a marine cover exposed to a large spillage of mineral oil. Pp. 335-354 in 

Proceedings of the Comm. Int. Explor. Scient. Mer. Merit. Symposium on marine 

pollution caused by micro-organisms and mineral oils, Monaco. 

Pacific Gas and Electric Company 1993. Thermal effects monitoring program. Diablo 

Canyon Power Plant 1992 Annual Report. Pacific Gas and Electric Co., San 

Francisco, California. 

Paine, R.T. 1966. Food web complexity and species diversity. Am. Nat. 100:65-75. 

Paine, R.T. 1974. Intertidal community structure: experimental studies on the relationship 

between a dominant competitor and its principal predator. Oecologia 15:93-120. 

Pearse, V., and L. Francis 2000. Anthopleura sola, a new species, solitary sibling species 

to the aggregating sea anemone, A. elegantissima (Cnidaria: Anthozoa: Actiniaria: 

Actiniidae). Proceedings of the Biological Society of Washington. 113(3):596-608 

Raimondi, P.T., C.M. Wilson, R.F. Ambrose, J.M. Engle, and T.E. Minchinton. 2002. 

Continued declines of black abalone along the coast of California: are mass 

mortalities related to El Nino events? Mar. Ecol. Prog. Ser. 242:143-152. 

Richards, D.V. and G.E. Davis. 1993. Early warnings of modern population collapse in 

black abalone Haliotis cracherodii, Leach, 1814 at the California Channel Islands. J. 

Shellfish Res. 12(2):189-194. 

B11 
Ricketts, E.G., J. Calvin, J. Hedgepeth, and D.W. Phillips. 1985. Between Pacific tides, 

5th ed., revised by J. Hedgepeth. Stanford University Press, Palo Alto, California. 

652 pp. 

Roy, K., A.G. Collins, B.J. Becker, E. Begovic, and J.M. Engle. 2003. Anthropogenic 

impacts and historical decline in body size of rocky intertidal gastropods in southern 

California. Ecology Letters 6:205-211. 

Sapper, S.A. and S.N. Murray 2003. Variation in structure of the subcanopy assemblage 

associated with southern California populations of the intertidal rockweed Silvetia 

compressa (Fucales). Pacific Science 57(4):433-462. 

Seapy, R.R., Littler, M.M. 1993. Rocky intertidal macroinvertebrates of the Southern

California Bight: an overview and checklist. In: Hockberg, F.G. (ed.), Third California 

Islands Symposium: recent advances in research on the California islands. Santa 

Barbara Museum of Natural History, Santa Barbara, California, pp. 293-322. 

Sebens, K.P. 1980. The regulation of asexual reproduction and indeterminate body size in 

the sea anemone Anthopleura elegantissima (Brandt). Biol. Bull. 158:370-382. 

Sebens, K.P. 1982. Asexual reproduction in Anthopleura elegantissima (Anthozoa: 

Actiniaria): seasonality and spatial extent of clones. Ecology 63:434-444. 

Shanks, A.L. 2002. Previous agnostic experience determines both foraging behavior and 

territoriality in the limpet Lottia gigantea (Sowerby). Behavioral Ecology 13(4):467-

471. 

Silva, P.C., F.F. Pedroche, M. E. Chacana, R.Aguilar-Rosas, L.E. Aguilar-Rosas, and J. 

Raum. Geographic correlation of morphological and molecular variation in Silvetia 

compressa (Fucaceae, Fucales, Phaeophyceae). Phycologia 43(2):204-214. 

Stewart, J.G. and B. Myers. 1980. Assemblages of algae and invertebrates in Southern 

California Phyllospadix-dominated intertidal habitats. Aquatic Botany 9:73-94. 

Stimpson, J. 1970. Territorial behavior of the owl limpet Lottia gigantea. Ecology 

51:113-118. 

Suchanek, T.H. 1979. The Mytilus californianus community: studies on composition, 

structure, organization, and dynamics of a mussel bed. Ph.D. Thesis, University of 

Washington, Seattle, Washington. 

Taylor, P.R. and M.M. Littler 1982. The roles of compensatory mortality, physical 

disturbance, and retention substrate in the development and organization of a sandinfluenced rocky-intertidal community. Ecology 63:135-146. 

Turner, T. 1983. Complexity of early and middle successional stages in a rocky intertidal 

surfgrass community. Oecologia 60:56-65. 

B12 
Turner, T. 1985. Stability of rocky intertidal surfgrass beds: persistence, preemption, and 

recovery. Ecology 66:83-92. 

VanBlaricom, G.R. 1993. Dynamics and distribution of black abalone populations at San 

Nicolas Island, California. In: Third California Islands Symposium: recent advances 

in research on the California Islands (F.G. Hochberg, ed.), Santa Barbara Museum of 

Natural History, Santa Barbara, pp. 323-334. 

Vesco, L.L. and R. Gillard. 1980. Recovery of benthic marine populations along the 

Pacific Coast of the United States following man-made and natural disturbances 

including pertinent life history information. U.S. Department of the Interior, Bureau 

of Land Management Service, POCS Reference Paper No. 53-4. 48 pp. 

Zupan, J.R. and J.A. West. 1988. Geographic variation in the life history of 

Mastocarpus papillatus (Rhodophyta). J. Phycol. 24: 223-229. 

B13 
 C-1 

Appendix C: Means and Standard Errors for All Target Species 

This appendix contains means and standard errors for all key species and for species that are 

abundant in target species plots. Mean percent cover or mean number across all plots is given for 

all species except black abalone. Total number for all plots combined is given for abalone. 

Abbreviations for sites used in the tables are as follows: 

PSN Pt. Sierra Nevada 

PBL Piedras Blancas 

RM Rancho Marino 

CAY Cayucos 

HAZ Hazards 

SHB Shell Beach 

OCC Occulto 

PUR Purisima Pt. 

STA Stairs 

BOA Boat House 

GPT Government Pt. 

ALEG Alegria 

ARHO Arroyo Hondo 

COPT Coal Oil Pt. 

CARP Carpinteria 

MUSH Mussel Shoals 

OLDS Old Stairs 

PCOV Paradise Cove 

WHPT White’s Pt. 

PTFM Pt. Fermin 

SHCO Shaw’s Cove 

CRCO Crystal Cove 

TRIS Treasure Island 

DAPT Dana Pt. 

CTBR Catalina Island, Bird Rock 

CTLH Catalina Island, Little Harbor 
 C-2 

SILVETIA 

 Silvetia Rock Silvetia Rock 

Sample Site mean se mean se Sample Site mean se mean se 

FA95 PSN 93.8 1.5 2.8 0.7 FA99 HAZ 79.4 5.3 3.2 0.7 

SP96 PSN 88.0 1.9 5.0 1.8 SP00 HAZ 72.8 5.8 2.4 0.9 

FA96 PSN 88.2 4.6 6.6 1.7 FA00 HAZ 72.2 4.5 2.6 0.7 

SP97 PSN 85.8 4.2 3.8 1.0 SP01 HAZ 65.2 7.4 3.6 1.1 

FA97 PSN 87.0 4.1 4.6 1.8 FA01 HAZ 67.8 4.2 5.2 1.1 

SP98 PSN 74.2 6.0 10.8 3.6 SP02 HAZ 66.6 6.2 3.2 0.9 

FA98 PSN 83.2 6.9 8.6 4.6 FA02 HAZ 69.0 5.0 6.6 2.6 

SP99 PSN 81.0 6.4 8.0 3.1 SP03 HAZ 56.4 7.7 6.2 2.1 

FA99 PSN 81.8 6.6 8.6 4.0 FA03 HAZ 61.4 11.5 9.6 4.3 

SP00 PSN 72.6 8.9 12.2 5.1 

FA00 PSN 61.8 14.1 15.0 6.9 FA95 SHB 93.0 2.2 1.6 0.5 

SP01 PSN 62.4 11.7 18.4 7.5 SP96 SHB 90.0 3.7 2.2 1.1 

FA01 PSN 51.4 12.7 23.8 7.8 FA96 SHB 88.6 1.9 2.2 0.4 

SP02 PSN 51.0 11.8 20.6 6.9 SP97 SHB 80.6 2.4 6.0 2.1 

FA02 PSN 57.4 12.8 14.6 5.1 FA97 SHB 85.8 3.4 5.2 2.5 

SP03 PSN 47.8 12.9 22.4 6.8 SP98 SHB 76.4 6.0 5.6 3.0 

FA03 PSN 59.0 9.2 20.6 4.5 FA98 SHB 76.2 7.4 9.0 2.3 

 SP99 SHB 74.6 9.7 7.0 1.9 

FA95 CAY 98.0 1.3 1.0 1.0 FA99 SHB 71.0 6.3 10.8 3.3 

SP96 CAY 76.6 4.6 7.4 2.3 SP00 SHB 60.4 11.0 12.4 5.2 

FA96 CAY 98.4 0.5 0.6 0.2 FA00 SHB 55.8 13.1 19.8 8.3 

SP97 CAY 75.2 2.2 11.2 2.4 SP01 SHB 58.0 12.9 17.0 6.8 

FA97 CAY 80.0 4.5 8.6 2.1 FA01 SHB 49.6 11.8 17.8 6.5 

SP98 CAY 66.8 6.1 12.2 2.3 SP02 SHB 49.0 11.8 18.4 5.0 

FA98 CAY 80.2 11.1 8.8 5.6 FA02 SHB 52.8 10.6 16.8 5.7 

SP99 CAY 71.6 8.9 11.2 3.5 SP03 SHB 46.0 9.6 18.2 3.9 

FA99 CAY 88.8 4.8 4.2 1.3 FA03 SHB 45.4 10.4 23.8 3.5 

SP00 CAY 69.0 5.8 15.2 4.2 

FA00 CAY 79.4 11.6 7.2 3.3 SP92 STA 86.8 3.1 5.6 0.7 

SP01 CAY 51.0 13.2 20.2 5.3 FA92 STA 79.0 8.7 15.2 6.9 

FA01 CAY 50.6 13.3 23.2 6.8 SP93 STA 70.6 8.4 16.8 6.3 

SP02 CAY 48.8 4.2 24.2 6.0 FA93 STA 92.2 2.7 4.6 2.3 

FA02 CAY 46.6 8.8 29.4 8.1 SP94 STA 76.6 6.7 10.6 6.0 

SP03 CAY 29.2 9.3 37.0 10.0 FA94 STA 86.8 1.7 2.8 1.2 

FA03 CAY 50.6 9.8 26.2 9.5 SP95 STA 68.8 9.4 9.4 3.7 

 FA95 STA 64.8 5.8 17.2 7.3 

FA95 HAZ 89.6 2.2 3.8 0.4 SP96 STA 52.6 7.9 20.4 7.6 

SP96 HAZ 78.6 1.5 4.8 1.0 FA96 STA 57.2 11.8 14.0 6.9 

FA96 HAZ 89.2 2.4 1.4 0.7 SP97 STA 55.2 11.2 17.8 6.9 

SP97 HAZ 88.8 4.7 2.0 0.8 FA97 STA 57.4 9.8 19.6 7.4 

FA97 HAZ 89.6 1.9 2.2 0.6 SP98 STA 34.2 7.8 29.8 10.1 

SP98 HAZ 73.0 4.9 4.6 1.8 FA98 STA 34.2 9.5 29.0 7.2 

FA98 HAZ 81.0 3.9 3.2 2.0 SP99 STA 26.2 8.1 29.8 8.6 

SP99 HAZ 73.2 7.5 3.8 1.9 FA99 STA 31.0 7.0 34.6 10.5 
 C-3 

 Silvetia Rock Silvetia Rock 

Sample Site mean se mean se Sample Site mean se mean se 

SP00 STA 21.0 7.1 32.4 7.7 FA97 GPT 80.0 3.1 12.4 2.6 

FA00 STA 25.2 12.1 34.2 10.6 SP98 GPT 60.4 11.2 28.0 8.8 

SP01 STA 17.6 8.4 30.0 8.5 FA98 GPT 65.4 9.7 22.0 7.3 

FA01 STA 33.6 9.0 28.4 6.7 SP99 GPT 64.2 11.2 22.4 8.6 

SP02 STA 21.0 8.1 33.8 6.2 FA99 GPT . . . . 

FA02 STA 34.0 10.1 25.2 6.5 SP00 GPT 67.6 13.3 15.8 7.2 

SP03 STA 19.8 7.9 29.8 2.9 FA00 GPT 70.6 9.8 19.6 7.9 

FA03 STA 29.6 11.0 21.6 2.2 SP01 GPT 63.2 11.5 21.2 7.1 

 FA01 GPT 64.6 8.1 22.2 6.5 

SP92 BOA 84.8 5.3 3.8 0.9 SP02 GPT 52.4 9.7 30.0 5.4 

FA92 BOA 91.6 4.0 2.8 1.2 FA02 GPT 66.6 7.7 16.8 4.3 

SP93 BOA 89.8 2.7 3.8 2.0 SP03 GPT 47.4 9.9 30.8 5.8 

FA93 BOA 88.2 5.1 5.0 1.9 FA03 GPT 59.4 7.9 20.8 2.6 

SP94 BOA 77.8 8.4 4.4 1.7 

FA94 BOA 87.0 7.8 2.4 1.3 FA99 PTFM 95.0 1.5 4.0 1.4 

SP95 BOA 71.0 5.0 8.6 2.7 SP00 PTFM 90.6 2.1 6.6 1.1 

FA95 BOA . . . . FA00 PTFM 97.8 2.0 2.0 2.0 

SP96 BOA 70.2 8.6 3.4 0.7 SP01 PTFM 88.8 3.4 11.2 3.4 

FA96 BOA 78.2 12.9 5.4 1.5 FA01 PTFM 98.4 0.9 1.2 1.0 

SP97 BOA 71.6 12.9 7.2 1.2 SP02 PTFM 92.4 2.2 7.0 2.3 

FA97 BOA 76.6 12.0 4.8 2.0 FA02 PTFM 100.0 0.0 0.0 0.0 

SP98 BOA 67.6 9.7 8.2 2.0 SP03 PTFM 96.4 1.4 3.4 1.2 

FA98 BOA 70.4 11.0 6.4 3.5 FA03 PTFM 99.4 0.4 0.4 0.2 

SP99 BOA 74.0 6.6 6.4 1.9 

FA99 BOA 73.2 9.0 8.4 3.2 FA96 CRCO 95.0 1.1 1.8 0.6 

SP00 BOA 76.0 6.5 3.0 2.0 SP97 CRCO 86.6 3.0 5.0 2.5 

FA00 BOA 91.8 3.6 1.0 0.8 FA97 CRCO 70.6 3.0 1.6 1.0 

SP01 BOA 76.8 4.3 5.4 1.6 SP98 CRCO 77.0 4.5 7.4 3.4 

FA01 BOA 92.6 6.2 1.0 0.6 FA98 CRCO 95.0 2.3 2.6 1.7 

SP02 BOA 90.4 5.4 1.4 0.5 SP99 CRCO 79.2 4.7 12.4 3.2 

FA02 BOA 91.2 4.1 1.4 1.0 FA99 CRCO 89.8 3.0 5.0 1.6 

SP03 BOA 84.0 6.7 3.2 1.6 SP00 CRCO 74.6 2.3 10.8 2.5 

FA03 BOA 93.2 4.7 0.8 0.5 FA00 CRCO 90.8 2.9 3.0 1.5 

 SP01 CRCO 84.6 5.1 7.2 3.9 

SP92 GPT 74.6 4.0 16.2 2.5 FA01 CRCO 92.8 2.2 3.0 1.0 

FA92 GPT 75.4 8.0 10.2 4.0 SP02 CRCO 67.4 4.1 8.4 1.7 

SP93 GPT 75.6 3.5 9.0 4.5 FA02 CRCO 88.6 1.5 2.6 1.6 

FA93 GPT 81.8 8.0 3.8 1.3 SP03 CRCO 87.4 2.2 3.0 0.6 

SP94 GPT 60.0 10.8 16.0 3.3 FA03 CRCO 94.0 1.8 1.6 0.4 

FA94 GPT 81.2 6.6 2.4 1.5 

SP95 GPT 63.4 7.5 18.2 4.7 FA96 SHCO 87.8 1.1 2.6 1.1 

FA95 GPT 71.8 5.6 13.8 2.9 SP97 SHCO 72.4 4.5 6.6 4.0 

SP96 GPT 68.8 4.9 18.4 3.9 FA97 SHCO 74.6 2.4 0.6 0.4 

FA96 GPT 80.4 4.7 10.0 4.8 SP98 SHCO 71.2 6.2 3.0 1.1 

SP97 GPT 64.8 7.0 22.8 5.1 FA98 SHCO 74.6 8.6 10.8 3.9 
 C-4 

 Silvetia Rock Silvetia Rock 

Sample Site mean se mean se Sample Site mean se mean se 

SP99 SHCO 65.8 9.3 1.4 1.2 FA94 CTBR 94.8 0.6 0.6 0.4 

FA99 SHCO 83.4 7.2 7.2 3.6 SP95 CTBR 91.4 1.8 0.8 0.6 

SP00 SHCO 76.8 7.9 13.0 6.8 FA95 CTBR 97.6 1.0 2.0 0.7 

FA00 SHCO 86.8 4.5 4.2 2.5 SP96 CTBR 89.8 3.1 1.2 0.7 

SP01 SHCO 89.4 0.9 3.4 1.2 FA96 CTBR 92.2 3.5 0.8 0.2 

FA01 SHCO 88.6 3.5 2.0 1.0 SP97 CTBR 84.4 5.2 2.6 1.9 

SP02 SHCO 72.8 5.6 7.0 2.7 FA97 CTBR 79.6 4.6 2.0 0.9 

FA02 SHCO 85.8 4.9 5.8 3.1 SP98 CTBR 79.0 5.4 1.0 0.5 

SP03 SHCO 82.6 2.1 4.8 2.2 FA98 CTBR 66.4 17.4 3.0 1.6 

FA03 SHCO 97.0 1.3 0.8 0.4 SP99 CTBR 67.6 13.9 0.0 0.0 

 FA99 CTBR 75.2 13.3 8.0 5.9 

FA96 TRIS 84.4 2.1 1.2 0.6 SP00 CTBR 72.2 12.6 3.6 2.5 

SP97 TRIS 53.8 7.4 10.8 4.3 FA00 CTBR 78.6 10.5 5.6 3.5 

FA97 TRIS 69.6 5.3 5.0 2.3 SP01 CTBR 79.4 9.6 2.6 1.5 

SP98 TRIS 63.8 4.7 14.8 6.9 FA01 CTBR 77.6 9.3 3.8 2.1 

FA98 TRIS 70.4 5.2 20.8 6.8 SP02 CTBR 75.8 10.1 0.6 0.6 

SP99 TRIS 62.2 8.7 22.0 8.8 FA02 CTBR 89.8 7.0 1.0 0.8 

FA99 TRIS 87.0 4.7 1.0 0.6 SP03 CTBR 76.6 8.5 1.8 0.6 

SP00 TRIS 62.2 2.7 14.0 2.0 

FA00 TRIS 85.4 5.0 7.4 3.8 FA94 CTLH 64.8 2.3 22.6 2.5 

SP01 TRIS 64.2 8.3 14.0 5.7 SP95 CTLH 67.2 3.8 14.8 2.1 

FA01 TRIS 69.6 6.0 7.0 1.9 FA95 CTLH 83.0 1.5 9.4 2.6 

SP02 TRIS 59.2 7.1 10.2 2.3 SP96 CTLH 82.8 1.2 8.8 2.7 

FA02 TRIS 69.2 6.5 8.8 3.1 FA96 CTLH 91.0 1.0 7.2 0.9 

SP03 TRIS 69.4 5.9 7.4 1.4 SP97 CTLH 86.8 1.6 9.2 2.3 

FA03 TRIS 92.4 1.7 3.2 1.4 FA97 CTLH 80.2 6.4 3.4 1.8 

 SP98 CTLH 64.8 11.2 9.8 3.8 

FA96 DAPT 92.6 1.5 2.8 1.2 FA98 CTLH 72.8 11.4 7.8 4.5 

SP97 DAPT 84.8 0.8 2.0 0.8 SP99 CTLH 67.8 10.2 1.8 1.2 

FA97 DAPT 84.2 3.8 1.0 0.6 FA99 CTLH 85.6 5.7 4.0 2.8 

SP98 DAPT 62.8 7.8 7.6 2.8 SP00 CTLH 86.4 3.1 6.8 2.5 

FA98 DAPT 85.8 3.9 6.8 2.5 FA00 CTLH 88.0 5.4 6.4 3.8 

SP99 DAPT 83.8 1.8 9.6 1.2 SP01 CTLH 92.6 2.7 3.4 1.5 

FA99 DAPT 87.6 2.6 1.6 1.1 FA01 CTLH 95.4 1.6 3.0 1.3 

SP00 DAPT 82.8 2.7 10.2 3.1 SP02 CTLH 90.4 1.9 4.0 2.1 

FA00 DAPT 93.4 0.8 3.6 1.1 FA02 CTLH 95.6 2.4 1.8 0.9 

SP01 DAPT 83.0 2.2 8.0 3.1 SP03 CTLH 85.6 3.9 10.4 2.5 

FA01 DAPT 94.8 1.8 1.4 0.9 

SP02 DAPT 81.6 3.2 9.0 1.9 

FA02 DAPT 94.0 1.5 2.8 1.0 

SP03 DAPT 83.0 1.4 5.8 1.4 

FA03 DAPT 94.4 1.9 3.2 1.7 
 C-5 

Hesperophycus 

 Hesperophycus Endocladia Barnacles Rock 

Sample Site mean se mean se mean se mean se 

FA95 PSN 89.4 2.0 3.6 1.4 2.0 1.0 5.0 1.7 

SP96 PSN 89.0 2.3 3.4 1.2 1.2 1.0 3.2 0.7 

FA96 PSN 85.2 3.2 1.8 0.7 3.8 3.1 8.4 1.5 

SP97 PSN 71.6 10.3 5.0 2.9 7.2 5.5 15.2 5.3 

FA97 PSN 58.8 12.1 2.8 0.7 2.8 2.6 30.8 9.0 

SP98 PSN 44.2 11.5 6.2 1.3 3.6 1.2 39.0 11.4 

FA98 PSN 32.8 8.4 4.2 1.2 1.2 0.5 49.2 10.6 

SP99 PSN 32.8 7.8 4.0 1.6 2.8 0.4 46.0 9.8 

FA99 PSN 16.8 7.7 6.2 2.0 4.8 2.4 59.4 8.8 

SP00 PSN 18.4 7.5 8.0 2.4 2.6 1.2 57.6 7.6 

FA00 PSN 14.2 9.4 9.2 3.3 5.2 2.6 67.4 9.8 

SP01 PSN 14.8 9.5 9.2 1.9 5.4 1.0 64.6 11.7 

FA01 PSN 10.2 6.5 11.2 3.8 3.2 1.5 71.4 7.0 

SP02 PSN 11.4 6.4 19.8 5.0 1.4 0.7 65.0 7.1 

FA02 PSN 10.6 5.0 15.8 4.6 8.0 2.8 59.6 8.0 

SP03 PSN 12.6 5.8 24.6 7.2 2.2 0.6 55.6 7.4 

FA03 PSN 23.0 11.2 14.4 5.1 1.0 0.3 56.2 8.4 

 

FA95 CAY 88.2 3.3 0.8 0.6 1.6 1.0 8.6 2.0 

SP96 CAY 62.0 2.8 6.6 1.6 10.4 3.3 20.8 3.2 

FA96 CAY 39.4 8.8 5.2 0.7 9.2 3.8 42.8 6.0 

SP97 CAY 17.8 5.6 7.2 1.2 24.8 6.3 48.0 1.8 

FA97 CAY 15.4 4.4 9.4 3.4 17.4 5.4 54.0 4.1 

SP98 CAY 17.4 5.7 16.2 8.7 9.8 2.5 51.6 9.8 

FA98 CAY 21.8 6.8 4.4 1.3 9.0 1.4 62.0 6.9 

SP99 CAY 14.0 4.5 7.4 2.6 7.8 3.1 69.4 6.8 

FA99 CAY 18.4 4.9 7.0 1.7 5.2 2.6 60.8 9.1 

SP00 CAY 25.6 6.9 8.8 2.7 2.4 1.9 61.0 9.2 

FA00 CAY 27.6 9.9 7.6 1.8 3.8 2.2 58.2 10.6 

SP01 CAY 33.8 13.4 5.4 2.6 4.0 2.5 55.4 13.5 

FA01 CAY 33.4 13.8 4.6 2.3 4.0 2.2 57.6 12.7 

SP02 CAY 29.0 12.3 7.4 1.9 3.0 1.8 58.8 11.9 

FA02 CAY 21.2 9.9 7.0 2.3 6.2 2.1 60.0 10.0 

SP03 CAY 25.8 11.3 9.0 3.3 3.4 1.9 60.0 11.9 

FA03 CAY 20.6 7.9 8.8 3.8 3.6 1.7 62.6 10.3 
 C-6 

Endocladia 

 Endocladia Silvetia Barnacles Rock 

Sample Site mean se mean se mean se mean se 

FA95 CAY 59.6 2.5 0.0 0.0 3.2 1.0 23.0 1.8 

SP96 CAY 65.0 8.1 0.0 0.0 1.2 0.7 26.2 6.1 

FA96 CAY 52.0 5.4 0.0 0.0 3.4 1.8 28.4 8.0 

SP97 CAY 61.0 8.2 0.0 0.0 4.0 1.7 21.4 5.8 

FA97 CAY 39.2 13.4 0.0 0.0 3.4 1.6 51.2 14.0 

SP98 CAY 54.8 9.9 0.0 0.0 0.0 0.0 32.4 8.9 

FA98 CAY 13.2 2.8 0.0 0.0 0.2 0.2 77.0 3.6 

SP99 CAY 21.0 2.7 0.0 0.0 3.4 2.9 64.2 5.5 

FA99 CAY 18.6 3.0 2.2 1.0 2.2 2.2 66.2 6.1 

SP00 CAY 23.8 4.7 0.0 0.0 0.6 0.6 68.0 6.6 

FA00 CAY 36.6 6.5 0.0 0.0 0.6 0.4 55.0 7.9 

SP01 CAY 37.0 8.4 0.0 0.0 0.8 0.4 52.4 10.0 

FA01 CAY 25.6 4.3 0.0 0.0 0.0 0.0 62.6 7.3 

SP02 CAY 42.4 6.6 0.0 0.0 0.0 0.0 44.4 9.6 

FA02 CAY 31.2 7.6 0.0 0.0 0.4 0.2 53.0 7.7 

SP03 CAY 45.8 9.5 0.0 0.0 0.8 0.4 42.4 10.1 

FA03 CAY 33.4 11.1 0.0 0.0 1.0 0.6 49.8 11.8 

 

FA95 HAZ 53.4 5.5 3.2 1.3 13.6 4.2 19.4 3.9 

SP96 HAZ 68.2 5.9 2.6 1.9 6.6 3.5 15.4 3.4 

FA96 HAZ 50.6 3.1 0.2 0.2 11.0 5.4 31.8 6.3 

SP97 HAZ 62.8 6.0 3.6 1.6 3.8 1.5 19.0 3.6 

FA97 HAZ 33.6 4.8 0.2 0.2 10.4 5.0 45.8 7.7 

SP98 HAZ 59.4 9.5 0.0 0.0 2.4 1.2 25.4 6.3 

FA98 HAZ 30.8 4.5 0.0 0.0 11.2 5.3 44.8 6.5 

SP99 HAZ 37.8 2.1 0.8 0.4 11.8 7.7 33.4 5.4 

FA99 HAZ 10.2 4.1 0.2 0.2 13.0 8.3 52.2 7.5 

SP00 HAZ 15.2 5.0 0.0 0.0 15.8 8.0 53.2 8.9 

FA00 HAZ 11.6 3.5 0.0 0.0 11.2 7.1 41.2 8.4 

SP01 HAZ 14.4 4.3 0.2 0.2 12.6 6.1 57.8 10.4 

FA01 HAZ 15.0 4.5 0.0 0.0 19.4 6.0 53.4 5.1 

SP02 HAZ 22.8 5.8 0.0 0.0 18.2 6.6 47.2 8.0 

FA02 HAZ 10.4 2.8 0.4 0.2 34.0 9.6 43.6 5.5 

SP03 HAZ 21.4 5.5 0.4 0.2 21.0 8.3 48.2 6.4 

FA03 HAZ 14.2 4.5 0.4 0.4 18.8 5.5 52.8 6.1 

 

FA95 SHB 64.8 4.8 0.0 0.0 8.4 8.4 18.6 3.5 

SP96 SHB 90.2 1.4 0.2 0.2 0.2 0.2 5.8 1.3 

FA96 SHB 45.2 10.1 0.0 0.0 8.8 6.2 43.4 5.8 

SP97 SHB 58.6 10.8 0.0 0.0 7.4 7.4 29.8 5.2 

FA97 SHB 56.4 13.7 0.0 0.0 16.2 16.2 22.2 3.1 

SP98 SHB 74.4 13.1 0.0 0.0 14.0 14.0 6.8 2.0 

FA98 SHB 28.8 4.4 0.0 0.0 16.8 15.1 47.4 9.7 

SP99 SHB 61.6 7.7 0.0 0.0 11.4 11.4 21.8 4.8 

FA99 SHB 65.4 7.2 0.0 0.0 10.8 10.8 19.8 4.5 
 C-7 

 Endocladia Silvetia Barnacles Rock 

Sample Site mean se mean se mean se mean se 

SP00 SHB 74.2 5.3 0.4 0.4 7.8 7.8 13.4 1.9 

FA00 SHB 16.6 4.3 0.0 0.0 56.8 6.9 20.4 3.1 

SP01 SHB 34.0 7.7 0.0 0.0 51.4 9.6 11.0 2.5 

FA01 SHB 24.6 6.3 0.0 0.0 62.8 8.3 8.2 4.0 

SP02 SHB 43.6 10.8 0.0 0.0 45.6 11.3 8.0 3.1 

FA02 SHB 29.0 6.4 0.4 0.4 57.2 10.4 10.0 3.5 

SP03 SHB 40.2 9.8 0.2 0.2 39.0 11.6 13.4 3.6 

FA03 SHB 24.4 9.5 0.0 0.0 48.6 10.7 20.6 3.7 

 

SP92 OCC 69.6 6.4 0.0 0.0 8.0 2.4 19.6 5.3 

FA92 OCC 32.8 7.1 0.8 0.6 19.4 2.4 40.6 6.5 

SP93 OCC 59.0 6.3 0.0 0.0 11.0 2.1 19.6 4.5 

FA93 OCC 23.8 7.5 0.0 0.0 13.2 5.1 29.6 9.7 

SP94 OCC 36.2 9.9 0.2 0.2 24.6 2.4 29.8 7.8 

FA94 OCC 17.4 5.8 0.2 0.2 46.0 4.9 9.8 4.9 

SP95 OCC 36.4 9.9 0.0 0.0 26.8 6.1 12.0 6.8 

FA95 OCC 30.0 5.9 0.0 0.0 16.0 5.5 15.0 6.0 

SP96 OCC 58.8 7.8 0.0 0.0 13.0 3.9 12.8 4.0 

FA96 OCC 26.4 8.7 0.0 0.0 19.0 2.7 18.4 2.5 

SP97 OCC 42.0 9.8 0.0 0.0 12.8 1.4 16.6 4.6 

FA97 OCC 28.4 4.5 0.0 0.0 4.8 1.5 27.2 7.1 

SP98 OCC 49.8 8.5 0.0 0.0 4.2 1.0 15.0 2.3 

FA98 OCC 16.6 2.4 0.0 0.0 12.0 3.3 31.6 4.3 

SP99 OCC 20.6 5.0 0.0 0.0 5.8 3.4 21.6 4.1 

FA99 OCC 16.0 6.0 0.0 0.0 4.5 2.7 16.5 4.9 

SP00 OCC 27.8 6.5 0.2 0.2 3.4 2.7 13.2 3.7 

FA00 OCC 22.4 8.4 0.0 0.0 5.2 2.9 15.6 4.8 

SP01 OCC 29.8 9.1 0.0 0.0 0.8 0.6 7.0 2.4 

FA01 OCC 17.6 6.9 0.0 0.0 2.0 1.5 8.6 2.9 

SP02 OCC 27.4 7.4 0.0 0.0 0.8 0.4 11.2 2.7 

FA02 OCC 8.4 5.8 0.0 0.0 1.6 1.4 14.6 3.9 

SP03 OCC 15.4 7.7 0.2 0.2 1.2 0.5 24.2 4.6 

FA03 OCC 14.2 6.0 0.0 0.0 2.4 0.9 28.4 7.5 

 

SP92 STA 79.6 2.5 0.8 0.2 0.0 0.0 18.0 1.6 

FA92 STA 19.8 6.7 0.0 0.0 0.0 0.0 66.8 5.0 

SP93 STA 40.8 8.1 1.4 0.7 0.0 0.0 45.8 7.9 

FA93 STA 40.8 5.1 0.8 0.6 0.0 0.0 53.8 4.4 

SP94 STA 56.6 3.0 0.6 0.6 0.0 0.0 38.2 3.6 

FA94 STA 62.0 6.2 2.0 1.0 0.0 0.0 32.4 4.9 

SP95 STA 73.0 3.3 3.2 1.0 0.0 0.0 19.4 4.3 

FA95 STA 47.8 5.8 0.4 0.2 0.0 0.0 41.4 5.2 

SP96 STA 68.4 3.5 0.2 0.2 0.0 0.0 25.4 3.3 

FA96 STA 43.8 6.2 0.0 0.0 0.8 0.5 44.6 7.0 

SP97 STA 62.6 4.6 0.0 0.0 0.0 0.0 30.0 4.6 

FA97 STA 37.6 3.5 0.2 0.2 0.0 0.0 49.4 6.9 
 C-8 

 Endocladia Silvetia Barnacles Rock 

Sample Site mean se mean se mean se mean se 

SP98 STA 71.4 4.6 0.4 0.2 0.0 0.0 22.6 5.2 

FA98 STA 36.0 3.0 0.8 0.6 0.0 0.0 48.6 4.1 

SP99 STA 47.0 3.3 0.4 0.4 0.0 0.0 45.4 2.7 

FA99 STA 39.2 1.8 0.4 0.2 0.0 0.0 47.8 3.7 

SP00 STA 58.0 1.4 0.0 0.0 0.0 0.0 32.2 2.2 

FA00 STA 29.4 4.4 0.0 0.0 0.0 0.0 60.2 5.5 

SP01 STA 45.0 5.4 0.2 0.2 0.0 0.0 40.0 3.8 

FA01 STA 34.2 8.6 2.0 1.0 0.2 0.2 48.2 10.4 

SP02 STA 53.2 13.9 0.6 0.2 0.0 0.0 32.4 8.4 

FA02 STA 31.4 8.2 6.2 1.6 0.0 0.0 35.4 2.9 

SP03 STA 57.2 14.4 2.2 1.0 0.2 0.2 21.4 7.7 

FA03 STA 36.2 10.2 2.6 1.3 0.0 0.0 36.8 8.8 

 

SP92 BOA 76.0 2.6 0.0 0.0 3.2 1.0 11.2 1.8 

FA92 BOA 48.0 7.6 3.0 1.3 4.6 2.0 26.8 10.4 

SP93 BOA 57.4 8.7 3.0 1.3 0.4 0.4 25.4 10.9 

FA93 BOA 54.6 8.0 6.8 3.3 0.0 0.0 19.4 6.1 

SP94 BOA 56.0 3.7 7.2 3.7 0.0 0.0 21.0 6.4 

FA94 BOA 40.4 10.1 25.0 11.3 0.0 0.0 12.6 5.7 

SP95 BOA 50.6 10.0 32.6 13.1 0.2 0.2 11.2 4.6 

FA95 BOA 41.4 14.5 39.4 19.4 0.6 0.6 10.6 5.6 

SP96 BOA 35.8 14.4 40.6 20.5 0.4 0.2 11.8 5.5 

FA96 BOA 36.2 15.2 48.6 21.4 1.2 0.5 5.8 2.4 

SP97 BOA 40.8 16.6 43.8 22.7 0.2 0.2 11.8 6.3 

FA97 BOA 35.6 15.4 47.2 21.6 0.2 0.2 9.8 5.0 

SP98 BOA 46.8 19.2 43.8 22.7 0.4 0.4 4.4 2.2 

FA98 BOA 19.0 9.3 49.0 21.5 0.2 0.2 21.6 10.8 

SP99 BOA 28.4 12.4 47.6 21.3 0.0 0.0 14.8 6.6 

FA99 BOA 11.6 7.3 67.0 12.6 0.0 0.0 14.8 7.9 

SP00 BOA 18.4 8.5 58.8 17.7 0.0 0.0 12.4 5.6 

FA00 BOA 19.0 12.9 63.8 18.3 0.6 0.4 10.8 5.9 

SP01 BOA 31.8 14.6 50.6 19.8 2.0 1.2 10.4 5.0 

FA01 BOA 23.0 11.2 55.6 16.9 1.0 1.0 16.2 5.2 

SP02 BOA 13.4 9.9 70.6 13.7 2.2 2.2 11.2 4.3 

FA02 BOA 16.4 7.3 60.2 18.4 1.6 1.6 8.4 3.7 

SP03 BOA 27.2 12.2 50.4 18.6 2.2 2.2 10.2 3.1 

FA03 BOA 21.8 10.4 55.4 18.0 2.4 1.9 10.2 3.8 

 

SP92 GPT 55.4 5.8 0.0 0.0 15.2 3.8 29.0 5.6 

FA92 GPT 8.6 0.9 0.0 0.0 35.6 6.7 55.2 6.2 

SP93 GPT 52.2 4.3 0.2 0.2 13.6 4.8 29.6 3.0 

FA93 GPT 33.8 2.7 0.8 0.8 8.8 3.6 44.2 1.9 

SP94 GPT 25.8 4.1 0.2 0.2 19.6 5.3 54.2 1.8 

FA94 GPT 38.2 5.9 0.8 0.8 11.4 3.1 46.2 5.8 

SP95 GPT 45.6 3.2 2.2 1.7 4.4 1.7 46.8 3.0 

FA95 GPT 37.4 3.3 3.0 2.8 10.0 3.5 40.0 4.8 
 C-9 

 Endocladia Silvetia Barnacles Rock 

Sample Site mean se mean se mean se mean se 

SP96 GPT 62.0 3.5 2.4 2.4 5.2 1.5 27.8 5.6 

FA96 GPT 22.4 3.0 0.2 0.2 10.6 3.6 58.0 3.9 

SP97 GPT 32.2 4.5 0.2 0.2 17.2 3.5 47.8 2.4 

FA97 GPT 13.8 1.6 0.2 0.2 19.4 5.8 61.0 6.1 

SP98 GPT 19.8 8.1 0.4 0.4 4.6 2.7 59.6 7.7 

FA98 GPT 10.4 2.2 0.6 0.4 8.8 3.9 69.6 6.1 

SP99 GPT 34.4 5.4 0.8 0.4 5.2 2.6 52.4 7.9 

FA99 GPT . . . . . . . . 

SP00 GPT 54.0 5.5 0.4 0.4 2.4 1.5 36.4 7.3 

FA00 GPT 16.4 3.9 0.0 0.0 6.6 4.5 71.4 5.5 

SP01 GPT 25.6 7.4 0.0 0.0 11.2 5.0 58.2 3.8 

FA01 GPT 15.4 5.5 0.0 0.0 17.4 7.5 53.6 2.6 

SP02 GPT 41.6 8.5 0.0 0.0 15.8 7.0 36.2 4.0 

FA02 GPT 16.2 4.3 0.0 0.0 18.8 8.9 55.2 5.6 

SP03 GPT 43.4 8.9 0.0 0.0 12.4 7.7 39.0 4.8 

FA03 GPT 9.6 4.0 0.0 0.0 25.6 12.0 55.2 7.7 

 

FA94 OLDS 25.6 6.2 0.0 0.0 55.2 7.4 7.4 2.4 

SP95 OLDS 29.8 4.1 0.0 0.0 47.8 6.3 11.6 4.9 

FA95 OLDS 37.2 6.6 0.0 0.0 36.0 5.2 17.6 5.7 

SP96 OLDS 45.2 6.8 0.0 0.0 35.8 3.4 12.6 3.2 

FA96 OLDS 29.2 7.8 0.0 0.0 31.0 7.9 15.4 4.6 

SP97 OLDS 27.6 5.8 0.0 0.0 48.6 6.6 8.0 2.1 

FA97 OLDS 50.2 2.2 0.0 0.0 18.6 6.1 13.0 5.0 

SP98 OLDS 12.2 4.0 0.0 0.0 38.6 8.2 37.0 7.1 

FA98 OLDS 18.6 4.0 0.0 0.0 63.8 3.0 6.6 2.2 

SP99 OLDS 18.0 3.7 0.0 0.0 46.0 6.1 20.2 3.8 

FA99 OLDS 25.8 3.2 0.0 0.0 37.6 3.8 16.4 2.8 

SP00 OLDS 23.2 3.0 0.0 0.0 36.0 7.1 25.2 5.7 

FA00 OLDS 19.0 3.1 0.0 0.0 35.4 3.7 25.0 4.6 

SP01 OLDS 36.6 4.0 0.0 0.0 24.0 2.4 23.4 3.6 

FA01 OLDS 28.0 2.7 0.0 0.0 22.8 2.0 17.8 3.9 

SP02 OLDS 14.0 3.3 0.0 0.0 34.8 3.3 41.0 4.3 

FA02 OLDS 11.0 3.7 0.0 0.0 38.6 8.3 25.6 5.5 

SP03 OLDS 20.6 4.0 0.0 0.0 29.0 6.8 42.2 8.4 

FA03 OLDS 21.6 5.6 0.0 0.0 16.8 3.5 11.4 2.6 

 

FA94 PCOV 18.0 2.1 0.0 0.0 43.6 6.6 35.4 4.8 

SP95 PCOV 21.8 3.1 0.0 0.0 17.0 5.4 58.0 5.1 

FA95 PCOV 36.0 6.5 0.0 0.0 16.0 4.6 43.2 4.8 

SP96 PCOV 42.2 5.6 0.0 0.0 7.4 3.1 47.0 5.6 

FA96 PCOV 53.4 6.5 0.0 0.0 4.0 1.2 39.6 6.8 

SP97 PCOV 50.8 6.9 0.0 0.0 1.4 0.6 40.4 6.6 

FA97 PCOV 52.2 8.3 0.0 0.0 2.4 1.9 33.8 7.1 

SP98 PCOV 53.0 9.1 0.0 0.0 1.0 0.3 40.6 9.3 

FA98 PCOV 49.2 5.3 0.0 0.0 12.0 4.1 33.2 5.1 
 C-10 

 Endocladia Silvetia Barnacles Rock 

Sample Site mean se mean se mean se mean se 

SP99 PCOV 34.2 6.3 0.0 0.0 14.2 4.6 45.8 3.6 

FA99 PCOV 55.2 7.5 0.0 0.0 3.0 1.4 33.2 7.9 

SP00 PCOV 51.8 9.4 0.0 0.0 0.4 0.2 39.8 8.8 

FA00 PCOV 60.2 10.5 0.0 0.0 4.6 3.6 29.2 5.7 

SP01 PCOV 54.4 9.8 0.0 0.0 6.4 3.7 29.4 6.2 

FA01 PCOV 57.6 8.0 0.0 0.0 1.2 0.7 21.6 6.5 

SP02 PCOV 40.8 5.7 0.0 0.0 7.2 2.9 40.0 8.1 

FA02 PCOV 60.8 6.9 0.0 0.0 2.4 1.0 24.2 5.0 

SP03 PCOV 57.6 12.5 0.0 0.0 3.4 1.2 32.6 10.9 

FA03 PCOV 54.6 9.1 0.0 0.0 1.6 0.5 27.4 9.3 

 

FA94 WHPT 59.8 2.7 0.0 0.0 8.0 1.9 27.4 2.9 

SP95 WHPT 83.6 4.2 0.0 0.0 5.6 2.5 8.0 1.1 

FA95 WHPT 56.2 2.2 0.0 0.0 4.0 2.4 27.4 2.5 

SP96 WHPT 76.6 4.7 0.0 0.0 8.6 2.2 8.0 1.5 

FA96 WHPT 61.6 1.8 0.0 0.0 7.2 0.9 18.2 4.0 

SP97 WHPT 70.2 4.4 0.0 0.0 9.0 2.4 10.8 2.8 

FA97 WHPT 46.6 7.1 0.0 0.0 18.2 2.1 23.6 5.5 

SP98 WHPT 71.0 4.1 0.0 0.0 9.0 2.7 7.8 2.1 

FA98 WHPT 45.2 5.9 0.0 0.0 11.8 3.1 29.8 2.9 

SP99 WHPT 36.4 4.2 0.0 0.0 8.4 3.5 37.4 3.1 

FA99 WHPT 32.4 3.6 0.0 0.0 19.0 3.4 39.4 0.7 

SP00 WHPT 35.6 2.3 0.0 0.0 18.8 1.8 7.6 1.4 

FA00 WHPT 34.2 4.5 0.0 0.0 10.6 1.9 48.6 2.6 

SP01 WHPT 37.2 4.9 0.0 0.0 16.6 1.6 40.8 4.0 

FA01 WHPT 45.2 3.5 0.0 0.0 16.0 2.1 29.0 1.9 

SP02 WHPT 45.2 12.6 0.0 0.0 18.4 1.8 21.2 3.8 

FA02 WHPT 42.6 3.3 0.0 0.0 19.2 3.6 30.6 1.7 

SP03 WHPT 69.6 3.2 0.0 0.0 12.2 2.2 8.2 1.8 

FA03 WHPT 30.2 4.4 0.0 0.0 14.4 2.2 11.4 1.1 

 

FA96 SHCO 42.0 4.8 0.2 0.2 25.4 4.8 28.0 3.1 

SP97 SHCO 31.6 6.7 0.0 0.0 37.4 6.5 29.2 2.8 

FA97 SHCO 28.2 3.9 1.2 1.2 59.6 4.7 6.4 1.8 

SP98 SHCO 31.8 5.5 1.2 1.2 46.8 6.2 15.6 3.0 

FA98 SHCO 35.2 4.9 1.2 1.0 49.8 4.8 12.0 2.0 

SP99 SHCO 38.4 5.8 2.0 1.8 48.8 4.9 7.2 1.8 

FA99 SHCO 43.0 3.9 1.2 0.8 37.4 4.5 15.8 4.3 

SP00 SHCO 39.8 5.6 2.2 2.0 21.4 3.6 35.0 6.9 

FA00 SHCO 21.0 3.4 2.4 2.4 35.8 5.9 38.0 4.6 

SP01 SHCO 40.4 7.3 1.8 0.8 33.8 4.7 23.2 3.2 

FA01 SHCO 51.6 4.0 1.6 0.8 31.6 4.7 13.4 1.3 

SP02 SHCO 43.6 8.3 1.2 1.0 37.2 6.8 16.4 3.2 

FA02 SHCO 49.0 2.4 4.2 3.1 21.0 4.2 12.4 2.0 

SP03 SHCO 44.6 6.5 4.4 3.1 21.0 5.3 18.6 3.1 

FA03 SHCO 19.6 4.1 13.0 8.4 29.8 4.9 28.2 2.5 
 C-11 

Mastocarpus 

 Mastocarpus Endocladia Rock 

Sample Site mean se mean se mean se 

FA95 PSN 83.4 3.2 5.8 0.7 5.8 2.1 

SP96 PSN 29.8 5.7 26.4 2.8 21.4 6.8 

FA96 PSN 57.0 6.6 18.6 4.8 13.2 6.0 

SP97 PSN 22.0 8.0 38.2 5.3 20.2 4.6 

FA97 PSN 39.6 8.4 24.0 4.1 22.4 3.7 

SP98 PSN 30.2 6.2 34.0 7.6 25.0 5.8 

FA98 PSN 67.4 7.5 10.0 4.7 11.8 2.4 

SP99 PSN 24.8 6.1 30.0 7.0 27.0 6.5 

FA99 PSN 21.4 6.4 16.8 3.0 41.0 10.8 

SP00 PSN 15.4 3.2 23.4 5.2 41.2 9.4 

FA00 PSN 27.2 6.3 17.4 4.4 35.8 7.1 

SP01 PSN 17.2 3.2 28.8 5.4 36.4 9.4 

FA01 PSN 38.0 5.7 22.4 5.4 25.8 9.9 

SP02 PSN 24.6 4.1 35.8 4.7 23.0 8.1 

FA02 PSN 47.8 6.6 13.8 3.8 14.0 7.9 

SP03 PSN 13.0 2.8 37.2 7.1 30.0 9.8 

FA03 PSN 52.2 8.1 17.8 2.0 16.8 6.7 

 

FA95 SHB 61.6 5.2 20.4 5.0 17.4 2.6 

SP96 SHB 35.8 6.7 47.8 6.7 14.0 2.6 

FA96 SHB 37.0 4.2 35.2 6.5 25.8 4.3 

SP97 SHB 35.2 6.5 50.2 5.8 13.2 3.3 

FA97 SHB 33.4 4.9 38.6 4.1 26.0 1.0 

SP98 SHB 30.2 3.7 57.8 5.5 9.4 1.4 

FA98 SHB 17.6 1.7 23.2 4.0 46.8 4.8 

SP99 SHB 21.6 5.2 52.6 3.9 21.8 4.7 

FA99 SHB 17.6 4.8 47.6 4.7 31.4 3.4 

SP00 SHB 16.0 5.0 58.4 6.9 22.6 2.4 

FA00 SHB 7.0 2.1 29.0 6.4 34.6 4.7 

SP01 SHB 15.8 5.5 47.6 7.7 18.0 3.0 

FA01 SHB 4.8 2.0 47.4 7.0 23.4 10.4 

SP02 SHB 9.8 3.7 59.2 6.0 18.6 8.1 

FA02 SHB 16.8 4.5 37.8 6.2 25.8 6.4 

SP03 SHB 21.6 5.3 51.8 5.4 21.2 6.1 

FA03 SHB 11.2 2.4 50.2 6.5 25.4 5.4 
 C-12 

Mazzaella 

 Mazzaella Artic. Coralline Other red 

Sample Site mean se mean se mean se 

FA95 PSN 66.4 4.4 16.0 1.1 12.8 2.3 

SP96 PSN 59.0 6.2 25.6 6.9 11.8 4.4 

FA96 PSN 50.2 6.9 22.2 4.5 23.4 3.4 

SP97 PSN 65.0 4.7 18.0 6.6 12.8 3.0 

FA97 PSN 48.6 5.8 20.4 3.6 28.6 5.5 

SP98 PSN 31.2 7.1 41.0 7.8 19.6 5.8 

FA98 PSN 43.2 3.9 13.6 2.0 38.4 4.8 

SP99 PSN 54.2 3.2 18.2 3.1 22.6 3.3 

FA99 PSN 44.6 3.4 26.0 4.9 20.4 3.3 

SP00 PSN 37.2 2.8 33.4 6.6 19.4 3.3 

FA00 PSN 46.4 6.1 20.4 6.2 25.0 3.4 

SP01 PSN 57.0 4.2 16.8 4.2 7.4 2.5 

FA01 PSN 71.8 3.3 10.6 2.0 4.0 1.8 

SP02 PSN 51.2 3.7 29.8 3.1 7.0 2.3 

FA02 PSN 69.4 6.6 11.0 1.2 10.6 5.5 

SP03 PSN 55.2 2.2 24.2 5.2 7.4 1.9 

FA03 PSN 75.8 6.4 6.8 0.7 7.8 2.8 

 

FA95 HAZ 54.2 3.2 7.6 1.6 16.6 3.9 

SP96 HAZ 37.8 6.3 18.4 1.9 9.6 2.8 

FA96 HAZ 52.4 6.4 10.2 2.9 21.4 5.1 

SP97 HAZ 48.8 4.7 13.4 3.1 13.4 3.0 

FA97 HAZ 45.0 5.1 15.8 3.7 21.6 3.6 

SP98 HAZ 34.0 6.4 8.6 2.9 19.2 4.4 

FA98 HAZ 51.4 3.9 3.6 1.4 30.0 6.3 

SP99 HAZ 49.2 8.3 8.4 2.7 15.4 5.1 

FA99 HAZ 46.8 6.7 9.8 2.7 14.4 4.7 

SP00 HAZ 44.2 5.2 10.8 1.7 17.4 4.7 

FA00 HAZ 46.0 6.0 10.0 0.8 16.2 5.1 

SP01 HAZ 44.2 4.5 6.6 1.4 12.6 3.1 

FA01 HAZ 38.6 8.4 4.2 1.2 10.0 2.4 

SP02 HAZ 40.4 6.7 5.4 0.9 12.2 2.4 

FA02 HAZ 41.4 5.0 2.8 0.9 10.8 1.9 

SP03 HAZ 40.4 7.7 4.0 0.8 10.8 1.7 

FA03 HAZ 57.0 9.8 1.8 0.7 7.4 2.0 
 C-13 

Anthopleura 

 Anthopleura Other red Sand Rock 

Sample Site mean se mean se mean se mean se 

SP92 BOA 14.8 0.9 62.4 10.2 0.0 0.0 5.2 1.7 

FA92 BOA 20.0 1.6 47.6 8.3 9.6 3.9 0.2 0.2 

SP93 BOA 15.2 1.5 74.4 3.6 1.0 1.0 0.8 0.6 

FA93 BOA 15.8 2.7 67.8 5.5 10.4 8.2 0.0 0.0 

SP94 BOA 13.6 1.9 76.2 3.5 0.6 0.6 2.0 1.0 

FA94 BOA 8.6 2.0 69.8 5.5 9.0 3.4 0.0 0.0 

SP95 BOA 15.4 2.2 71.2 2.6 1.0 0.5 0.8 0.5 

FA95 BOA 11.2 1.7 83.8 1.6 3.2 1.9 0.0 0.0 

SP96 BOA 16.8 3.3 78.6 3.8 0.0 0.0 0.8 0.5 

FA96 BOA 10.2 2.2 87.2 2.1 1.4 0.9 0.0 0.0 

SP97 BOA 13.6 1.5 83.0 2.3 0.0 0.0 1.8 1.4 

FA97 BOA 10.8 1.7 86.6 2.5 0.4 0.2 1.0 1.0 

SP98 BOA 12.8 1.4 71.6 5.5 0.0 0.0 11.2 5.9 

FA98 BOA 6.6 2.4 76.4 8.5 0.0 0.0 10.4 10.4 

SP99 BOA 9.6 1.3 64.4 5.5 2.0 0.7 7.8 5.9 

FA99 BOA 10.2 2.5 63.4 9.1 3.4 1.2 9.0 8.0 

SP00 BOA 11.6 1.9 73.6 4.5 0.4 0.2 4.8 4.1 

FA00 BOA 8.2 1.5 91.0 1.7 0.0 0.0 0.4 0.4 

SP01 BOA 12.6 2.0 34.4 4.3 0.8 0.8 1.6 1.1 

FA01 BOA 10.0 2.8 78.8 3.5 3.8 1.2 1.0 0.8 

SP02 BOA 9.6 2.3 32.6 3.7 1.8 1.6 5.6 3.5 

FA02 BOA 16.2 4.2 30.4 6.7 3.0 1.5 3.0 1.4 

SP03 BOA 11.4 2.0 21.4 4.5 0.0 0.0 3.0 1.0 

FA03 BOA 10.6 2.8 27.4 5.5 0.0 0.0 5.8 2.1 

 

SP92 ALEG 94.0 2.4 0.0 0.0 0.0 0.0 4.6 2.2 

FA92 ALEG 85.0 1.9 0.0 0.0 0.8 0.6 12.0 2.1 

SP93 ALEG 76.0 3.9 0.0 0.0 0.0 0.0 21.4 4.7 

FA93 ALEG 86.0 3.0 0.0 0.0 0.0 0.0 12.0 2.3 

SP94 ALEG 88.2 4.6 0.0 0.0 0.8 0.6 7.0 1.5 

FA94 ALEG 72.6 10.3 0.0 0.0 21.0 11.1 1.2 0.6 

SP95 ALEG 77.0 4.0 3.4 2.3 0.0 0.0 13.6 4.1 

FA95 ALEG 69.4 6.3 3.2 2.0 1.4 0.6 13.4 3.4 

SP96 ALEG 74.8 8.4 0.0 0.0 0.0 0.0 16.6 3.7 

FA96 ALEG 65.2 8.5 0.0 0.0 0.0 0.0 19.0 6.0 

SP97 ALEG 67.0 9.8 0.0 0.0 0.0 0.0 21.2 7.3 

FA97 ALEG 72.0 10.1 0.0 0.0 1.2 1.0 12.6 4.8 

SP98 ALEG 53.0 8.7 0.0 0.0 0.0 0.0 34.0 10.2 

FA98 ALEG 40.4 7.1 0.0 0.0 0.0 0.0 29.6 4.3 

SP99 ALEG 42.0 7.2 0.2 0.2 0.8 0.4 44.4 9.4 

FA99 ALEG 68.0 10.0 0.2 0.2 0.6 0.6 11.4 4.2 

SP00 ALEG 72.0 8.8 0.0 0.0 0.6 0.4 25.0 9.3 

FA00 ALEG 72.8 8.5 0.2 0.2 0.8 0.4 12.0 6.0 

SP01 ALEG 79.4 10.5 0.0 0.0 0.2 0.2 6.0 3.1 

FA01 ALEG 79.6 10.1 0.2 0.2 0.4 0.2 6.4 2.8 
 C-14 

 Anthopleura Other red Sand Rock 

Sample Site mean se mean se mean se mean se 

SP02 ALEG 76.8 10.4 0.0 0.0 1.0 0.8 8.6 3.9 

FA02 ALEG 72.0 8.7 0.0 0.0 5.8 2.4 9.0 4.5 

SP03 ALEG 64.4 11.5 0.0 0.0 0.6 0.4 20.4 8.3 

FA03 ALEG 30.2 9.8 0.2 0.2 56.2 16.8 0.2 0.2 

 

SP92 COPT 24.6 2.0 53.0 7.0 1.8 1.3 16.4 4.5 

FA92 COPT 17.4 2.2 72.2 3.4 3.2 1.3 5.6 1.6 

SP93 COPT 22.0 2.1 55.0 9.6 0.0 0.0 20.4 10.6 

FA93 COPT 16.2 3.4 82.2 2.7 1.0 1.0 0.2 0.2 

SP94 COPT 18.6 2.1 61.6 9.0 6.2 4.0 9.2 5.6 

FA94 COPT 0.0 0.0 0.0 0.0 100.0 0.0 0.0 0.0 

SP95 COPT 17.6 4.1 20.2 4.6 3.8 2.3 50.4 8.7 

FA95 COPT 10.6 2.2 77.6 3.1 6.4 1.3 2.2 0.4 

SP96 COPT 13.8 3.2 26.0 4.9 22.8 4.6 10.2 5.0 

FA96 COPT 11.4 3.3 51.2 7.6 23.8 11.2 2.4 0.9 

SP97 COPT 13.6 5.1 14.6 4.2 65.0 10.0 0.0 0.0 

FA97 COPT 14.0 3.6 33.0 4.4 21.0 9.2 23.4 11.0 

SP98 COPT 11.8 1.6 72.6 10.3 0.0 0.0 15.4 8.9 

FA98 COPT 38.0 7.5 37.0 6.6 3.0 3.0 0.0 0.0 

SP99 COPT 2.4 2.4 0.2 0.2 96.4 3.6 0.0 0.0 

FA99 COPT 14.8 2.9 68.0 2.6 6.4 2.0 3.6 1.5 

SP00 COPT 12.6 2.7 41.6 8.3 33.4 9.7 6.0 3.0 

FA00 COPT 15.0 1.9 60.6 3.1 7.2 1.4 10.0 2.3 

SP01 COPT 15.2 2.2 34.6 5.2 25.4 7.5 8.6 4.9 

FA01 COPT 14.8 2.0 67.2 3.1 0.8 0.4 9.8 2.1 

SP02 COPT 15.0 1.8 48.8 4.5 17.0 6.2 7.6 2.3 

FA02 COPT 5.8 5.8 6.0 5.0 86.2 12.8 1.8 1.8 

SP03 COPT 15.8 2.6 68.8 3.4 7.0 1.4 1.4 0.5 

FA03 COPT 18.6 3.6 65.8 3.8 1.0 0.4 11.6 2.9 

 

SP92 CARP 36.6 4.0 10.0 5.8 45.4 11.4 3.4 1.5 

FA92 CARP 33.0 3.7 19.0 4.4 36.2 5.3 5.6 1.9 

SP93 CARP 26.0 1.0 57.4 1.5 10.6 2.2 1.0 0.8 

FA93 CARP 23.8 3.6 57.2 7.6 9.8 4.3 0.8 0.5 

SP94 CARP 25.8 2.2 45.8 7.7 20.0 7.2 2.2 1.3 

FA94 CARP 17.8 3.8 11.6 6.3 67.6 7.7 0.0 0.0 

SP95 CARP 32.0 3.6 44.4 10.6 5.4 3.4 2.2 1.4 

FA95 CARP 23.6 1.7 46.6 9.1 16.8 6.8 3.6 3.6 

SP96 CARP 22.6 2.2 70.8 4.5 1.4 0.5 0.4 0.4 

FA96 CARP 23.0 5.8 16.0 6.9 55.2 11.8 0.0 0.0 

SP97 CARP 29.2 2.0 60.0 3.4 7.2 2.5 0.0 0.0 

FA97 CARP 24.2 3.3 56.4 5.7 7.8 2.0 7.8 3.0 

SP98 CARP 22.4 2.5 74.6 4.2 0.0 0.0 0.4 0.4 

FA98 CARP 16.3 2.1 59.0 4.4 12.3 3.2 0.8 0.8 

SP99 CARP 26.2 1.9 65.6 3.3 4.0 2.3 0.2 0.2 

FA99 CARP 21.0 1.8 61.0 4.5 11.4 3.6 2.0 0.9 
 C-15 

 Anthopleura Other red Sand Rock 

Sample Site mean se mean se mean se mean se 

SP00 CARP 26.8 1.6 56.0 3.1 8.6 1.5 1.6 0.6 

FA00 CARP 29.2 2.6 48.2 6.3 18.6 3.2 3.0 0.7 

SP01 CARP 26.2 3.4 60.6 4.8 7.8 0.9 0.2 0.2 

FA01 CARP 27.8 2.6 38.6 7.3 31.2 5.9 0.4 0.2 

SP02 CARP 18.8 1.0 60.4 3.7 10.0 3.5 0.8 0.8 

FA02 CARP 27.8 1.9 64.0 4.3 1.8 0.6 2.4 1.9 

SP03 CARP 26.0 3.2 49.6 6.0 12.0 7.9 4.2 2.5 

FA03 CARP 28.0 3.2 44.8 5.3 21.4 5.1 1.4 0.7 

 

FA94 MUSH 97.2 1.3 0.4 0.2 0.0 0.0 1.2 0.7 

SP95 MUSH 94.6 1.7 0.2 0.2 0.0 0.0 4.2 1.7 

FA95 MUSH 85.2 9.3 0.0 0.0 0.0 0.0 9.4 7.7 

SP96 MUSH 81.2 8.2 0.0 0.0 0.0 0.0 10.4 4.3 

FA96 MUSH 69.6 14.8 0.0 0.0 14.0 10.4 12.6 5.6 

SP97 MUSH 79.6 5.8 0.0 0.0 0.0 0.0 5.6 2.6 

FA97 MUSH 75.2 9.5 0.4 0.2 6.0 6.0 9.4 2.5 

SP98 MUSH 59.2 4.9 0.0 0.0 0.0 0.0 33.4 4.4 

FA98 MUSH 66.6 5.5 0.0 0.0 0.0 0.0 14.8 3.3 

SP99 MUSH 77.8 4.2 0.0 0.0 0.0 0.0 8.6 3.1 

FA99 MUSH 72.0 7.6 0.0 0.0 0.0 0.0 11.4 2.8 

SP00 MUSH 75.4 11.5 0.0 0.0 0.4 0.4 6.6 2.3 

FA00 MUSH 70.2 15.7 1.6 0.9 3.0 1.7 3.2 1.7 

SP01 MUSH 75.6 16.6 0.0 0.0 0.0 0.0 3.4 1.9 

FA01 MUSH 78.4 15.9 0.4 0.4 0.0 0.0 2.8 0.7 

SP02 MUSH 78.0 17.0 0.4 0.2 0.0 0.0 1.6 0.9 

FA02 MUSH 80.8 17.0 5.2 4.7 0.8 0.6 1.4 0.9 

SP03 MUSH 77.6 17.7 2.8 2.8 0.0 0.0 4.0 1.3 

FA03 MUSH 81.6 15.9 4.0 2.9 0.0 0.0 4.4 3.4 

 
 C-16 

 Anthopleura Other red Sand Rock 

Sample Site mean se mean se mean se mean se 

FA94 OLDS 68.2 10.3 0.0 0.0 2.6 2.6 8.8 4.4 

SP95 OLDS 72.8 10.6 0.0 0.0 4.0 3.5 11.4 10.7 

FA95 OLDS 61.6 8.5 3.2 2.3 1.2 1.2 17.8 3.9 

SP96 OLDS 45.4 13.8 3.2 2.0 35.0 17.3 4.8 2.4 

FA96 OLDS 46.0 14.4 1.4 0.9 38.4 19.2 4.2 2.1 

SP97 OLDS 44.3 21.7 1.3 0.8 49.5 23.4 2.5 2.5 

FA97 OLDS 68.2 5.6 1.0 1.0 9.6 8.9 5.6 2.2 

SP98 OLDS 58.4 10.5 1.0 1.0 20.2 13.0 9.4 6.0 

FA98 OLDS 53.0 8.9 1.4 0.9 0.0 0.0 13.4 7.7 

SP99 OLDS 42.8 14.4 1.2 1.2 27.0 14.6 12.2 5.8 

FA99 OLDS 49.0 12.7 0.2 0.2 5.2 4.7 16.6 12.9 

SP00 OLDS 51.8 13.7 0.8 0.8 5.4 4.3 18.0 10.7 

FA00 OLDS 54.0 6.3 0.8 0.5 0.8 0.8 4.2 1.1 

SP01 OLDS 63.6 6.5 3.2 0.7 2.2 2.2 6.4 1.8 

FA01 OLDS 52.6 10.1 1.2 0.7 0.0 0.0 4.0 1.2 

SP02 OLDS 36.0 12.2 2.2 1.7 30.8 18.6 8.6 4.0 

FA02 OLDS 55.6 12.5 5.8 2.5 2.6 2.6 7.0 3.1 

SP03 OLDS 51.6 10.3 7.0 5.1 1.2 1.2 12.2 4.6 

FA03 OLDS 48.4 9.8 7.2 1.9 0.6 0.6 7.8 2.7 

 
 C-17 

Barnacles 

 Barnacles Endocladia Chthamalus Balanus 

Sample Site mean se mean se mean se mean se 

FA95 PSN 73.4 4.6 5.4 1.2 . . . . 

SP96 PSN 72.2 6.4 5.0 1.4 . . . . 

FA96 PSN 76.8 8.8 3.0 2.0 . . . . 

SP97 PSN 66.6 5.1 3.6 2.0 . . . . 

FA97 PSN 57.2 4.1 1.4 1.2 . . . . 

SP98 PSN 44.6 5.8 2.4 1.6 . . . . 

FA98 PSN 54.0 5.6 1.6 1.0 . . . . 

SP99 PSN 52.0 10.4 2.8 1.7 . . . . 

FA99 PSN 50.4 8.9 2.4 1.9 . . . . 

SP00 PSN 48.8 9.1 3.8 2.5 . . . . 

FA00 PSN 52.8 5.2 0.0 0.0 . . . . 

SP01 PSN 23.0 4.6 2.6 1.4 23.0 4.6 0.0 0.0 

FA01 PSN 26.4 5.1 1.0 0.8 26.4 5.1 0.0 0.0 

SP02 PSN 30.0 4.0 1.2 1.2 29.4 3.8 0.6 0.4 

SU02 PSN 25.0 3.8 0.4 0.2 24.8 3.9 0.2 0.2 

FA02 PSN 30.4 3.2 0.0 0.0 30.4 3.2 0.0 0.0 

SP03 PSN 36.8 3.7 1.8 1.1 36.8 3.7 0.0 0.0 

FA03 PSN 27.6 4.2 0.4 0.4 27.6 4.2 0.0 0.0 

 

FA95 CAY 81.2 2.6 0.4 0.4 . . . . 

SP96 CAY 77.0 2.7 1.8 1.8 . . . . 

FA96 CAY 83.6 1.4 0.0 0.0 . . . . 

SP97 CAY 81.6 2.9 0.2 0.2 . . . . 

FA97 CAY 78.8 4.1 1.2 1.2 . . . . 

SP98 CAY 61.2 7.2 4.6 4.6 . . . . 

FA98 CAY 53.2 4.5 0.0 0.0 . . . . 

SP99 CAY 48.2 6.3 0.0 0.0 . . . . 

FA99 CAY 45.4 5.2 0.0 0.0 . . . . 

SP00 CAY 36.2 5.1 0.0 0.0 . . . . 

FA00 CAY 34.6 6.4 0.0 0.0 . . . . 

SP01 CAY 26.0 3.3 0.0 0.0 24.2 3.5 1.8 1.1 

FA01 CAY 36.4 3.7 0.0 0.0 34.0 4.3 2.4 1.0 

SP02 CAY 31.6 5.4 0.0 0.0 30.8 5.5 0.8 0.2 

SU02 CAY 37.8 6.5 0.0 0.0 37.2 6.7 0.6 0.6 

FA02 CAY 50.4 5.4 0.0 0.0 49.2 5.2 1.2 1.2 

SP03 CAY 51.6 3.8 0.0 0.0 50.2 4.3 1.4 1.0 

FA03 CAY 47.8 4.3 0.0 0.0 46.2 4.1 1.6 0.7 

 

FA95 HAZ 67.0 3.6 4.8 3.8 . . . . 

SP96 HAZ 73.4 3.5 5.4 2.9 . . . . 

FA96 HAZ 68.2 7.2 13.4 7.5 . . . . 

SP97 HAZ 65.2 11.4 20.4 11.4 . . . . 

FA97 HAZ 63.2 10.9 7.4 5.6 . . . . 

SP98 HAZ 42.6 8.4 22.2 11.7 . . . . 

FA98 HAZ 49.4 9.9 9.0 4.2 . . . . 

SP99 HAZ 50.2 10.6 10.0 5.1 . . . . 
 C-18 

 Barnacles Endocladia Chthamalus Balanus 

Sample Site mean se mean se mean se mean se 

FA99 HAZ 47.4 8.6 8.0 3.9 . . . . 

SP00 HAZ 50.4 4.4 9.0 3.9 . . . . 

FA00 HAZ 55.2 7.7 11.6 6.5 . . . . 

SP01 HAZ 51.8 7.1 14.6 8.8 27.2 11.1 24.6 10.6 

FA01 HAZ 58.8 4.1 7.8 5.0 35.2 13.1 23.6 10.1 

SP02 HAZ 49.0 9.9 15.8 10.2 34.6 13.1 14.4 5.1 

SU02 HAZ 42.2 9.8 8.4 5.1 26.0 12.8 16.2 6.2 

FA02 HAZ 41.2 10.1 0.0 0.0 16.4 7.4 24.8 11.2 

SP03 HAZ 19.4 7.4 0.0 0.0 12.0 7.4 7.4 2.5 

FA03 HAZ 46.6 7.8 0.0 0.0 35.8 9.8 10.8 4.6 

 

FA95 SHB 92.4 2.1 0.0 0.0 . . . . 

SP96 SHB 92.4 1.8 3.0 1.3 . . . . 

FA96 SHB 94.0 3.6 0.0 0.0 . . . . 

SP97 SHB 79.4 19.1 0.2 0.2 . . . . 

FA97 SHB 77.8 19.5 0.0 0.0 . . . . 

SP98 SHB 75.6 18.9 0.8 0.6 . . . . 

FA98 SHB 76.8 19.0 0.0 0.0 . . . . 

SP99 SHB 78.0 19.5 0.4 0.4 . . . . 

FA99 SHB 75.4 17.6 0.0 0.0 . . . . 

SP00 SHB 75.4 16.2 0.4 0.4 . . . . 

FA00 SHB 90.6 5.7 0.0 0.0 . . . . 

SP01 SHB 89.8 7.0 0.0 0.0 89.8 7.0 0.0 0.0 

FA01 SHB 86.2 1.7 0.0 0.0 86.2 1.7 0.0 0.0 

SP02 SHB 87.0 2.1 0.0 0.0 87.0 2.1 0.0 0.0 

SU02 SHB 79.8 2.9 0.0 0.0 79.8 2.9 0.0 0.0 

FA02 SHB 90.8 1.7 0.0 0.0 90.8 1.7 0.0 0.0 

SP03 SHB 76.8 1.4 0.0 0.0 76.8 1.4 0.0 0.0 

FA03 SHB 89.6 3.4 0.0 0.0 89.6 3.4 0.0 0.0 

 

SP92 OCC 73.8 3.8 3.8 1.4 . . . . 

FA92 OCC 61.0 11.8 17.2 7.4 . . . . 

SP93 OCC 44.2 12.8 37.0 13.3 . . . . 

FA93 OCC 29.8 6.7 16.4 3.4 . . . . 

SP94 OCC 33.4 4.2 18.2 5.5 . . . . 

FA94 OCC 62.2 5.3 12.0 5.8 . . . . 

SP95 OCC 44.4 12.8 24.6 11.7 . . . . 

FA95 OCC 20.4 7.0 33.4 6.1 . . . . 

SP96 OCC 9.0 1.9 54.0 8.2 . . . . 

FA96 OCC 18.0 2.6 24.0 7.4 . . . . 

SP97 OCC 10.0 2.8 26.8 5.2 . . . . 

FA97 OCC 20.6 6.1 30.0 6.5 . . . . 

SP98 OCC 9.2 4.4 52.6 6.5 . . . . 

FA98 OCC 8.3 1.7 18.8 3.7 . . . . 

SP99 OCC 9.4 1.8 32.4 4.3 . . . . 

FA99 OCC 11.1 1.3 26.6 6.2 . . . . 
 C-19 

 Barnacles Endocladia Chthamalus Balanus 

Sample Site mean se mean se mean se mean se 

SP00 OCC 14.8 1.9 48.0 9.9 . . . . 

FA00 OCC 23.4 2.8 30.8 7.2 . . . . 

SP01 OCC 17.2 3.2 26.4 7.2 

FA01 OCC 9.6 1.6 25.0 3.4 1.6 0.7 8.0 2.0 

SP02 OCC 6.8 2.0 31.8 4.5 0.8 0.8 6.0 1.8 

SU02 OCC 11.2 2.7 22.8 2.1 1.4 0.7 9.8 2.5 

FA02 OCC 7.6 1.6 15.0 2.3 1.8 0.6 5.8 1.6 

SP03 OCC 11.0 3.1 14.6 3.8 3.2 1.1 7.8 2.7 

FA03 OCC 13.6 2.7 10.4 3.2 4.2 1.5 9.4 2.0 

 

SP92 STA 77.6 2.9 0.6 0.4 . . . . 

FA92 STA 82.8 4.9 0.0 0.0 . . . . 

SP93 STA 85.4 2.3 0.6 0.4 . . . . 

FA93 STA 84.4 3.1 4.0 1.7 . . . . 

SP94 STA 78.2 3.6 6.6 2.6 . . . . 

FA94 STA 77.4 6.5 8.0 3.7 . . . . 

SP95 STA 60.8 4.3 16.0 5.3 . . . . 

FA95 STA 70.4 6.5 7.8 3.4 . . . . 

SP96 STA 65.6 6.2 12.6 5.1 . . . . 

FA96 STA 73.8 6.3 2.0 1.3 . . . . 

SP97 STA 75.8 6.8 2.6 1.5 . . . . 

FA97 STA 53.2 5.2 0.0 0.0 . . . . 

SP98 STA 52.4 7.9 3.2 1.6 . . . . 

FA98 STA 34.2 10.0 6.4 2.9 . . . . 

SP99 STA 19.8 9.9 9.6 5.1 . . . . 

FA99 STA 10.2 7.8 5.2 3.6 . . . . 

SP00 STA 4.0 2.5 9.0 6.0 . . . . 

FA00 STA 4.4 1.2 1.6 1.6 . . . . 

SP01 STA 1.2 0.7 4.6 4.1 1.2 0.7 0.0 0.0 

FA01 STA 0.4 0.2 0.4 0.2 0.4 0.2 0.0 0.0 

SP02 STA 0.4 0.2 0.4 0.4 0.2 0.2 0.2 0.2 

SU02 STA 2.8 0.6 0.4 0.4 2.8 0.6 0.0 0.0 

FA02 STA 4.0 1.4 0.8 0.8 4.0 1.4 0.0 0.0 

SP03 STA 3.2 1.3 1.2 1.0 3.2 1.3 0.0 0.0 

FA03 STA 1.0 0.6 1.2 1.0 1.0 0.6 0.0 0.0 

 

SP92 BOA 78.6 3.5 2.8 1.2 . . . . 

FA92 BOA 52.6 8.6 33.0 8.9 . . . . 

SP93 BOA 31.8 12.2 46.0 11.1 . . . . 

FA93 BOA 12.4 10.2 47.6 5.4 . . . . 

SP94 BOA 10.0 9.0 43.2 3.8 . . . . 

FA94 BOA 11.2 7.8 48.8 6.2 . . . . 

SP95 BOA 6.8 4.3 67.2 7.5 . . . . 

FA95 BOA 11.2 5.2 45.2 8.9 . . . . 

SP96 BOA 14.4 8.1 41.0 12.6 . . . . 

FA96 BOA 27.6 14.8 33.4 14.3 . . . . 
 C-20 

 Barnacles Endocladia Chthamalus Balanus 

Sample Site mean se mean se mean se mean se 

SP97 BOA 26.2 14.6 36.0 15.0 . . . . 

FA97 BOA 22.8 15.0 30.0 7.6 . . . . 

SP98 BOA 23.6 14.6 40.0 10.4 . . . . 

FA98 BOA 15.0 8.8 26.6 9.7 . . . . 

SP99 BOA 17.8 11.2 34.2 11.1 . . . . 

FA99 BOA 17.4 11.4 27.6 7.7 . . . . 

SP00 BOA 16.8 12.1 33.8 12.8 . . . . 

FA00 BOA 20.8 11.9 16.6 8.3 . . . . 

SP01 BOA 22.4 12.6 13.8 7.0 22.4 12.6 0.0 0.0 

FA01 BOA 44.0 9.9 4.4 1.8 44.0 9.9 0.0 0.0 

SP02 BOA 45.0 11.7 5.4 2.2 45.0 11.7 0.0 0.0 

SU02 BOA 53.0 11.3 3.2 1.5 52.6 11.4 0.4 0.4 

FA02 BOA 48.6 11.7 3.2 1.8 48.0 11.7 0.6 0.4 

SP03 BOA 49.0 13.7 3.2 2.2 48.6 13.9 0.4 0.4 

FA03 BOA 43.4 12.0 4.2 3.5 43.0 11.9 0.4 0.2 

 

SP92 GPT 65.2 3.3 0.0 0.0 . . . . 

FA92 GPT 73.4 3.3 1.4 0.9 . . . . 

SP93 GPT 72.8 3.6 0.0 0.0 . . . . 

FA93 GPT 60.2 3.3 0.0 0.0 . . . . 

SP94 GPT 53.8 4.3 1.6 1.1 . . . . 

FA94 GPT 39.0 3.8 2.6 1.1 . . . . 

SP95 GPT 49.6 4.3 2.4 1.9 . . . . 

FA95 GPT 54.0 2.8 6.0 1.8 . . . . 

SP96 GPT 49.4 5.4 12.8 3.7 . . . . 

FA96 GPT 56.8 4.4 5.2 3.0 . . . . 

SP97 GPT 58.0 7.8 17.2 5.0 . . . . 

FA97 GPT 61.2 5.5 5.4 2.2 . . . . 

SP98 GPT 37.2 10.6 26.4 8.8 . . . . 

FA98 GPT 27.8 9.8 2.4 1.6 . . . . 

SP99 GPT 27.2 11.8 7.4 2.9 . . . . 

FA99 GPT . . . . . . . . 

SP00 GPT 29.0 10.3 22.2 4.9 . . . . 

FA00 GPT 38.0 11.8 1.6 0.8 . . . . 

SP01 GPT 38.2 10.5 7.2 2.1 19.2 7.4 19.0 3.2 

FA01 GPT 43.2 10.1 12.2 3.6 . . . . 

SP02 GPT 40.0 8.3 20.4 4.3 20.6 4.5 19.4 4.1 

SU02 GPT 37.0 6.1 17.4 4.4 23.2 4.9 13.8 2.6 

FA02 GPT 40.6 4.9 1.0 0.5 14.8 1.5 25.8 4.4 

SP03 GPT 43.2 9.3 13.0 4.4 2.6 0.9 40.6 8.4 

FA03 GPT 41.4 6.4 2.8 1.5 14.6 2.2 26.8 5.2 
 C-21 

 Barnacles Endocladia Barnacles Endocladia 

Sample Site mean se mean se Sample Site mean se mean se 

SP92 ALEG 63.8 4.5 12.6 4.3 FA02 ARHO 84.0 6.4 1.8 1.0 

FA92 ALEG 58.6 8.3 12.2 5.4 SP03 ARHO 75.0 7.4 3.6 2.0 

SP93 ALEG 9.0 5.3 7.4 2.3 FA03 ARHO 80.8 8.3 3.2 1.2 

FA93 ALEG 41.0 12.3 0.2 0.2 

SP94 ALEG 49.0 10.2 3.6 1.1 SP92 CARP 66.6 8.1 2.8 1.2 

FA94 ALEG 43.6 6.1 8.4 3.0 FA92 CARP 69.0 7.1 0.0 0.0 

SP95 ALEG 8.6 2.9 1.2 0.6 SP93 CARP 65.0 3.4 6.2 3.7 

FA95 ALEG 32.4 2.5 2.4 0.9 FA93 CARP 54.8 9.9 1.4 0.7 

SP96 ALEG 21.8 5.2 6.8 2.0 SP94 CARP 58.6 10.1 0.6 0.6 

FA96 ALEG 36.0 9.6 3.6 1.3 FA94 CARP 88.0 2.7 0.4 0.2 

SP97 ALEG 45.6 11.4 3.2 1.2 SP95 CARP 68.4 5.5 4.0 2.5 

FA97 ALEG 52.8 11.2 3.6 1.1 FA95 CARP 57.2 13.2 1.2 0.8 

SP98 ALEG 11.8 6.5 6.0 2.3 SP96 CARP 52.2 8.4 1.8 1.1 

FA98 ALEG 16.8 8.4 1.3 0.8 FA96 CARP 49.6 9.0 0.0 0.0 

SP99 ALEG 36.0 9.0 0.8 0.5 SP97 CARP 46.8 5.1 2.2 1.7 

FA99 ALEG 42.0 9.3 3.4 1.8 FA97 CARP 44.8 5.7 0.0 0.0 

SP00 ALEG 32.8 8.7 10.0 2.9 SP98 CARP 35.6 4.9 1.6 0.7 

FA00 ALEG 41.2 9.4 6.8 2.4 FA98 CARP 32.6 4.6 0.0 0.0 

SP01 ALEG 27.2 6.5 20.8 7.3 SP99 CARP 31.2 9.8 2.2 1.7 

FA01 ALEG 33.6 7.2 8.0 2.8 FA99 CARP 43.2 9.8 0.2 0.2 

SP02 ALEG 23.0 6.5 11.4 3.1 SP00 CARP 44.4 9.9 0.4 0.2 

FA02 ALEG 39.4 10.0 12.4 5.5 FA00 CARP 48.8 11.0 0.0 0.0 

SP03 ALEG 18.6 5.1 24.4 11.0 SP01 CARP 47.6 11.1 1.0 0.5 

FA03 ALEG 56.0 8.3 1.6 1.4 FA01 CARP 50.0 8.8 0.0 0.0 

 SP02 CARP 48.6 9.8 1.4 0.6 

SP92 ARHO 80.6 5.0 5.4 2.4 FA02 CARP 53.0 11.5 0.0 0.0 

FA92 ARHO 80.2 4.6 3.6 2.3 SP03 CARP 46.6 7.5 5.0 2.4 

SP93 ARHO 51.8 7.3 7.8 4.0 FA03 CARP 61.2 13.5 0.2 0.2 

FA93 ARHO 75.2 3.1 7.2 3.6 

SP94 ARHO 77.0 7.5 5.6 3.7 FA94 MUSH 94.4 3.9 0.0 0.0 

FA94 ARHO 54.8 3.2 5.8 2.7 SP95 MUSH 68.0 5.6 0.0 0.0 

SP95 ARHO 46.2 7.5 11.6 4.8 FA95 MUSH 90.0 2.9 0.0 0.0 

FA95 ARHO 73.2 2.9 2.4 1.5 SP96 MUSH 5.4 3.2 0.0 0.0 

SP96 ARHO 64.2 4.7 8.8 4.1 FA96 MUSH 75.4 10.2 0.0 0.0 

FA96 ARHO 78.6 5.5 10.4 4.4 SP97 MUSH 84.0 4.8 0.0 0.0 

SP97 ARHO 65.0 12.5 11.4 5.6 FA97 MUSH 81.0 3.8 0.0 0.0 

FA97 ARHO 21.4 5.0 0.6 0.4 SP98 MUSH 58.4 4.2 0.0 0.0 

SP98 ARHO 18.0 6.1 4.8 2.4 FA98 MUSH 87.0 2.5 0.0 0.0 

FA98 ARHO 0.4 0.2 2.6 1.7 SP99 MUSH 85.0 1.3 0.0 0.0 

SP99 ARHO 13.4 6.0 3.2 1.6 FA99 MUSH 45.6 6.7 0.0 0.0 

FA99 ARHO 28.6 10.5 1.4 0.7 SP00 MUSH 54.0 9.3 0.0 0.0 

SP00 ARHO 34.5 12.8 3.5 1.9 FA00 MUSH 77.6 4.1 0.0 0.0 

FA00 ARHO 77.0 4.0 0.2 0.2 SP01 MUSH 74.4 14.5 0.0 0.0 

SP01 ARHO 68.8 4.9 3.6 2.0 FA01 MUSH 75.6 2.6 0.0 0.0 

FA01 ARHO 56.2 7.7 3.2 1.4 SP02 MUSH 41.0 6.6 0.0 0.0 

SP02 ARHO 62.6 11.8 3.0 1.4 FA02 MUSH 85.4 2.6 0.0 0.0 
 C-22 

 Barnacles Endocladia Barnacles Endocladia 

Sample Site mean se mean se Sample Site mean se mean se 

SP03 MUSH 63.6 9.9 0.0 0.0 SP96 WHPT 82.6 4.1 1.6 1.6 

FA03 MUSH 92.6 4.1 0.0 0.0 FA96 WHPT 71.5 14.7 0.0 0.0 

 SP97 WHPT 73.8 6.0 0.0 0.0 

FA94 OLDS 96.2 2.0 1.0 0.8 FA97 WHPT 64.4 10.7 0.0 0.0 

SP95 OLDS 91.8 2.0 1.8 1.1 SP98 WHPT 76.0 3.4 0.8 0.6 

FA95 OLDS 86.0 4.3 1.0 0.8 FA98 WHPT 24.6 2.7 0.0 0.0 

SP96 OLDS 85.2 3.4 2.2 1.4 SP99 WHPT 35.0 2.2 0.0 0.0 

FA96 OLDS 80.8 5.5 3.0 1.8 FA99 WHPT 60.4 4.0 0.0 0.0 

SP97 OLDS 71.6 6.3 4.8 3.0 SP00 WHPT 51.0 8.1 0.0 0.0 

FA97 OLDS 71.6 10.9 10.8 6.6 FA00 WHPT 40.4 5.7 0.0 0.0 

SP98 OLDS 56.4 11.0 3.2 2.0 SP01 WHPT 59.4 7.4 0.0 0.0 

FA98 OLDS 85.8 2.6 2.4 1.5 FA01 WHPT 61.2 2.3 0.0 0.0 

SP99 OLDS 46.2 10.5 1.6 1.0 SP02 WHPT 56.8 2.0 0.0 0.0 

FA99 OLDS 74.2 5.2 0.8 0.4 FA02 WHPT 70.2 4.0 0.0 0.0 

SP00 OLDS 69.4 4.4 1.6 1.0 SP03 WHPT 57.0 5.1 0.0 0.0 

FA00 OLDS 68.4 6.9 2.2 1.5 FA03 WHPT 61.0 5.3 0.0 0.0 

SP01 OLDS 69.4 8.4 5.4 3.0 

FA01 OLDS 55.0 9.4 5.2 2.0 FA99 PTFM 56.6 3.2 0.0 0.0 

SP02 OLDS 64.2 8.6 1.6 0.8 SP00 PTFM 47.2 4.0 0.0 0.0 

FA02 OLDS 80.6 6.6 2.2 0.7 FA00 PTFM 33.0 2.8 0.0 0.0 

SP03 OLDS 75.6 6.6 8.4 3.4 SP01 PTFM 34.2 2.7 0.0 0.0 

FA03 OLDS 55.0 11.0 15.0 3.6 FA01 PTFM 42.0 2.7 0.0 0.0 

 SP02 PTFM 39.8 5.3 0.0 0.0 

FA94 PCOV 80.8 3.8 0.0 0.0 FA02 PTFM 36.6 5.5 0.0 0.0 

SP95 PCOV 72.2 5.1 0.4 0.4 SP03 PTFM 33.6 6.7 0.0 0.0 

FA95 PCOV 72.4 6.5 4.0 4.0 FA03 PTFM 64.0 8.1 0.0 0.0 

SP96 PCOV 70.0 4.0 4.4 2.8 

FA96 PCOV 85.0 3.6 2.0 2.0 FA96 CRCO 80.8 1.7 0.0 0.0 

SP97 PCOV 74.6 5.4 3.4 3.4 SP97 CRCO 70.6 3.8 0.0 0.0 

FA97 PCOV 71.6 5.0 6.4 4.2 FA97 CRCO 69.4 2.6 0.0 0.0 

SP98 PCOV 62.4 7.2 13.6 9.8 SP98 CRCO 57.6 11.0 0.4 0.4 

FA98 PCOV 66.8 4.7 5.6 4.1 FA98 CRCO 58.8 15.1 0.0 0.0 

SP99 PCOV 70.8 8.1 6.4 4.1 SP99 CRCO 82.0 10.5 0.0 0.0 

FA99 PCOV 73.0 6.8 5.0 4.5 FA99 CRCO 63.2 7.2 0.0 0.0 

SP00 PCOV 74.8 3.6 3.6 2.7 SP00 CRCO 73.8 4.7 0.0 0.0 

FA00 PCOV 57.2 5.2 5.8 5.3 FA00 CRCO 70.0 4.9 0.0 0.0 

SP01 PCOV 62.8 3.2 1.6 1.2 SP01 CRCO 77.8 3.5 0.0 0.0 

FA01 PCOV 62.0 4.6 4.6 3.1 FA01 CRCO 67.8 4.2 6.8 1.7 

SP02 PCOV 69.4 3.8 2.6 1.7 SP02 CRCO 72.2 2.9 0.2 0.2 

FA02 PCOV 70.2 4.3 3.2 1.8 FA02 CRCO 61.0 6.3 0.0 0.0 

SP03 PCOV 63.0 5.8 13.8 8.3 SP03 CRCO 58.0 6.8 0.0 0.0 

FA03 PCOV 62.4 9.2 16.2 12.1 FA03 CRCO 55.0 7.4 0.0 0.0 

 

FA94 WHPT 79.4 5.4 0.0 0.0 FA96 SHCO 84.4 1.4 0.0 0.0 

SP95 WHPT 84.4 4.6 0.0 0.0 SP97 SHCO 77.2 1.0 0.0 0.0 

FA95 WHPT 77.4 5.8 1.0 1.0 FA97 SHCO 87.4 1.1 0.0 0.0 
 C-23 

 Barnacles Endocladia Barnacles Endocladia 

Sample Site mean se mean se Sample Site mean se mean se 

SP98 SHCO 81.0 2.2 0.0 0.0 FA94 CTBR 65.8 3.0 0.0 0.0 

FA98 SHCO 85.4 2.8 2.4 2.4 SP95 CTBR 61.8 5.4 0.0 0.0 

SP99 SHCO 90.0 1.7 0.2 0.2 FA95 CTBR 52.6 6.4 0.0 0.0 

FA99 SHCO 72.8 3.3 0.0 0.0 SP96 CTBR 61.8 3.3 0.0 0.0 

SP00 SHCO 65.2 3.4 0.0 0.0 FA96 CTBR 54.2 4.9 0.0 0.0 

FA00 SHCO 50.4 6.0 0.2 0.2 SP97 CTBR 58.6 3.1 0.0 0.0 

SP01 SHCO 67.0 1.4 0.2 0.2 FA97 CTBR 60.6 3.2 0.0 0.0 

FA01 SHCO 70.8 4.5 0.4 0.4 SP98 CTBR 62.2 3.9 0.0 0.0 

SP02 SHCO 69.8 1.6 0.0 0.0 FA98 CTBR 66.2 3.3 0.0 0.0 

FA02 SHCO 59.2 0.4 0.8 0.6 SP99 CTBR 62.4 5.1 0.0 0.0 

SP03 SHCO 60.2 3.1 0.8 0.6 FA99 CTBR 44.2 2.8 0.0 0.0 

FA03 SHCO 43.6 3.7 1.0 0.8 SP00 CTBR 34.4 4.5 0.0 0.0 

 FA00 CTBR 30.6 5.5 0.0 0.0 

FA96 TRIS 86.8 0.9 0.0 0.0 SP01 CTBR 36.2 4.9 0.0 0.0 

SP97 TRIS 78.8 3.7 0.0 0.0 FA01 CTBR 26.8 2.3 0.0 0.0 

FA97 TRIS 83.8 1.2 0.0 0.0 SP02 CTBR 36.2 3.6 0.0 0.0 

SP98 TRIS 81.2 3.6 0.0 0.0 FA02 CTBR 32.4 3.0 0.0 0.0 

FA98 TRIS 69.0 5.3 0.0 0.0 SP03 CTBR 28.4 4.0 0.0 0.0 

SP99 TRIS 78.6 2.9 0.0 0.0 

FA99 TRIS 92.2 1.3 0.0 0.0 FA94 CTLH 80.8 2.1 0.0 0.0 

SP00 TRIS 78.2 4.2 0.0 0.0 SP95 CTLH 77.0 3.1 0.0 0.0 

FA00 TRIS 65.8 5.9 0.0 0.0 FA95 CTLH 73.4 10.9 0.0 0.0 

SP01 TRIS 73.0 5.5 0.0 0.0 SP96 CTLH 66.4 12.6 0.0 0.0 

FA01 TRIS 58.2 5.8 0.0 0.0 FA96 CTLH 85.8 3.3 0.0 0.0 

SP02 TRIS 61.2 5.1 0.0 0.0 SP97 CTLH 66.4 4.2 0.0 0.0 

FA02 TRIS 37.2 6.3 0.0 0.0 FA97 CTLH 62.4 4.1 0.0 0.0 

SP03 TRIS 40.6 8.9 0.0 0.0 SP98 CTLH 49.2 9.9 0.8 0.6 

FA03 TRIS 37.8 6.7 0.0 0.0 FA98 CTLH 77.8 3.9 0.0 0.0 

 SP99 CTLH 84.4 1.9 0.0 0.0 

FA96 DAPT 85.8 1.6 0.0 0.0 FA99 CTLH 71.2 5.1 0.0 0.0 

SP97 DAPT 88.4 2.4 0.0 0.0 SP00 CTLH 61.4 5.4 0.0 0.0 

FA97 DAPT 84.6 3.1 0.0 0.0 FA00 CTLH 62.2 3.0 0.0 0.0 

SP98 DAPT 75.0 1.8 0.0 0.0 SP01 CTLH 79.0 4.9 0.0 0.0 

FA98 DAPT 90.6 1.8 0.0 0.0 FA01 CTLH 80.8 1.9 0.0 0.0 

SP99 DAPT 95.4 2.0 0.0 0.0 SP02 CTLH 77.2 3.1 0.0 0.0 

FA99 DAPT 94.8 2.5 0.0 0.0 FA02 CTLH 71.2 4.3 0.0 0.0 

SP00 DAPT 87.8 2.1 0.0 0.0 SP03 CTLH 60.6 6.6 0.0 

FA00 DAPT 78.0 1.8 0.0 0.0 

SP01 DAPT 86.2 1.6 0.0 0.0 

FA01 DAPT 90.4 1.4 0.0 0.0 

SP02 DAPT 89.0 2.0 0.0 0.0 

FA02 DAPT 80.6 1.2 0.0 0.0 

SP03 DAPT 64.4 1.1 0.0 0.0 

FA03 DAPT 62.2 3.5 0.0 0.0 

 
 C-24 

Pollicipes 

 Pollicipes Mytilus Rock 

Sample Site mean se mean se mean se 

SP92 GPT 37.4 4.0 4.2 2.6 38.6 3.7 

FA92 GPT 40.4 5.6 4.2 1.7 33.4 5.9 

SP93 GPT 41.6 6.7 5.6 2.7 27.6 4.0 

FA93 GPT 38.6 8.4 4.4 2.6 34.8 8.6 

SP94 GPT 33.6 5.8 5.0 2.0 32.8 8.1 

FA94 GPT 32.2 5.5 5.4 2.0 35.8 8.4 

SP95 GPT 36.2 7.7 5.6 2.7 29.4 3.9 

FA95 GPT 30.2 6.2 8.8 3.8 30.8 7.1 

SP96 GPT 31.6 7.1 8.4 3.7 23.6 6.3 

FA96 GPT 30.0 8.2 7.4 2.8 33.2 8.7 

SP97 GPT 30.0 8.0 7.4 2.9 35.0 7.3 

FA97 GPT 27.6 7.9 9.4 3.9 33.8 6.5 

SP98 GPT 33.4 9.5 7.2 2.9 26.8 6.4 

FA98 GPT 28.6 7.9 7.6 3.1 26.8 6.0 

SP99 GPT 30.2 7.1 8.4 3.4 30.4 6.4 

FA99 GPT . . . . . . 

SP00 GPT 30.2 6.8 13.2 5.2 28.4 8.4 

FA00 GPT 32.6 6.9 10.4 4.4 27.2 8.0 

SP01 GPT 29.2 6.5 13.8 5.2 30.2 5.8 

FA01 GPT 29.2 4.9 12.4 5.6 32.8 7.6 

SP02 GPT 29.0 6.1 14.6 6.3 35.8 9.5 

FA02 GPT 27.0 5.6 13.4 6.6 27.8 7.4 

SP03 GPT 22.6 4.4 16.2 6.8 35.4 6.0 

FA03 GPT 29.6 5.7 13.2 5.6 31.0 7.7 

 

SP93 ALEG 17.4 4.8 38.2 13.7 42.6 15.1 

FA93 ALEG 23.0 7.0 41.8 14.6 31.8 14.4 

SP94 ALEG 19.0 5.6 44.2 14.2 31.6 12.3 

FA94 ALEG 18.2 6.0 42.2 13.2 30.0 9.4 

SP95 ALEG 17.2 5.9 44.6 14.3 30.8 10.7 

FA95 ALEG 22.8 6.5 46.8 14.3 22.4 9.8 

SP96 ALEG 19.2 5.4 49.8 14.4 28.4 14.1 

FA96 ALEG 17.8 5.4 46.8 14.4 27.6 9.7 

SP97 ALEG 15.8 5.5 46.2 14.3 29.8 8.5 

FA97 ALEG 18.2 6.0 43.8 14.9 29.0 8.9 

SP98 ALEG 11.0 3.8 48.0 15.5 37.2 15.1 

FA98 ALEG 14.6 4.3 48.6 16.2 30.4 12.0 

SP99 ALEG 13.4 4.5 54.2 17.1 27.4 13.1 

FA99 ALEG 20.6 6.4 53.0 16.3 21.2 12.5 

SP00 ALEG 15.0 5.8 51.0 15.5 28.6 11.0 

FA00 ALEG 16.6 6.6 49.2 14.8 24.6 7.5 

SP01 ALEG 16.8 4.4 43.4 12.8 33.4 8.9 

FA01 ALEG 19.6 6.7 41.6 13.5 27.8 6.1 

SP02 ALEG 20.8 6.5 41.4 14.3 26.0 6.0 

FA02 ALEG 19.2 7.5 45.0 14.0 22.4 5.0 
 C-25 

 Pollicipes Mytilus Rock 

Sample Site mean se mean se mean se 

SP03 ALEG 18.4 5.5 46.2 14.2 25.8 6.2 

FA03 ALEG 23.8 7.6 49.0 12.8 14.2 5.1 

 

SP92 CARP 54.2 5.5 8.0 3.2 28.8 4.4 

FA92 CARP 61.4 6.7 13.6 4.4 16.0 2.7 

SP93 CARP 51.6 5.1 19.0 5.7 21.2 2.9 

FA93 CARP 62.4 6.4 22.8 6.4 9.2 3.1 

SP94 CARP 48.0 4.8 24.6 5.8 22.8 3.7 

FA94 CARP 56.8 8.5 26.4 5.4 10.6 5.8 

SP95 CARP 41.6 8.5 31.0 7.6 16.2 7.6 

FA95 CARP 53.8 3.0 22.2 4.1 13.6 3.4 

SP96 CARP 32.2 8.1 12.8 3.8 28.2 5.7 

FA96 CARP 46.8 6.3 18.2 3.7 15.6 4.8 

SP97 CARP 46.6 4.8 23.0 3.2 16.6 4.3 

FA97 CARP 34.4 1.9 32.6 5.7 24.0 5.6 

SP98 CARP 29.2 3.5 24.2 3.5 36.2 6.3 

FA98 CARP 41.0 4.4 28.2 3.4 19.6 4.7 

SP99 CARP 41.4 4.8 34.2 5.5 17.2 6.7 

FA99 CARP 42.2 5.9 29.4 3.4 8.6 3.7 

SP00 CARP 32.6 2.9 33.4 3.0 16.8 3.1 

FA00 CARP 32.0 4.3 43.0 7.9 13.0 4.2 

SP01 CARP 27.6 3.0 39.2 4.5 17.4 1.1 

FA01 CARP 35.6 5.8 31.0 8.0 13.8 2.6 

SP02 CARP 36.2 2.0 27.4 6.0 19.8 3.9 

FA02 CARP 35.0 3.8 39.2 6.7 11.8 2.6 

SP03 CARP 35.0 2.9 44.2 7.0 11.2 5.8 

FA03 CARP 32.0 5.0 52.4 5.7 7.4 4.2 

 

FA94 CTBR 26.8 3.1 43.2 4.2 5.0 1.9 

SP95 CTBR 38.4 3.4 30.6 2.7 4.4 2.2 

FA95 CTBR 25.8 3.1 63.6 3.9 3.8 2.0 

SP96 CTBR 20.4 2.6 68.2 4.5 4.2 2.4 

FA96 CTBR 27.0 2.3 60.0 4.1 4.0 1.3 

SP97 CTBR 27.2 3.1 60.8 5.8 3.8 2.1 

FA97 CTBR 29.0 2.4 52.8 4.9 3.8 1.9 

SP98 CTBR 38.8 3.6 37.6 3.7 4.4 1.5 

FA98 CTBR 33.6 2.9 47.8 4.4 2.2 1.3 

SP99 CTBR 39.2 3.7 45.6 2.5 2.0 0.9 

FA99 CTBR 43.0 4.1 45.8 3.4 3.8 1.7 

SP00 CTBR 49.6 3.9 41.6 4.0 0.4 0.2 

FA00 CTBR 40.2 3.0 50.6 4.0 4.2 1.7 

SP01 CTBR 43.4 4.1 50.6 3.1 2.4 1.7 

FA01 CTBR 33.8 2.7 58.4 2.9 2.6 1.2 

SP02 CTBR 41.2 2.2 51.2 3.5 3.4 1.8 

FA02 CTBR 35.4 4.0 56.0 3.8 2.8 1.6 

SP03 CTBR 39.6 2.3 51.8 3.7 3.4 2.1 
 C-26 

Mytilus 

 Mytilus Pollicipes Rock 

Sample Site mean se mean se mean se 

FA95 PSN 84.0 1.9 9.2 1.7 4.2 2.5 

SP96 PSN 78.4 3.8 11.6 2.7 5.6 2.6 

FA96 PSN 78.6 3.0 8.8 1.4 7.0 3.5 

SP97 PSN 79.6 2.5 9.4 2.1 3.8 1.8 

FA97 PSN 80.8 2.5 7.8 1.5 4.8 1.6 

SP98 PSN 80.6 5.4 6.6 1.4 4.8 2.1 

FA98 PSN 83.8 4.1 5.4 1.5 3.6 1.2 

SP99 PSN 82.8 3.4 8.8 1.6 4.6 2.1 

FA99 PSN 87.2 3.0 7.4 2.0 2.8 1.4 

SP00 PSN 84.0 3.7 8.8 2.7 3.4 1.4 

FA00 PSN 84.0 4.1 8.4 1.5 3.4 1.9 

SP01 PSN 83.0 2.9 9.0 1.4 4.2 2.0 

FA01 PSN 88.8 3.6 5.2 2.0 1.4 0.7 

SP02 PSN 85.6 2.5 9.0 1.9 2.0 1.3 

FA02 PSN 77.8 5.1 8.8 2.6 2.4 0.9 

SP03 PSN 87.8 3.1 7.2 1.5 1.2 1.2 

FA03 PSN 88.2 3.1 6.8 1.7 2.2 1.1 

 

FA95 CAY 69.4 4.9 8.0 3.1 19.6 1.8 

SP96 CAY 72.2 4.7 6.8 2.6 18.8 3.2 

FA96 CAY 68.2 5.5 6.8 3.1 22.6 3.5 

SP97 CAY 70.6 5.3 4.8 2.4 21.4 3.1 

FA97 CAY 67.6 4.7 5.8 2.0 22.6 3.6 

SP98 CAY 67.0 5.0 5.2 2.1 23.2 3.0 

FA98 CAY 66.0 5.0 3.4 1.5 27.8 4.7 

SP99 CAY 64.6 4.5 4.4 1.9 27.6 3.7 

FA99 CAY 57.8 5.6 5.4 2.7 32.4 3.9 

SP00 CAY 56.8 3.4 4.0 1.3 35.8 3.8 

FA00 CAY 48.0 5.9 5.4 2.6 41.8 5.3 

SP01 CAY 46.4 3.7 3.2 1.6 44.8 3.9 

FA01 CAY 49.8 6.7 1.0 0.6 34.6 4.9 

SP02 CAY 52.6 7.6 1.4 0.9 31.6 5.4 

FA02 CAY 56.0 8.4 0.8 0.8 27.2 5.3 

SP03 CAY 60.6 7.9 1.2 0.6 25.6 5.4 

FA03 CAY 66.8 7.4 0.6 0.6 26.2 6.0 

 

FA95 HAZ 47.4 4.3 17.6 4.2 20.4 1.7 

SP96 HAZ 46.0 5.3 16.4 4.8 14.2 1.7 

FA96 HAZ 44.8 5.3 18.8 5.5 9.2 0.8 

SP97 HAZ 45.6 4.5 16.4 3.9 12.0 2.3 

FA97 HAZ 50.8 8.1 17.4 4.0 10.6 2.0 

SP98 HAZ 54.4 5.9 14.8 3.3 11.2 2.2 

FA98 HAZ 41.4 4.9 13.8 3.9 19.4 2.6 

SP99 HAZ 41.2 5.1 12.4 2.9 20.4 2.6 

FA99 HAZ 31.2 3.7 16.8 2.9 25.8 2.7 
 C-27 

 Mytilus Pollicipes Rock 

Sample Site mean se mean se mean se 

SP00 HAZ 34.4 4.5 13.8 2.9 26.0 2.2 

FA00 HAZ 26.6 3.8 16.0 2.1 16.4 2.2 

SP01 HAZ 31.2 5.1 12.0 2.4 13.6 1.9 

FA01 HAZ 32.2 4.2 17.0 2.7 13.2 3.0 

SP02 HAZ 32.2 5.7 17.0 3.1 14.6 2.4 

FA02 HAZ 34.8 5.9 17.0 3.9 13.0 3.1 

SP03 HAZ 40.0 6.3 16.4 4.2 17.4 1.7 

FA03 HAZ 35.4 5.4 16.6 2.7 16.0 1.4 

 

FA95 SHB 91.2 2.5 0.0 0.0 2.0 0.7 

SP96 SHB 88.6 3.9 0.0 0.0 3.0 1.2 

FA96 SHB 90.4 2.4 0.0 0.0 2.8 0.8 

SP97 SHB 91.8 2.6 0.0 0.0 1.0 0.4 

FA97 SHB 41.8 8.5 0.0 0.0 20.0 4.4 

SP98 SHB 35.0 8.2 0.0 0.0 16.4 3.6 

FA98 SHB 37.2 6.7 0.0 0.0 18.4 7.9 

SP99 SHB 37.4 7.6 0.0 0.0 25.2 8.8 

FA99 SHB 33.6 7.3 0.0 0.0 34.8 11.4 

SP00 SHB 33.2 8.1 0.0 0.0 35.6 9.0 

FA00 SHB 27.2 6.0 0.0 0.0 40.8 7.8 

SP01 SHB 32.8 6.4 0.0 0.0 35.0 6.1 

FA01 SHB 39.0 6.1 0.0 0.0 26.8 4.1 

SP02 SHB 37.2 6.2 0.0 0.0 27.4 3.9 

FA02 SHB 42.4 6.0 0.0 0.0 14.2 2.7 

SP03 SHB 52.2 6.3 0.0 0.0 15.6 1.7 

FA03 SHB 66.8 2.1 0.0 0.0 8.4 0.9 

 

SP92 OCC 85.8 4.5 2.4 1.9 7.0 2.8 

FA92 OCC 87.6 3.4 1.4 1.4 4.8 3.0 

SP93 OCC 80.2 5.8 1.8 0.9 10.4 4.7 

FA93 OCC 85.8 5.1 2.2 1.0 7.4 4.7 

SP94 OCC 79.4 4.9 2.4 1.2 9.6 4.2 

FA94 OCC 78.0 4.1 3.6 1.6 4.6 2.6 

SP95 OCC 71.0 12.0 2.8 1.7 17.2 9.1 

FA95 OCC 77.8 10.7 2.0 1.0 3.8 1.7 

SP96 OCC 74.2 11.3 3.6 1.2 4.8 2.4 

FA96 OCC 78.0 10.0 3.4 1.8 3.2 1.8 

SP97 OCC 85.4 5.2 2.2 1.0 6.6 3.3 

FA97 OCC 80.0 4.7 1.4 0.9 7.4 3.7 

SP98 OCC 79.4 5.1 2.0 0.9 12.8 4.2 

FA98 OCC 84.0 5.9 2.8 1.8 8.8 4.7 

SP99 OCC 82.2 3.5 3.4 1.1 9.0 2.6 

FA99 OCC 85.4 3.7 3.2 1.6 5.6 1.5 

SP00 OCC 78.0 3.8 4.0 1.3 10.6 2.6 

FA00 OCC 78.8 2.9 2.2 1.5 8.0 2.8 
 C-28 

 Mytilus Pollicipes Rock 

Sample Site mean se mean se mean se 

SP01 OCC 77.6 4.9 3.8 1.9 10.6 4.1 

FA01 OCC 77.6 6.3 2.6 1.9 7.2 3.7 

SP02 OCC 74.4 6.0 4.4 2.1 14.4 5.5 

FA02 OCC 79.6 6.3 3.4 1.5 4.6 1.8 

SP03 OCC 80.4 6.6 3.8 1.5 10.8 5.4 

FA03 OCC 80.6 5.3 4.6 2.2 5.8 2.8 

 

SP92 STA 84.6 4.0 0.6 0.4 10.0 3.8 

FA92 STA 85.0 4.2 0.6 0.4 6.0 1.5 

SP93 STA 86.0 4.6 0.2 0.2 6.4 2.5 

FA93 STA 91.4 4.1 0.2 0.2 5.0 2.8 

SP94 STA 88.4 4.2 0.0 0.0 6.2 2.4 

FA94 STA 85.2 4.7 0.0 0.0 5.2 1.9 

SP95 STA 87.4 3.8 0.4 0.2 4.8 1.4 

FA95 STA 92.2 2.8 0.6 0.2 4.0 2.1 

SP96 STA 93.8 2.2 0.8 0.6 2.4 1.0 

FA96 STA 94.8 2.1 0.2 0.2 2.2 1.1 

SP97 STA 93.6 2.3 0.4 0.2 4.0 1.7 

FA97 STA 75.4 7.0 0.0 0.0 21.6 6.0 

SP98 STA 63.8 12.5 0.2 0.2 21.0 7.5 

FA98 STA 49.8 16.4 0.2 0.2 26.8 6.2 

SP99 STA 51.6 17.6 0.2 0.2 23.0 6.4 

FA99 STA 48.6 18.0 0.6 0.6 14.2 4.3 

SP00 STA 48.2 17.9 0.2 0.2 13.8 6.1 

FA00 STA 44.0 18.3 0.0 0.0 17.8 5.0 

SP01 STA 43.6 17.4 0.4 0.4 13.6 4.7 

FA01 STA 43.2 17.5 0.0 0.0 18.0 5.2 

SP02 STA 39.2 16.9 0.2 0.2 18.8 5.2 

FA02 STA 36.2 16.0 0.6 0.4 23.2 5.3 

SP03 STA 30.2 16.8 0.4 0.4 18.4 4.3 

FA03 STA 28.0 17.9 0.8 0.8 35.6 11.6 

 

SP92 BOA 77.0 3.9 7.2 1.4 12.6 4.0 

FA92 BOA 78.2 5.0 8.2 2.0 12.0 4.1 

SP93 BOA 75.4 3.8 7.2 1.5 10.8 2.8 

FA93 BOA 73.2 4.7 5.8 2.4 15.4 4.2 

SP94 BOA 72.0 4.6 6.4 1.6 13.2 4.8 

FA94 BOA 74.8 4.3 5.6 1.2 9.6 2.8 

SP95 BOA 67.0 10.6 6.8 1.6 20.6 10.7 

FA95 BOA 67.2 12.3 8.2 3.6 13.6 6.0 

SP96 BOA 61.8 14.1 8.2 2.3 24.0 13.3 

FA96 BOA 59.0 12.9 6.4 1.7 24.0 11.2 

SP97 BOA 62.4 14.0 4.8 1.6 26.0 11.3 

FA97 BOA 61.2 12.9 7.0 1.8 21.4 8.9 

SP98 BOA 62.8 13.1 4.8 1.1 20.8 9.4 
 C-29 

 Mytilus Pollicipes Rock 

Sample Site mean se mean se mean se 

FA98 BOA 58.2 12.7 5.6 1.2 23.4 9.4 

SP99 BOA 52.2 11.5 5.4 2.9 32.4 9.5 

FA99 BOA 56.0 13.8 5.0 1.8 26.2 11.5 

SP00 BOA 54.6 11.5 6.6 2.2 28.2 10.8 

FA00 BOA 54.0 12.2 8.0 1.1 23.0 8.8 

SP01 BOA 51.6 13.4 7.8 1.4 27.6 10.8 

FA01 BOA 57.2 12.5 6.6 1.5 24.0 9.8 

SP02 BOA 59.2 11.9 5.8 1.5 24.8 9.5 

FA02 BOA 61.8 12.3 7.4 1.5 18.2 8.9 

SP03 BOA 65.0 12.4 4.2 0.7 18.6 8.9 

FA03 BOA 71.0 12.9 4.4 1.0 16.6 10.8 

 

SP92 GPT 77.6 1.1 8.4 3.0 12.8 3.2 

FA92 GPT 82.2 1.4 8.4 2.9 8.0 2.4 

SP93 GPT 84.4 1.6 6.2 1.8 8.0 2.4 

FA93 GPT 89.0 0.3 3.6 1.6 6.8 1.3 

SP94 GPT 88.4 0.9 5.8 2.1 4.6 1.3 

FA94 GPT 90.8 1.8 4.4 1.4 2.4 0.8 

SP95 GPT 87.8 3.1 5.6 1.4 5.4 2.4 

FA95 GPT 91.0 2.1 5.6 1.5 2.8 1.4 

SP96 GPT 92.0 1.9 5.4 2.0 2.0 0.7 

FA96 GPT 91.6 2.4 4.4 1.9 3.8 1.9 

SP97 GPT 86.8 2.4 6.6 2.7 6.0 2.3 

FA97 GPT 73.4 3.0 6.6 1.7 15.4 2.1 

SP98 GPT 65.4 3.0 6.8 2.2 19.2 2.1 

FA98 GPT 66.8 5.3 5.0 1.3 22.6 4.0 

SP99 GPT 76.0 1.8 6.2 1.5 13.2 2.4 

FA99 GPT . . . . . . 

SP00 GPT 81.4 5.2 4.2 0.9 10.8 3.3 

FA00 GPT 80.0 4.9 4.0 0.9 12.8 3.2 

SP01 GPT 77.6 4.2 5.2 1.1 13.8 3.8 

FA01 GPT 77.6 5.6 5.2 1.8 13.4 3.7 

SP02 GPT 76.0 6.0 5.4 1.2 15.8 4.5 

FA02 GPT 71.6 5.8 5.4 2.4 16.6 4.6 

SP03 GPT 73.8 8.0 5.8 1.1 15.6 5.8 

FA03 GPT 76.4 5.6 7.4 2.2 11.6 4.4 

 

SP92 ALEG 76.8 2.8 0.0 0.0 3.6 1.3 

FA92 ALEG 78.8 1.8 0.0 0.0 1.4 0.7 

SP93 ALEG 74.0 1.9 0.0 0.0 6.8 1.6 

FA93 ALEG 79.4 3.9 0.0 0.0 4.6 1.6 

SP94 ALEG 68.4 7.4 0.0 0.0 6.0 1.7 

FA94 ALEG 70.4 8.5 0.0 0.0 4.6 1.3 

SP95 ALEG 57.0 7.5 0.0 0.0 9.2 2.3 

FA95 ALEG 60.8 9.7 0.0 0.0 10.6 3.3 
 C-30 

 Mytilus Pollicipes Rock 

Sample Site mean se mean se mean se 

SP96 ALEG 54.8 9.5 0.0 0.0 11.8 3.0 

FA96 ALEG 52.0 10.6 0.0 0.0 8.2 1.3 

SP97 ALEG 45.6 11.5 0.0 0.0 11.6 3.3 

FA97 ALEG 42.8 11.9 0.0 0.0 7.2 1.4 

SP98 ALEG 40.4 10.9 0.0 0.0 20.2 3.2 

FA98 ALEG 40.6 11.1 0.0 0.0 10.8 2.6 

SP99 ALEG 42.6 10.3 0.0 0.0 11.4 2.6 

FA99 ALEG 47.6 9.9 0.0 0.0 0.6 0.4 

SP00 ALEG 50.2 11.0 0.0 0.0 2.8 0.9 

FA00 ALEG 62.6 6.6 0.0 0.0 6.4 2.3 

SP01 ALEG 51.6 4.4 0.0 0.0 13.0 3.5 

FA01 ALEG 54.2 2.2 0.0 0.0 8.4 1.6 

SP02 ALEG 55.6 1.5 0.0 0.0 9.2 1.8 

FA02 ALEG 61.2 1.5 0.0 0.0 4.0 1.8 

SP03 ALEG 61.6 2.9 0.0 0.0 7.0 1.7 

FA03 ALEG 62.8 2.7 0.0 0.0 2.8 1.0 

 

SP92 ARHO 79.0 5.1 1.0 1.0 15.6 3.4 

FA92 ARHO 81.2 5.1 2.0 2.0 9.8 3.6 

SP93 ARHO 67.8 6.0 0.6 0.6 20.2 3.9 

FA93 ARHO 60.0 14.2 0.4 0.4 23.8 12.0 

SP94 ARHO 63.8 16.2 0.0 0.0 28.4 13.3 

FA94 ARHO 63.2 16.4 0.0 0.0 24.8 11.8 

SP95 ARHO 55.8 14.6 0.2 0.2 31.2 8.3 

FA95 ARHO 62.6 15.1 0.2 0.2 14.4 5.6 

SP96 ARHO 59.0 14.9 0.2 0.2 25.0 8.8 

FA96 ARHO 61.4 15.4 0.2 0.2 21.2 9.4 

SP97 ARHO 55.2 13.7 0.6 0.6 23.6 9.9 

FA97 ARHO 52.0 13.5 0.0 0.0 28.8 13.2 

SP98 ARHO 16.0 4.9 0.0 0.0 71.4 5.3 

FA98 ARHO 7.8 4.9 0.0 0.0 28.4 6.1 

SP99 ARHO 18.6 3.9 0.0 0.0 60.4 2.4 

FA99 ARHO 26.0 6.7 0.0 0.0 52.2 3.9 

SP00 ARHO 32.2 8.5 0.0 0.0 52.4 6.6 

FA00 ARHO 50.2 11.0 0.4 0.4 21.0 3.5 

SP01 ARHO 57.2 5.6 0.2 0.2 26.6 3.1 

FA01 ARHO 68.6 3.1 0.2 0.2 13.6 1.2 

SP02 ARHO 67.2 7.2 0.0 0.0 20.0 5.4 

FA02 ARHO 69.4 6.8 0.0 0.0 15.2 3.6 

SP03 ARHO 65.8 6.5 0.0 0.0 26.4 5.3 

FA03 ARHO 63.6 15.0 0.0 0.0 12.2 5.4 

 

SP03 COPT 79.8 4.4 0.0 0.0 14.6 4.2 
 C-31 

 Mytilus Pollicipes Rock 

Sample Site mean se mean se mean se 

SP92 CARP 83.4 4.5 0.0 0.0 7.2 3.1 

FA92 CARP 82.0 7.1 0.0 0.0 7.5 2.5 

SP93 CARP 87.8 6.6 0.0 0.0 1.3 1.3 

FA93 CARP 94.6 2.8 0.0 0.0 2.0 1.2 

SP94 CARP 89.8 9.5 0.0 0.0 0.0 0.0 

FA94 CARP 78.2 19.6 0.0 0.0 8.0 6.8 

SP96 CARP 34.4 11.0 0.0 0.0 9.4 3.8 

FA96 CARP 

SP97 CARP 11.4 5.8 0.0 0.0 22.0 10.7 

FA97 CARP 44.0 11.7 0.0 0.0 27.2 13.0 

SP98 CARP 19.2 4.5 0.0 0.0 31.8 13.9 

FA98 CARP 

SP99 CARP 77.8 7.3 0.0 0.0 3.2 1.3 

FA99 CARP 

SP00 CARP 46.8 19.3 0.0 0.0 6.2 2.2 

FA00 CARP 65.0 17.8 0.0 0.0 4.0 1.4 

SP01 CARP 53.0 15.5 0.0 0.0 10.4 4.4 

FA01 CARP . . . . . . 

SP02 CARP 56.0 15.1 0.0 0.0 5.8 3.8 

FA02 CARP . . . . . . 

SP03 CARP 52.0 5.0 0.0 0.0 9.4 2.9 

FA03 CARP 

 

FA94 MUSH 85.2 2.6 4.8 2.7 3.2 0.8 

SP95 MUSH 84.4 2.5 3.4 1.4 8.2 2.9 

FA95 MUSH 91.0 2.3 1.4 1.4 3.0 1.2 

SP96 MUSH 92.6 2.6 1.8 1.6 4.2 2.1 

FA96 MUSH 87.4 5.3 0.6 0.6 10.0 5.0 

SP97 MUSH 90.2 2.3 1.4 0.7 4.4 1.5 

FA97 MUSH 86.2 4.6 1.0 0.5 8.4 4.0 

SP98 MUSH 78.4 5.9 0.0 0.0 17.4 5.1 

FA98 MUSH 87.0 5.2 0.4 0.4 9.8 4.7 

SP99 MUSH 89.6 3.5 0.4 0.4 7.4 3.1 

FA99 MUSH 88.4 3.5 0.0 0.0 8.8 2.7 

SP00 MUSH 88.8 4.1 1.6 1.6 7.0 2.2 

FA00 MUSH 86.2 4.2 0.6 0.6 6.4 2.2 

SP01 MUSH 84.8 4.7 0.2 0.2 10.2 3.9 

FA01 MUSH 78.8 7.8 0.4 0.4 10.6 4.1 

SP02 MUSH 77.8 6.4 0.8 0.8 15.0 5.0 

FA02 MUSH 71.0 8.1 1.0 1.0 16.4 4.5 

SP03 MUSH 76.0 6.3 1.0 0.8 17.2 5.2 

FA03 MUSH 79.2 7.1 0.2 0.2 10.2 4.0 

 

FA94 OLDS 81.4 3.0 1.8 0.9 7.4 2.0 

SP95 OLDS 77.8 7.0 1.8 1.0 10.4 6.1 
 C-32 

 Mytilus Pollicipes Rock 

Sample Site mean se mean se mean se 

FA95 OLDS 33.6 11.5 3.0 1.8 34.2 8.7 

SP96 OLDS 30.8 7.0 1.6 1.0 25.4 5.3 

FA96 OLDS 40.0 8.1 3.6 1.2 15.0 7.4 

SP97 OLDS 28.0 7.6 3.2 0.7 30.4 4.3 

FA97 OLDS 40.8 6.0 0.8 0.6 22.2 7.0 

SP98 OLDS 45.6 5.7 2.2 0.5 28.6 6.0 

FA98 OLDS 41.4 6.8 3.8 0.9 32.2 7.9 

SP99 OLDS 36.8 6.4 1.4 0.7 34.2 8.8 

FA99 OLDS 57.2 8.6 3.0 0.6 21.8 7.9 

SP00 OLDS 53.0 6.4 4.6 0.5 25.2 5.7 

FA00 OLDS 55.0 8.0 1.0 0.3 20.6 5.4 

SP01 OLDS 52.2 8.9 1.8 0.9 18.2 4.2 

FA01 OLDS 54.2 10.1 2.4 0.9 17.4 3.9 

SP02 OLDS 47.8 8.6 1.8 0.6 26.6 3.0 

FA02 OLDS 53.2 6.3 0.8 0.5 20.2 5.6 

SP03 OLDS 51.8 5.4 1.6 0.7 24.2 6.6 

FA03 OLDS 63.6 4.8 3.6 1.5 6.4 2.8 

 

FA94 PCOV 91.8 2.7 0.8 0.8 4.8 1.0 

SP95 PCOV 90.0 2.5 1.4 1.2 5.2 1.1 

FA95 PCOV 96.4 2.3 0.8 0.8 2.4 1.3 

SP96 PCOV 94.8 1.5 1.0 1.0 2.4 1.5 

FA96 PCOV 92.8 2.0 0.6 0.6 1.6 1.6 

SP97 PCOV 82.4 10.8 0.0 0.0 12.4 10.2 

FA97 PCOV 81.4 9.7 0.0 0.0 11.8 7.2 

SP98 PCOV 73.0 8.6 0.0 0.0 21.0 7.0 

FA98 PCOV 79.4 9.3 0.0 0.0 7.8 5.5 

SP99 PCOV 82.0 6.5 0.0 0.0 13.0 5.9 

FA99 PCOV 83.4 5.4 0.0 0.0 13.2 5.2 

SP00 PCOV 82.4 5.4 0.0 0.0 14.4 5.2 

FA00 PCOV 78.8 7.8 0.4 0.2 16.0 6.2 

SP01 PCOV 78.6 6.1 0.4 0.2 15.4 5.6 

FA01 PCOV 82.8 5.6 0.2 0.2 11.6 4.8 

SP02 PCOV 85.0 4.7 0.0 0.0 11.8 4.9 

FA02 PCOV 89.0 4.0 0.0 0.0 9.0 3.8 

SP03 PCOV 87.8 4.5 0.0 0.0 9.6 3.7 

FA03 PCOV 88.6 3.5 0.2 0.2 8.6 2.9 

 

FA94 WHPT 93.4 1.6 0.0 0.0 3.4 1.6 

SP95 WHPT 94.6 2.1 1.2 0.5 2.0 1.0 

FA95 WHPT 95.0 2.1 0.0 0.0 3.6 1.6 

SP96 WHPT 81.0 15.8 1.0 0.8 6.0 5.0 

FA96 WHPT 76.6 11.5 0.0 0.0 9.8 4.1 

SP97 WHPT 79.2 10.3 0.4 0.2 8.4 2.8 

FA97 WHPT 80.6 10.3 0.0 0.0 5.6 3.5 
 C-33 

 Mytilus Pollicipes Rock 

Sample Site mean se mean se mean se 

SP98 WHPT 81.8 10.0 0.4 0.4 5.8 2.7 

FA98 WHPT 88.2 7.3 0.2 0.2 8.6 5.4 

SP99 WHPT 87.6 7.2 1.0 0.4 9.2 6.6 

FA99 WHPT 82.8 10.3 0.4 0.2 8.0 5.2 

SP00 WHPT 91.8 3.4 1.2 1.0 2.6 1.1 

FA00 WHPT 95.0 2.2 0.0 0.0 2.0 0.8 

SP01 WHPT 94.6 1.9 0.0 0.0 1.4 0.4 

FA01 WHPT 90.2 5.6 0.0 0.0 4.8 2.9 

SP02 WHPT 88.6 5.2 0.0 0.0 3.6 1.4 

FA02 WHPT 83.2 5.2 0.2 0.2 3.8 1.8 

SP03 WHPT 84.2 4.7 0.0 0.0 4.0 1.9 

FA03 WHPT 85.6 5.6 0.0 0.0 2.4 1.1 

 

FA99 PTFM 93.6 2.2 0.0 0.0 5.0 1.5 

SP00 PTFM 78.0 3.4 0.0 0.0 17.8 2.7 

FA00 PTFM 51.6 6.6 0.0 0.0 25.0 2.3 

SP01 PTFM 40.0 4.9 0.0 0.0 37.8 3.8 

FA01 PTFM 37.0 4.5 0.0 0.0 22.8 3.0 

SP02 PTFM 32.8 3.6 0.0 0.0 37.4 3.5 

FA02 PTFM 25.2 6.2 0.0 0.0 25.8 3.8 

SP03 PTFM 19.6 3.9 0.0 0.0 35.0 2.5 

FA03 PTFM 12.0 7.4 0.0 0.0 28.8 1.5 

 

FA96 CRCO 94.0 2.5 0.0 0.0 3.0 1.7 

SP97 CRCO 96.8 2.2 0.0 0.0 1.6 1.2 

FA97 CRCO 92.0 3.3 0.0 0.0 5.0 2.2 

SP98 CRCO 48.0 18.6 0.0 0.0 12.2 5.4 

FA98 CRCO 70.6 10.3 0.0 0.0 6.4 3.0 

SP99 CRCO 74.4 9.6 0.2 0.2 9.2 1.9 

FA99 CRCO 90.4 6.4 0.4 0.2 2.6 1.9 

SP00 CRCO 94.0 5.0 0.0 0.0 0.0 0.0 

FA00 CRCO 95.2 3.9 0.0 0.0 0.2 0.2 

SP01 CRCO 98.2 1.8 0.0 0.0 0.4 0.4 

FA01 CRCO 99.2 0.5 0.0 0.0 0.0 0.0 

SP02 CRCO 99.4 0.4 0.0 0.0 0.4 0.2 

FA02 CRCO 76.6 13.2 0.0 0.0 3.0 1.1 

SP03 CRCO 74.6 13.5 0.0 0.0 3.4 1.4 

FA03 CRCO 82.8 12.1 0.0 0.0 0.6 0.4 

 

FA96 SHCO 79.8 2.6 1.6 1.0 9.2 1.2 

SP97 SHCO 80.6 2.3 2.4 1.0 11.4 2.2 

FA97 SHCO 78.0 4.0 0.6 0.4 6.2 1.9 

SP98 SHCO 51.2 3.8 0.8 0.6 33.2 4.1 

FA98 SHCO 58.0 2.2 0.0 0.0 22.8 5.7 

SP99 SHCO 61.0 2.6 0.2 0.2 3.2 1.8 
 C-34 

 Mytilus Pollicipes Rock 

Sample Site mean se mean se mean se 

FA99 SHCO 55.0 6.3 2.0 0.4 15.8 3.2 

SP00 SHCO 57.8 2.4 3.2 0.7 26.4 2.5 

FA00 SHCO 58.0 5.4 1.6 0.4 23.6 3.8 

SP01 SHCO 64.8 5.7 2.4 1.0 20.6 4.1 

FA01 SHCO 65.6 5.1 2.2 0.9 12.0 2.0 

SP02 SHCO 67.4 6.2 1.4 0.9 22.6 5.1 

FA02 SHCO 64.2 7.4 3.4 1.2 12.2 3.4 

SP03 SHCO 63.2 7.5 3.4 0.7 18.8 5.0 

FA03 SHCO 82.8 3.7 3.4 0.9 3.8 1.2 

 

FA96 TRIS 67.2 3.8 0.6 0.4 9.6 2.1 

SP97 TRIS 56.8 7.3 0.0 0.0 29.4 4.8 

FA97 TRIS 39.8 8.9 0.0 0.0 7.8 3.4 

SP98 TRIS 22.2 7.0 0.2 0.2 65.2 7.4 

FA98 TRIS 22.8 5.8 0.0 0.0 67.6 5.7 

SP99 TRIS 22.6 4.0 0.4 0.4 69.2 3.9 

FA99 TRIS 29.8 3.4 3.4 1.5 9.8 2.7 

SP00 TRIS 26.0 3.8 4.4 1.4 40.2 4.9 

FA00 TRIS 24.6 1.4 2.8 1.0 50.8 4.1 

SP01 TRIS 30.4 2.6 4.2 1.4 39.4 4.6 

FA01 TRIS 26.4 3.4 3.0 1.4 34.2 6.5 

SP02 TRIS 32.4 2.1 4.8 1.6 17.0 1.5 

FA02 TRIS 31.0 1.7 4.0 2.3 24.8 4.2 

SP03 TRIS 34.2 3.9 3.0 0.9 28.6 5.2 

FA03 TRIS 62.8 5.5 5.0 2.2 13.0 3.8 

 

FA96 DAPT 96.6 1.0 0.0 0.0 1.8 0.6 

SP97 DAPT 95.4 2.9 0.0 0.0 3.0 2.3 

FA97 DAPT 84.8 4.3 0.0 0.0 8.0 2.6 

SP98 DAPT 59.6 6.4 0.0 0.0 27.8 5.9 

FA98 DAPT 62.4 8.9 0.0 0.0 19.4 2.9 

SP99 DAPT 67.0 5.7 0.0 0.0 7.2 3.7 

FA99 DAPT 66.6 6.3 0.0 0.0 18.8 5.0 

SP00 DAPT 70.0 6.8 1.0 0.6 15.0 3.3 

FA00 DAPT 71.6 4.3 0.0 0.0 21.6 2.3 

SP01 DAPT 69.4 3.0 0.0 0.0 22.0 3.0 

FA01 DAPT 77.2 2.3 0.0 0.0 9.2 1.3 

SP02 DAPT 73.8 2.7 0.0 0.0 16.6 2.3 

FA02 DAPT 79.6 2.8 0.0 0.0 12.0 2.0 

SP03 DAPT 81.2 2.6 0.4 0.2 12.4 1.6 

FA03 DAPT 92.8 1.9 0.2 0.2 4.2 1.1 

 
 C-35 

 Mytilus Pollicipes Rock 

Sample Site mean se mean se mean se 

FA94 CTBR 8.8 3.1 0.2 0.2 1.0 0.5 

SP95 CTBR 32.8 8.3 1.8 0.9 0.8 0.5 

FA95 CTBR 52.8 6.2 3.6 0.7 2.0 1.0 

SP96 CTBR 61.2 6.3 4.4 1.9 0.4 0.4 

FA96 CTBR 50.8 6.2 3.2 1.2 2.8 0.8 

SP97 CTBR 54.0 6.8 9.6 3.4 2.2 0.8 

FA97 CTBR 39.0 10.4 4.0 1.6 1.0 0.6 

SP98 CTBR 24.2 6.0 5.2 1.6 2.2 0.6 

FA98 CTBR 13.4 7.5 1.8 1.2 6.6 2.1 

SP99 CTBR 11.8 3.5 7.4 4.0 0.6 0.2 

FA99 CTBR 10.8 6.2 3.0 2.5 1.8 0.7 

SP00 CTBR 12.8 3.3 8.4 3.5 1.0 0.4 

FA00 CTBR 15.2 5.1 5.4 2.7 1.6 0.5 

SP01 CTBR 27.0 9.2 3.6 2.7 2.6 0.9 

FA01 CTBR 11.6 4.8 2.6 2.1 2.6 0.8 

SP02 CTBR 28.0 4.9 4.2 1.4 2.4 0.9 

FA02 CTBR 12.4 3.9 0.8 0.5 2.4 1.7 

SP03 CTBR 18.6 3.3 2.2 0.7 2.2 1.0 

 

FA94 CTLH 21.0 4.1 0.2 0.2 12.6 3.3 

SP95 CTLH 17.2 2.2 0.6 0.2 7.0 1.8 

FA95 CTLH 37.2 6.8 0.4 0.4 5.4 1.7 

SP96 CTLH 38.6 4.5 0.6 0.4 10.4 2.2 

FA96 CTLH 39.6 4.2 0.0 0.0 6.0 2.3 

SP97 CTLH 38.8 5.0 0.4 0.4 12.0 3.8 

FA97 CTLH 13.4 2.9 0.2 0.2 8.6 4.4 

SP98 CTLH 19.4 1.9 0.0 0.0 14.4 5.5 

FA98 CTLH 23.6 2.1 0.2 0.2 0.0 0.0 

SP99 CTLH 24.0 3.6 0.4 0.4 0.8 0.8 

FA99 CTLH 14.6 4.9 0.0 0.0 7.0 2.1 

SP00 CTLH 19.0 7.4 1.4 0.9 10.4 1.2 

FA00 CTLH 15.6 7.1 0.2 0.2 4.8 1.7 

SP01 CTLH 17.6 8.0 0.2 0.2 6.8 3.0 

FA01 CTLH . . . . . . 

SP02 CTLH 16.4 7.3 0.2 0.2 9.4 3.5 

FA02 CTLH 19.8 9.2 0.0 0.0 4.6 1.9 

SP03 CTLH 18.0 8.2 0.0 0.0 15.6 3.5 

 
 C-36 

Recovery  Barnacles Other Reds Endocladia Cladophora Silvetia Ulva Non-Sample Site mean se mean se mean se mean se mean se mean se meaSP98 STA 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 6.1 4.0 0.1 FA98 STA 0.1 0.1 1.9 1.9 0.0 0.0 10.4 5.5 0.0 0.0 0.0 0.0 18.7 SP99 STA 0.0 0.0 0.7 0.4 0.0 0.0 0.0 0.0 0.0 0.0 4.4 3.3 22.9 FA99 STA 1.6 0.8 1.6 1.3 0.0 0.0 0.0 0.0 1.0 0.7 0.0 0.0 18.9 SP00 STA 0.1 0.1 0.0 0.0 0.3 0.2 0.0 0.0 0.3 0.2 0.0 0.0 29.9 FA00 STA 6.1 4.4 8.7 4.4 1.7 1.3 0.0 0.0 1.9 1.5 3.9 1.7 15.4 SP01 STA 7.6 6.6 2.1 1.3 2.9 2.1 3.3 1.8 1.1 0.7 1.0 1.0 23.1 FA01 STA 4.9 3.7 5.9 2.6 1.4 0.6 0.6 0.4 5.9 2.8 8.0 2.8 22.3 SP02 STA 4.0 3.5 3.0 2.2 7.9 3.5 1.0 1.0 1.7 1.4 8.1 5.1 23.9 FA02 STA 5.0 3.0 2.7 0.9 5.1 3.1 2.1 0.8 8.7 5.2 5.3 3.2 28.6 SP03 STA 1.3 1.0 4.7 3.1 6.6 4.0 0.3 0.2 2.9 1.5 19.1 2.9 32.7 FA03 STA 3.0 1.5 2.6 1.2 2.4 1.3 0.7 0.2 8.7 3.6 2.9 0.9 21.9 
 C-37 

Phyllospadix 

 Phyllospadix Egregia Other algae Sand 

Sample Site mean se mean se mean se mean se 

FA95 PSN 83.7 7.2 0.7 0.7 14.3 6.2 1.3 1.3 

SP96 PSN 70.7 8.4 0.0 0.0 24.7 10.3 1.0 0.6 

FA96 PSN 81.3 7.2 2.7 2.7 14.3 4.8 1.0 0.6 

SP97 PSN 68.3 8.7 0.7 0.7 24.7 10.7 1.0 0.6 

FA97 PSN 85.3 7.8 0.7 0.7 8.0 2.5 0.0 0.0 

SP98 PSN 78.0 4.5 0.0 0.0 14.0 7.2 1.0 0.6 

FA98 PSN 79.3 12.8 0.0 0.0 18.3 12.5 0.7 0.7 

SP99 PSN 78.3 6.7 0.0 0.0 17.3 7.4 1.7 1.2 

FA99 PSN 77.3 5.2 0.0 0.0 15.0 8.0 0.7 0.7 

SP00 PSN 76.7 5.8 0.0 0.0 18.3 4.7 1.3 0.9 

FA00 PSN 86.7 7.4 3.7 3.7 9.3 3.9 0.0 0.0 

SP01 PSN 78.7 5.6 0.7 0.7 17.3 6.2 0.0 0.0 

FA01 PSN 82.0 6.0 5.7 5.2 10.7 4.7 1.7 0.9 

SP02 PSN 72.7 9.9 2.7 2.7 21.0 9.6 1.0 0.6 

FA02 PSN 88.0 4.0 11.0 7.0 10.0 3.5 0.3 0.3 

SP03 PSN 74.7 7.1 4.3 4.3 14.3 3.9 1.3 1.3 

FA03 PSN 84.7 7.9 16.0 16.0 11.3 5.3 0.3 0.3 

 

FA95 CAY 90.3 1.2 0.0 0.0 3.3 0.3 0.7 0.7 

SP96 CAY 92.3 2.7 0.0 0.0 5.0 0.0 0.0 0.0 

FA96 CAY 87.7 3.8 0.0 0.0 9.3 2.3 0.0 0.0 

SP97 CAY 91.7 3.0 0.0 0.0 7.3 3.3 0.0 0.0 

FA97 CAY 74.0 14.2 0.0 0.0 11.0 2.3 0.3 0.3 

SP98 CAY 83.0 4.6 0.0 0.0 10.3 1.8 0.0 0.0 

FA98 CAY 89.7 3.8 0.0 0.0 4.0 0.6 0.0 0.0 

SP99 CAY 94.0 1.0 0.0 0.0 2.3 1.5 0.0 0.0 

FA99 CAY 90.0 1.5 0.0 0.0 5.7 0.9 1.7 0.9 

SP00 CAY 92.7 0.7 0.0 0.0 3.7 1.9 1.0 1.0 

FA00 CAY 86.7 1.5 0.0 0.0 8.3 1.2 0.0 0.0 

SP01 CAY 85.3 2.9 0.0 0.0 9.0 4.2 0.0 0.0 

FA01 CAY 72.7 8.4 0.0 0.0 10.3 3.8 1.3 0.7 

SP02 CAY 85.0 5.5 0.0 0.0 11.3 5.8 0.0 0.0 

FA02 CAY 84.0 1.2 0.0 0.0 8.7 1.5 0.0 0.0 

SP03 CAY 81.7 2.9 0.0 0.0 8.3 3.7 0.0 0.0 

FA03 CAY 88.0 4.2 0.0 0.0 9.0 4.5 0.0 0.0 

 

FA01 HAZ 89.7 4.1 0.3 0.3 2.7 0.9 0.0 0.0 

SP02 HAZ . . . . . . . . 

FA02 HAZ 93.0 . 1.0 . 6.0 . 0.0 . 

SP03 HAZ 73.0 . 0.0 . 15.0 . 1.0 . 

FA03 HAZ 85.7 7.5 1.7 0.9 7.0 6.5 0.0 0.0 

 

FA99 SHB 93.3 1.7 0.0 0.0 4.7 1.5 0.0 0.0 

SP00 SHB 75.3 2.8 0.0 0.0 15.7 0.7 0.0 0.0 

FA00 SHB 89.3 4.8 2.3 1.9 5.3 3.9 0.0 0.0 
 C-38 

 Phyllospadix Egregia Other algae Sand 

Sample Site mean se mean se mean se mean se 

SP01 SHB 77.3 4.8 0.0 0.0 7.3 1.5 0.0 0.0 

FA01 SHB 83.5 5.5 0.0 0.0 10.0 3.0 0.0 0.0 

SP02 SHB 77.0 3.5 0.0 0.0 12.0 2.6 0.0 0.0 

FA02 SHB 70.3 5.8 0.0 0.0 20.7 6.4 0.0 0.0 

SP03 SHB 55.7 11.6 0.0 0.0 26.0 8.7 0.0 0.0 

FA03 SHB 69.0 6.4 0.0 0.0 28.0 6.8 0.0 0.0 

 

SP92 STA 77.3 3.3 0.0 0.0 13.7 2.3 4.0 1.5 

FA92 STA 90.3 0.9 0.0 0.0 7.0 1.2 0.0 0.0 

SP93 STA 74.7 2.7 0.0 0.0 12.7 3.7 8.7 3.8 

FA93 STA 94.7 1.8 0.0 0.0 3.3 1.3 1.0 0.6 

SP94 STA 89.3 2.4 0.0 0.0 5.7 1.8 0.3 0.3 

FA94 STA 98.7 0.9 0.0 0.0 0.7 0.7 0.7 0.3 

SP95 STA 77.0 11.5 0.0 0.0 8.3 3.8 0.7 0.7 

FA95 STA 80.3 8.5 0.0 0.0 13.3 6.2 0.3 0.3 

SP96 STA 67.3 9.8 0.0 0.0 15.7 5.8 0.0 0.0 

FA96 STA 83.7 4.8 0.0 0.0 7.0 1.2 0.3 0.3 

SP97 STA 71.7 3.3 0.0 0.0 17.0 6.0 0.0 0.0 

FA97 STA 85.7 4.3 0.0 0.0 8.0 1.5 0.3 0.3 

SP98 STA 7.0 3.0 0.0 0.0 3.3 3.3 0.0 0.0 

FA98 STA 11.5 8.5 0.0 0.0 43.0 34.0 1.0 0.0 

SP99 STA 21.7 8.4 0.0 0.0 42.7 18.4 1.0 0.6 

FA99 STA 33.7 13.4 0.0 0.0 42.3 18.5 0.0 0.0 

SP00 STA 32.3 15.0 0.0 0.0 35.7 19.4 0.0 0.0 

FA00 STA 44.7 12.0 0.0 0.0 34.3 8.1 1.0 1.0 

SP01 STA 35.3 15.3 0.0 0.0 21.7 10.8 0.0 0.0 

FA01 STA 40.7 18.8 0.7 0.7 39.0 20.1 0.0 0.0 

SP02 STA 39.7 18.0 0.0 0.0 43.0 19.5 0.0 0.0 

FA02 STA 56.0 21.6 0.0 0.0 35.7 20.9 0.7 0.3 

SP03 STA 37.7 18.0 0.0 0.0 47.7 16.2 0.7 0.7 

FA03 STA 48.0 21.4 1.0 1.0 37.3 20.8 0.0 0.0 

 

SP92 GPT 72.7 12.2 0.7 0.7 0.7 0.7 24.0 11.5 

FA92 GPT 93.7 3.4 2.0 2.0 4.3 1.5 0.0 0.0 

SP93 GPT 67.7 10.1 0.7 0.3 8.0 1.7 21.0 9.5 

FA93 GPT 98.7 0.9 0.0 0.0 0.3 0.3 1.0 0.6 

SP94 GPT 95.3 1.2 0.0 0.0 1.7 0.9 2.3 1.5 

FA94 GPT 98.3 0.9 1.0 1.0 0.0 0.0 0.7 0.7 

SP95 GPT 81.7 8.4 0.0 0.0 3.0 1.5 4.7 2.7 

FA95 GPT 99.0 0.6 0.0 0.0 0.0 0.0 1.0 0.6 

SP96 GPT 86.0 6.4 0.0 0.0 1.7 1.7 2.7 2.2 

FA96 GPT 97.7 2.3 2.3 2.3 0.0 0.0 0.0 0.0 

SP97 GPT 88.0 4.0 0.0 0.0 1.0 0.0 1.7 0.9 

FA97 GPT 98.7 0.7 0.0 0.0 0.0 0.0 0.0 0.0 

SP98 GPT 73.0 4.7 12.0 3.6 12.3 4.4 0.0 0.0 
 C-39 

 Phyllospadix Egregia Other algae Sand 

Sample Site mean se mean se mean se mean se 

FA98 GPT 74.3 18.9 19.7 19.7 1.7 0.3 4.3 3.0 

SP99 GPT 79.0 12.1 15.3 12.9 2.3 1.9 1.0 0.6 

SP00 GPT 94.7 2.4 2.0 1.2 1.7 1.2 1.7 0.9 

FA00 GPT 99.0 0.6 0.7 0.7 0.3 0.3 0.0 0.0 

SP01 GPT 97.3 1.2 0.0 0.0 0.7 0.3 1.0 1.0 

FA01 GPT 95.0 3.2 0.0 0.0 0.0 0.0 5.0 3.2 

SP02 GPT 91.7 1.9 1.3 1.3 2.7 0.7 5.7 1.2 

FA02 GPT 98.3 1.7 1.7 1.7 0.7 0.7 1.0 1.0 

SP03 GPT 91.0 4.0 0.0 0.0 3.0 2.5 5.0 2.6 

FA03 GPT 100.0 0.0 1.0 1.0 0.0 0.0 0.0 0.0 

 

FA01 ALEG 83.3 8.6 3.0 3.0 8.3 5.4 0.3 0.3 

SP02 ALEG 80.7 12.4 1.3 0.9 14.0 10.0 0.3 0.3 

FA02 ALEG 85.3 9.5 1.7 0.3 10.3 7.4 0.0 0.0 

SP03 ALEG 70.0 9.3 0.3 0.3 19.0 7.2 5.0 2.1 

FA03 ALEG 78.3 8.5 24.3 12.5 8.3 5.4 1.3 0.9 

 

SP92 ARHO 77.0 12.0 0.0 0.0 8.5 2.5 4.0 4.0 

FA92 ARHO 94.0 3.0 0.0 0.0 4.5 1.5 0.0 0.0 

SP93 ARHO 89.5 7.5 0.0 0.0 3.0 1.0 3.5 3.5 

FA93 ARHO 88.5 3.5 0.0 0.0 3.0 0.0 7.0 2.0 

SP94 ARHO 73.5 15.5 0.0 0.0 14.0 5.0 9.5 8.5 

FA94 ARHO 87.5 4.5 0.0 0.0 5.5 0.5 5.5 4.5 

SP95 ARHO 46.5 5.5 0.0 0.0 11.0 3.0 14.5 5.5 

FA95 ARHO 88.0 1.0 0.0 0.0 6.5 3.5 3.5 1.5 

SP96 ARHO 88.5 3.5 0.0 0.0 8.5 4.5 2.5 0.5 

FA96 ARHO 86.0 1.0 0.0 0.0 11.5 0.5 1.0 0.0 

SP97 ARHO 64.5 0.5 0.0 0.0 21.0 9.0 7.0 7.0 

FA97 ARHO 71.0 1.0 0.0 0.0 20.5 2.5 3.0 2.0 

SP98 ARHO 34.5 9.5 0.0 0.0 52.5 8.5 2.5 2.5 

FA98 ARHO 75.0 1.0 0.5 0.5 22.5 1.5 1.5 0.5 

SP99 ARHO 64.0 2.0 1.0 1.0 23.5 5.5 7.0 4.0 

FA99 ARHO 84.0 5.0 0.0 0.0 7.0 1.0 2.5 0.5 

SP00 ARHO 75.0 4.0 0.0 0.0 17.0 2.0 7.0 5.0 

FA00 ARHO 93.5 4.5 0.0 0.0 4.5 4.5 1.0 1.0 

SP01 ARHO 80.0 1.5 0.7 0.7 14.7 0.9 0.7 0.3 

FA01 ARHO 87.0 5.2 0.0 0.0 9.7 4.3 1.3 0.7 

SP02 ARHO 78.0 3.2 0.0 0.0 16.0 1.0 1.3 1.3 

FA02 ARHO 62.3 2.8 0.0 0.0 28.3 1.2 3.7 1.2 

SP03 ARHO 54.0 2.1 0.0 0.0 30.3 5.2 4.3 2.0 

FA03 ARHO 58.7 7.7 2.3 1.5 28.3 2.6 6.3 1.8 

 

SP92 COPT 61.7 8.7 1.0 1.0 16.7 4.4 13.3 2.6 

FA92 COPT 81.7 5.2 0.0 0.0 5.3 1.9 11.0 3.5 

SP93 COPT 61.3 9.8 0.3 0.3 17.3 3.8 15.0 4.0 

FA93 COPT 78.0 3.2 0.0 0.0 11.0 5.8 4.7 0.3 
 C-40 

 Phyllospadix Egregia Other algae Sand 

Sample Site mean se mean se mean se mean se 

SP94 COPT 61.3 6.4 0.0 0.0 9.0 2.3 26.0 5.2 

FA94 COPT 0.0 0.0 0.0 0.0 0.0 0.0 100.0 0.0 

SP95 COPT 49.3 4.3 0.0 0.0 17.3 8.4 21.0 5.9 

FA95 COPT 60.3 11.5 10.3 8.0 21.3 7.8 2.0 0.0 

SP96 COPT 54.3 9.2 5.0 3.2 15.3 4.3 21.7 4.7 

FA96 COPT 84.0 7.5 0.0 0.0 4.7 2.6 10.3 4.3 

SP97 COPT 52.3 7.8 0.0 0.0 9.7 7.2 33.0 9.0 

FA97 COPT 59.0 6.6 0.0 0.0 7.3 5.4 30.7 10.7 

SP98 COPT 45.3 9.7 0.0 0.0 42.7 9.7 5.3 1.5 

FA98 COPT 70.7 8.4 0.0 0.0 11.7 4.5 12.0 4.5 

SP99 COPT 23.7 9.6 0.3 0.3 0.3 0.3 75.7 9.9 

FA99 COPT 67.0 7.2 1.0 1.0 14.7 4.8 13.3 3.8 

SP00 COPT 45.3 8.3 0.0 0.0 14.7 4.9 34.0 12.3 

FA00 COPT 62.3 11.9 0.3 0.3 12.3 6.4 20.3 8.1 

SP01 COPT 50.0 4.5 0.0 0.0 3.3 1.5 43.0 3.5 

FA01 COPT 86.7 7.1 0.0 0.0 7.0 3.6 3.0 1.2 

SP02 COPT 48.7 6.6 0.0 0.0 4.0 2.6 41.7 10.2 

FA02 COPT 41.3 14.9 0.0 0.0 0.3 0.3 57.3 14.5 

SP03 COPT 49.7 8.9 0.0 0.0 31.0 3.5 15.3 7.4 

FA03 COPT 64.3 9.9 0.0 0.0 23.7 8.2 5.0 2.1 

 

SP92 CARP 78.7 1.2 1.0 0.6 7.7 2.4 7.7 2.3 

FA92 CARP 87.7 0.7 5.3 2.7 6.0 2.0 0.0 0.0 

SP93 CARP 70.3 4.3 0.0 0.0 10.3 3.5 18.7 0.9 

FA93 CARP 90.3 1.7 5.0 1.7 3.0 2.1 1.3 0.9 

SP94 CARP 84.0 2.5 1.7 0.9 2.3 1.2 11.3 1.2 

FA94 CARP 79.0 5.6 1.7 1.7 1.7 0.9 16.3 4.3 

SP95 CARP 63.0 4.0 0.0 0.0 24.0 4.2 3.7 2.2 

FA95 CARP 66.7 10.9 16.7 6.4 16.0 4.5 0.3 0.3 

SP96 CARP 57.0 2.5 0.7 0.3 36.0 4.6 3.3 1.8 

FA96 CARP 72.3 5.0 1.3 1.3 4.0 2.3 21.7 6.9 

SP97 CARP 61.3 5.8 0.0 0.0 22.3 1.5 14.7 3.8 

FA97 CARP 55.7 5.4 0.0 0.0 21.7 3.8 1.7 0.7 

SP98 CARP 41.7 9.9 0.0 0.0 50.0 10.5 6.3 1.2 

FA98 CARP 9.7 1.7 82.7 3.7 6.7 2.6 0.3 0.3 

SP99 CARP 11.3 6.9 41.7 14.2 29.3 4.9 13.3 6.4 

FA99 CARP 21.0 11.5 66.3 10.9 10.3 1.9 1.7 0.7 

SP00 CARP 16.7 5.2 33.7 4.4 41.0 4.5 3.3 1.3 

FA00 CARP 39.7 3.2 29.7 3.8 16.0 1.5 11.7 3.2 

SP01 CARP 43.3 3.8 17.7 6.7 29.7 0.9 5.3 2.8 

FA01 CARP 52.7 1.7 23.3 4.4 17.0 4.6 6.3 4.4 

SP02 CARP 46.3 5.2 13.0 2.6 35.3 5.6 4.0 1.2 

FA02 CARP 51.0 1.0 11.5 5.5 28.5 2.5 4.5 1.5 

SP03 CARP 44.7 5.4 8.3 3.0 34.3 2.3 9.3 3.8 

FA03 CARP 46.0 4.0 5.5 1.5 33.5 4.5 11.0 2.0 
 C-41 

 Phyllospadix Egregia Other algae Sand 

Sample Site mean se mean se mean se mean se 

FA94 MUSH 72.7 5.0 4.3 2.2 7.0 3.6 12.7 1.2 

SP95 MUSH 68.0 8.3 0.0 0.0 6.3 3.2 0.0 0.0 

FA95 MUSH 81.3 3.2 5.0 2.1 1.7 0.9 2.3 1.5 

SP96 MUSH 72.0 7.2 3.0 1.2 4.0 0.6 0.0 0.0 

FA96 MUSH 51.7 6.7 24.7 2.8 10.0 1.0 0.7 0.7 

SP97 MUSH 46.7 11.2 11.3 6.4 8.7 3.5 6.3 2.3 

FA97 MUSH 60.0 3.6 5.3 2.6 2.7 1.2 14.0 5.6 

SP98 MUSH 37.3 13.9 0.3 0.3 8.7 4.2 0.0 0.0 

FA98 MUSH 43.0 10.0 4.7 3.7 32.0 5.5 5.3 1.3 

SP99 MUSH 42.0 4.4 6.3 3.5 37.3 6.2 2.0 2.0 

FA99 MUSH 38.7 7.2 27.3 14.3 9.3 4.9 13.7 5.8 

SP00 MUSH 39.7 3.5 16.7 0.9 23.7 3.0 2.7 1.7 

FA00 MUSH 43.7 10.4 30.0 11.9 7.3 4.9 17.3 7.3 

SP01 MUSH 38.7 13.4 19.7 7.4 24.3 2.7 0.3 0.3 

FA01 MUSH 51.3 15.0 19.7 10.1 17.7 2.0 3.7 1.8 

SP02 MUSH 44.0 17.0 26.0 8.7 28.0 8.1 0.0 0.0 

FA02 MUSH 52.3 15.2 20.7 8.8 23.0 6.6 7.3 0.3 

SP03 MUSH 45.0 15.7 8.0 6.0 37.0 9.3 0.7 0.7 

FA03 MUSH 47.7 15.5 26.0 13.0 16.7 6.5 11.0 0.0 

 

FA94 PCOV 80.3 2.3 0.0 0.0 11.3 1.5 4.7 2.4 

SP95 PCOV 58.7 7.6 1.0 0.6 9.7 3.3 14.3 6.1 

FA95 PCOV 77.0 4.5 2.0 1.2 4.3 2.0 1.7 0.9 

SP96 PCOV 61.0 8.3 5.3 2.7 5.3 2.7 20.0 9.5 

FA96 PCOV 79.0 4.4 4.0 2.0 2.3 1.9 4.7 0.3 

SP97 PCOV 52.3 9.4 0.3 0.3 6.0 2.5 20.0 8.5 

FA97 PCOV 30.7 5.7 8.3 2.3 29.3 5.0 29.3 6.4 

SP98 PCOV 15.0 5.0 5.0 2.6 12.7 5.2 3.7 1.5 

FA98 PCOV 39.0 12.5 10.0 3.8 18.7 1.3 28.7 9.8 

SP99 PCOV 34.0 10.0 1.7 0.7 33.0 4.7 15.0 2.5 

FA99 PCOV 49.7 11.2 4.0 1.0 18.7 1.5 21.0 10.2 

SP00 PCOV 34.0 9.0 0.7 0.7 36.7 1.5 24.7 7.5 

FA00 PCOV 50.0 12.2 0.0 0.0 30.3 5.7 15.7 8.4 

SP01 PCOV 38.0 13.3 0.0 0.0 20.0 2.5 34.7 15.6 

FA01 PCOV 61.0 10.6 0.7 0.7 21.3 4.3 11.7 8.4 

SP02 PCOV 50.3 11.8 1.3 1.3 35.3 9.9 7.7 3.3 

FA02 PCOV 84.0 6.6 0.0 0.0 9.7 2.8 5.0 4.0 

SP03 PCOV 53.0 10.3 4.0 3.5 18.0 4.5 20.7 9.3 

FA03 PCOV 67.0 11.0 9.0 4.5 20.3 7.0 6.7 5.2 

 

FA99 PTFM 54.0 13.5 26.7 15.9 16.7 5.4 1.3 0.7 

SP00 PTFM 23.0 3.8 17.3 16.8 54.3 11.7 0.3 0.3 

FA00 PTFM 58.7 13.0 19.7 4.4 19.7 9.8 2.0 0.6 

SP01 PTFM 51.3 8.0 13.3 1.8 32.0 9.3 2.3 1.9 

FA01 PTFM 78.0 10.3 8.0 3.6 12.3 6.8 0.3 0.3 

SP02 PTFM 66.7 14.1 1.7 1.7 32.0 13.6 0.3 0.3 
 C-42 

 Phyllospadix Egregia Other algae Sand 

Sample Site mean se mean se mean se mean se 

FA02 PTFM 83.7 7.1 15.3 4.6 11.3 9.4 0.0 0.0 

SP03 PTFM 61.3 13.9 2.7 1.8 32.3 15.9 0.0 0.0 

FA03 PTFM 55.7 7.1 16.7 3.8 22.3 10.3 0.7 0.3 

 

FA96 CRCO 84.3 3.9 1.8 1.0 6.0 1.1 2.5 0.2 

SP97 CRCO 61.5 7.8 2.3 2.1 11.2 3.8 22.2 2.7 

FA97 CRCO 69.5 5.2 0.7 0.7 14.7 4.8 12.2 1.8 

SP98 CRCO 56.2 7.5 7.0 1.7 14.7 3.1 20.7 5.3 

FA98 CRCO 59.8 6.1 26.8 6.1 6.5 2.1 4.2 2.5 

SP99 CRCO 43.8 3.7 28.7 4.3 10.0 2.8 13.5 2.6 

FA99 CRCO 69.6 3.4 16.2 3.7 9.1 3.2 5.7 1.4 

SP00 CRCO 60.0 6.0 14.2 8.4 20.1 3.5 11.3 2.2 

FA00 CRCO 81.3 4.3 14.5 11.5 7.7 3.0 5.7 2.4 

SP01 CRCO 60.0 3.8 5.8 3.1 14.2 4.4 15.3 2.5 

FA01 CRCO 78.3 6.2 6.2 3.6 9.7 4.0 2.8 1.6 

SP02 CRCO 64.2 6.5 2.5 0.8 7.8 3.7 20.0 3.0 

FA02 CRCO 77.8 6.6 3.7 2.3 8.3 3.7 5.8 2.0 

SP03 CRCO 71.0 7.6 2.8 1.3 10.7 4.1 12.7 2.7 

FA03 CRCO 82.5 4.7 1.2 0.8 3.3 1.8 9.3 3.0 
 C-43 

Seastars 

 Pisaster Pisaster

Sample Site mean se Sample Site mean se 

FA95 PSN 15.3 2.3 . FA00 HAZ 38.0 17.3 

SP96 PSN 8.3 2.6 . SP01 HAZ 30.7 15.4 

FA96 PSN 15.7 1.5 . FA01 HAZ 41.3 18.2 

SP97 PSN 6.3 1.9 . SP02 HAZ 19.7 5.9 

FA97 PSN 9.0 0.6 . FA02 HAZ 24.0 13.5 

SP98 PSN 11.3 2.3 . SP03 HAZ 16.0 7.0 

FA98 PSN 18.0 3.5 . FA03 HAZ 53.3 29.0 

SP99 PSN 15.3 3.5 . 

FA99 PSN 18.0 1.5 . FA95 SHB 5.3 0.3 

SP00 PSN 23.0 4.4 . SP96 SHB 3.3 1.2 

FA00 PSN 30.0 8.6 . FA96 SHB 1.7 0.7 

SP01 PSN 19.0 3.2 . SP97 SHB 2.7 1.2 

FA01 PSN 22.0 1.5 . FA97 SHB 3.7 0.3 

SP02 PSN 15.3 2.9 . SP98 SHB 3.7 0.3 

FA02 PSN 24.3 3.4 . FA98 SHB 1.7 0.9 

SP03 PSN 25.3 5.5 . SP99 SHB 2.3 0.3 

FA03 PSN 26.0 2.5 . FA99 SHB 2.7 1.2 

 . SP00 SHB 2.7 1.2 

FA95 CAY 14.3 4.7 . FA00 SHB 6.3 1.7 

SP96 CAY 3.3 1.9 . SP01 SHB 1.3 0.3 

FA96 CAY 4.0 1.2 . FA01 SHB 0.7 0.3 

SP97 CAY 4.0 2.0 . SP02 SHB 0.3 0.3 

FA97 CAY 9.3 1.5 . FA02 SHB 3.3 0.7 

SP98 CAY 3.7 0.9 . SP03 SHB 1.3 0.3 

FA98 CAY 9.7 1.3 . FA03 SHB 7.0 1.5 

SP99 CAY 3.3 1.9 . 

FA99 CAY 4.3 1.5 . SP00 OCC 120.0 . 

SP00 CAY 4.3 1.5 . FA00 OCC 67.0 . 

FA00 CAY 11.0 1.2 . SP01 OCC 60.0 . 

SP01 CAY 9.3 0.9 . FA01 OCC 18.0 . 

FA01 CAY 13.0 2.1 . SP02 OCC 74.0 

SP02 CAY 10.3 2.4 . FA02 OCC 134.0 

FA02 CAY 19.3 2.2 . SP03 OCC 219.0 

SP03 CAY 20.0 4.2 . FA03 OCC 152.0 

FA03 CAY 21.7 4.4 . 

 . SP92 STA 20.7 7.5 

FA95 HAZ 11.3 4.7 . FA92 STA 10.0 5.7 

SP96 HAZ 54.0 19.9 . SP93 STA 12.3 7.8 

FA96 HAZ 22.3 3.2 . FA93 STA 9.0 6.2 

SP97 HAZ 22.3 10.0 . SP94 STA 9.0 4.2 

FA97 HAZ 38.3 19.8 . FA94 STA 2.7 1.3 

SP98 HAZ 28.3 8.6 . SP95 STA 8.3 3.5 

FA98 HAZ 50.7 16.7 . FA95 STA 9.3 5.0 

SP99 HAZ 55.0 17.5 . SP96 STA 21.3 8.7 

FA99 HAZ 22.3 11.1 . FA96 STA 11.3 6.6 

SP00 HAZ 23.3 6.2 . SP97 STA 29.0 13.7 
 C-44 

 Pisaster Asterina Pisaster

Sample Site mean se mean se Sample Site mean se 

FA97 STA 24.7 2.3 . . FA95 GPT 101.0 23.5 

SP98 STA 43.3 18.3 . . SP96 GPT 70.0 21.5 

FA98 STA 27.3 12.4 . . FA96 GPT 76.0 17.7 

SP99 STA 20.3 4.8 . . SP97 GPT 63.0 6.6 

FA99 STA 17.0 6.0 . . FA97 GPT 75.3 15.3 

SP00 STA 12.0 2.1 . . SP98 GPT 39.7 20.7 

FA00 STA 18.7 2.9 . . FA98 GPT 40.0 12.2 

SP01 STA 13.3 2.3 . . SP99 GPT 40.0 11.8 

FA01 STA 5.7 2.9 FA99 GPT . . 

SP02 STA 13.7 3.8 SP00 GPT 39.0 14.4 

FA02 STA 12.0 5.5 FA00 GPT 80.3 9.0 

SP03 STA 13.7 3.0 SP01 GPT 62.5 7.5 

FA03 STA 10.3 3.2 FA01 GPT 57.0 9.0 

 SP02 GPT 93.3 22.0 

SP92 BOA 34.7 13.2 4.3 2.3 FA02 GPT 112.0 19.5 

FA92 BOA 48.7 24.9 11.7 5.5 SP03 GPT 72.0 16.3 

SP93 BOA 33.0 9.1 6.0 2.1 FA03 GPT 94.3 10.8 

FA93 BOA 30.3 11.4 6.7 3.8 

SP94 BOA 23.3 7.1 8.3 4.3 SP02 ALEG 41.0 10.6 

FA94 BOA 35.0 11.0 15.3 6.9 FA02 ALEG 67.3 17.9 

SP95 BOA 31.0 11.0 10.3 5.8 SP03 ALEG 51.0 12.8 

FA95 BOA 25.3 8.8 6.3 3.4 FA03 ALEG 26.0 11.5 

SP96 BOA 34.0 11.0 14.7 3.5 

FA96 BOA 31.0 9.1 12.0 5.5 SP92 ARHO 20.3 5.5 

SP97 BOA 35.7 14.5 8.0 1.0 FA92 ARHO 12.0 1.2 

FA97 BOA 30.7 8.5 6.0 1.0 SP93 ARHO 7.7 5.4 

SP98 BOA 22.3 3.7 1.7 0.3 FA93 ARHO 6.7 4.3 

FA98 BOA 28.0 11.9 3.3 1.2 SP94 ARHO 2.0 1.5 

SP99 BOA 9.7 4.9 0.0 0.0 FA94 ARHO 4.0 1.5 

FA99 BOA 28.7 10.1 4.7 3.7 SP95 ARHO 2.0 0.6 

SP00 BOA 26.7 12.4 2.3 2.3 FA95 ARHO 1.3 0.3 

FA00 BOA 46.7 21.2 5.7 2.2 SP96 ARHO 1.0 0.6 

SP01 BOA 50.0 16.0 2.3 1.3 FA96 ARHO 7.0 4.4 

FA01 BOA 29.0 13.7 0.0 0.0 SP97 ARHO 12.0 8.5 

SP02 BOA 22.3 11.9 3.0 1.7 FA97 ARHO 6.3 4.1 

FA02 BOA 36.3 12.7 2.0 0.6 SP98 ARHO 3.0 2.5 

SP03 BOA 25.7 9.2 3.0 2.1 FA98 ARHO 1.0 0.6 

FA03 BOA 22.0 8.5 5.3 1.2 SP99 ARHO 0.7 0.3 

 FA99 ARHO 1.3 0.9 

SP92 GPT 26.3 13.9 . . SP00 ARHO 3.3 1.2 

FA92 GPT 66.3 11.0 . . FA00 ARHO 3.3 1.5 

SP93 GPT . . . . SP01 ARHO 2.3 1.9 

FA93 GPT 48.3 14.4 . . FA01 ARHO 14.7 10.3 

SP94 GPT 66.3 14.8 . . SP02 ARHO 9.0 5.6 

FA94 GPT 43.7 9.3 . . FA02 ARHO 18.3 9.8 

SP95 GPT 78.0 18.9 . . SP03 ARHO 13.7 6.8 
 C-45 

 Pisaster Pisaster

Sample Site mean se Sample Site mean se 

FA03 ARHO 11.7 6.3 FA94 OLDS 10.3 2.2 

 SP95 OLDS 6.7 3.4 

SP92 CARP 12.7 5.8 FA95 OLDS 6.7 2.7 

FA92 CARP 13.3 4.6 SP96 OLDS 3.3 2.3 

SP93 CARP 12.3 0.3 FA96 OLDS 2.7 1.8 

FA93 CARP 8.0 3.6 SP97 OLDS 5.7 3.8 

SP94 CARP 8.0 4.0 FA97 OLDS 1.3 1.3 

FA94 CARP 3.0 3.0 SP98 OLDS 1.7 1.7 

SP95 CARP . . FA98 OLDS 1.7 1.7 

FA95 CARP . . SP99 OLDS 0.0 0.0 

SP96 CARP 2.7 2.7 FA99 OLDS 2.0 1.0 

FA96 CARP . . SP00 OLDS 3.3 0.7 

SP97 CARP 1.0 0.0 FA00 OLDS . . 

FA97 CARP 1.3 0.9 SP01 OLDS . . 

SP98 CARP 0.0 0.0 FA01 OLDS 16.3 5.2 

FA98 CARP . . SP02 OLDS 6.3 6.3 

SP99 CARP 0.3 0.3 FA02 OLDS 11.3 5.9 

FA99 CARP . . SP03 OLDS 24.3 9.3 

SP00 CARP 6.3 6.3 FA03 OLDS 23.3 9.7 

FA00 CARP 3.3 3.3 

SP01 CARP 5.3 4.8 SP02 PCOV 59.7 28.8 

FA01 CARP . . FA02 PCOV 97.3 31.2 

SP02 CARP 24.0 15.1 SP03 PCOV 94.7 39.0 

FA02 CARP . . FA03 PCOV 54.0 9.0 

SP03 CARP 16.0 7.8 

FA03 CARP . . SP03 PTFM 25.3 6.7 

 FA03 PTFM 29.0 6.1 

FA94 MUSH 24.3 8.0 

SP95 MUSH 22.3 10.7 SP97 CRCO 13.0 

FA95 MUSH 24.3 13.7 FA97 CRCO 7.0 

SP96 MUSH 14.0 8.1 SP98 CRCO 2.0 

FA96 MUSH 30.0 9.6 FA98 CRCO 6.0 

SP97 MUSH 23.0 11.4 SP99 CRCO 42.0 

FA97 MUSH 12.0 7.2 FA99 CRCO 89.0 

SP98 MUSH 4.3 3.8 SP00 CRCO . 

FA98 MUSH 4.3 2.3 FA00 CRCO 41.0 

SP99 MUSH 4.7 1.5 SP01 CRCO 27.0 

FA99 MUSH 6.3 1.8 FA01 CRCO 30.0 

SP00 MUSH 7.7 2.6 SP02 CRCO 36.0 

FA00 MUSH FA02 CRCO 30.0 

SP01 MUSH SP03 CRCO 34.0 

FA01 MUSH 17.7 8.6 FA03 CRCO 40.0 

SP02 MUSH 12.3 5.9 

FA02 MUSH 16.7 6.4 SP97 SHCO 3.0 

SP03 MUSH 12.3 5.2 FA97 SHCO 1.0 

FA03 MUSH 21.7 11.6 SP98 SHCO 13.0 
 C-46 

 Pisaster Pisaster

Sample Site mean Sample Site mean 

FA98 SHCO 7.0 SP02 CTBR 1.0 

SP99 SHCO 39.0 FA02 CTBR . 

FA99 SHCO 15.0 SP03 CTBR 2.0 

SP00 SHCO 60.0 FA03 CTBR 0.0 

FA00 SHCO 22.0 SP02 CTLH 0.0 

SP01 SHCO 48.0 FA02 CTLH . 

FA01 SHCO 15.0 SP03 CTLH 3.0 

SP02 SHCO 12.0 FA03 CTLH 1.0 

FA02 SHCO 52.0 

SP03 SHCO 24.0 

FA03 SHCO 33.0 

 

SP97 TRIS 2.0 

FA97 TRIS 0.0 

SP98 TRIS 5.0 

FA98 TRIS 1.0 

SP99 TRIS 14.0 

FA99 TRIS 56.0 

SP00 TRIS 16.0 

FA00 TRIS 1.0 

SP01 TRIS 2.0 

FA01 TRIS 0.0 

SP02 TRIS 8.0 

FA02 TRIS 13.0 

SP03 TRIS 30.0 

FA03 TRIS 34.0 

 

FA96 DAPT 0.0 

SP97 DAPT 1.0 

FA97 DAPT 0.0 

SP98 DAPT 1.0 

FA98 DAPT 2.0 

SP99 DAPT 4.0 

FA99 DAPT 10.0 

SP00 DAPT 2.0 

FA00 DAPT 2.0 

SP01 DAPT 1.0 

FA01 DAPT 3.0 

SP02 DAPT 2.0 

FA02 DAPT 1.0 

SP03 DAPT . 

FA03 DAPT 18.0 

 
 C-47 

Lottia

 Count Size Count Size 

Sample Site mean se mean se Sample Site mean se mean se 

SP02 RMR 42.0 4.9 37.1 0.7 SP95 STA 40.8 4.4 41.7 1.1 

FA02 RMR 36.8 3.2 36.4 0.8 FA95 STA 35.8 3.6 40.5 1.1 

SP03 RMR 39.4 4.0 36.2 0.8 SP96 STA 32.2 3.4 43.0 1.1 

FA03 RMR 41.0 6.2 32.1 0.9 FA96 STA 28.8 3.2 45.3 1.2 

 SP97 STA 28.8 3.2 42.2 1.2 

FA95 CAY 46.6 4.0 31.8 0.6 FA97 STA 29.4 3.0 40.7 1.3 

SP96 CAY 58.8 6.4 32.4 0.5 SP98 STA 29.8 2.0 39.7 1.2 

FA96 CAY 54.8 10.4 30.5 0.6 FA98 STA 45.4 4.1 35.4 1.2 

SP97 CAY 47.2 10.4 29.9 0.6 SP99 STA 40.8 4.1 35.8 1.3 

FA97 CAY 38.6 8.7 30.0 0.7 FA99 STA 44.2 2.2 38.7 1.2 

SP98 CAY 35.6 9.3 32.6 0.6 SP00 STA 37.8 0.7 44.6 1.3 

FA98 CAY 73.0 15.8 25.9 0.5 FA00 STA 38.0 1.7 46.6 1.3 

SP99 CAY 55.8 8.7 24.5 0.5 SP01 STA 41.6 2.6 46.8 1.2 

FA99 CAY 45.0 7.0 30.8 0.6 FA01 STA 39.6 1.3 49.5 1.3 

SP00 CAY 59.0 9.3 30.7 0.5 SP02 STA 39.8 2.3 46.8 1.3 

FA00 CAY 46.0 9.7 31.2 0.7 FA02 STA 33.8 1.7 51.3 1.4 

SP01 CAY 38.6 6.7 35.5 0.7 SP03 STA 26.0 9.1 47.6 1.7 

FA01 CAY 30.4 6.3 35.7 1.0 FA03 STA 29.0 6.5 49.1 1.3 

SP02 CAY 22.2 5.1 36.8 0.7 

FA02 CAY 34.0 8.1 29.8 1.0 SP92 BOA 36.6 6.1 46.0 1.0 

SP03 CAY 32.2 7.5 32.4 0.9 FA92 BOA 42.2 7.8 46.9 1.0 

FA03 CAY 35.8 8.4 25.5 0.8 SP93 BOA 34.6 5.9 45.6 1.1 

 FA93 BOA 47.6 8.5 42.4 1.1 

SP96 HAZ 21.6 2.0 32.5 1.0 SP94 BOA 39.8 7.8 44.8 1.1 

FA96 HAZ 18.4 2.3 30.7 1.1 FA94 BOA 41.8 6.2 45.8 1.1 

SP97 HAZ 15.8 1.3 33.2 1.2 SP95 BOA 44.6 4.9 45.1 1.0 

FA97 HAZ 17.6 2.2 34.4 1.2 FA95 BOA 44.0 3.7 46.0 1.0 

SP98 HAZ 18.2 1.6 34.3 1.2 SP96 BOA 41.0 3.4 46.3 1.0 

FA98 HAZ 20.4 3.8 25.0 0.8 FA96 BOA 21.0 2.7 41.6 1.2 

SP99 HAZ 30.0 4.5 26.0 0.7 SP97 BOA 23.0 3.6 42.7 1.1 

FA99 HAZ 20.6 3.1 31.6 0.8 FA97 BOA 30.8 1.9 37.1 1.3 

SP00 HAZ 20.8 2.7 35.2 0.9 SP98 BOA 30.6 2.7 38.3 1.3 

FA00 HAZ 21.8 1.5 32.7 1.1 FA98 BOA 34.2 3.7 36.4 1.1 

SP01 HAZ 21.8 2.2 33.5 1.2 SP99 BOA 41.0 4.9 35.9 0.9 

FA01 HAZ 18.2 1.3 38.8 1.1 FA99 BOA 34.6 4.2 43.2 1.0 

SP02 HAZ 17.8 1.5 41.0 1.2 SP00 BOA 39.0 4.6 43.6 0.9 

FA02 HAZ 23.0 2.3 36.2 1.5 FA00 BOA 45.2 3.6 45.3 0.9 

SP03 HAZ 23.2 1.9 36.6 1.4 SP01 BOA 43.2 2.8 45.0 0.9 

FA03 HAZ 22.2 2.0 39.7 1.5 FA01 BOA 47.8 5.8 45.4 0.9 

 SP02 BOA 48.6 5.5 45.8 0.9 

SP92 STA 25.0 1.4 44.0 1.2 FA02 BOA 41.6 6.3 44.5 1.0 

FA92 STA 24.6 2.1 46.4 1.2 SP03 BOA 41.2 4.8 44.5 1.0 

SP93 STA 31.8 3.2 44.9 1.2 FA03 BOA 45.0 6.6 42.7 1.0 

FA93 STA 23.4 3.1 42.9 1.4 

SP94 STA 38.4 3.3 37.0 1.1 

FA94 STA 36.4 3.8 41.7 1.3 
 C-48 

 Count Size Count Size 

Sample Site mean se mean se Sample Site mean se mean se 

SP92 GPT 27.4 1.3 34.6 1.0 SP03 ALEG 24.4 3.9 43.7 0.9 

FA92 GPT 27.0 4.0 35.3 0.9 FA03 ALEG 19.6 3.7 46.5 1.1 

SP93 GPT 27.4 3.7 32.2 0.9 

FA93 GPT 24.0 5.6 37.1 1.0 FA01 CARP 29.8 7.1 27.7 0.5 

SP94 GPT 40.0 7.9 28.7 0.8 SP02 CARP 32.4 6.1 28.4 0.5 

FA94 GPT 33.8 5.9 30.0 0.9 FA02 CARP 49.6 6.5 28.5 0.4 

SP95 GPT 42.8 5.9 28.4 0.7 SP03 CARP 28.6 5.6 26.3 0.6 

FA95 GPT 33.0 6.1 31.6 0.7 FA03 CARP 45.4 11.1 22.3 0.6 

SP96 GPT 30.0 4.6 32.2 0.8 

FA96 GPT 33.4 3.4 31.6 1.0 SP02 MUSH 52.4 6.6 36.6 0.7 

SP97 GPT 31.2 3.4 32.3 1.0 FA02 MUSH 64.0 5.6 33.4 0.6 

FA97 GPT 24.8 1.4 35.4 1.2 SP03 MUSH 20.0 2.5 35.6 1.0 

SP98 GPT 25.0 4.0 37.5 1.2 FA03 MUSH 24.0 1.8 35.1 1.3 

FA98 GPT 37.6 5.2 30.3 1.1 

SP99 GPT 43.2 11.0 30.3 0.8 FA94 OLDS 15.8 2.6 32.6 1.0 

FA99 GPT . . . . SP95 OLDS 18.4 3.8 32.6 0.9 

SP00 GPT 38.4 7.7 29.8 0.9 FA95 OLDS 11.4 3.8 31.7 1.1 

FA00 GPT 35.2 6.0 32.8 0.9 SP96 OLDS 13.2 4.1 30.6 1.0 

SP01 GPT 38.8 7.9 28.5 0.7 FA96 OLDS 17.4 4.3 29.7 0.9 

FA01 GPT 27.0 5.3 34.0 0.9 SP97 OLDS 20.8 8.0 25.7 0.9 

SP02 GPT 28.8 5.4 33.7 0.9 FA97 OLDS 15.8 5.4 26.9 0.9 

FA02 GPT 45.8 10.5 29.0 0.7 SP98 OLDS 15.4 5.1 29.5 0.7 

SP03 GPT 50.8 12.3 26.9 0.7 FA98 OLDS 21.0 4.6 28.7 0.7 

FA03 GPT 56.8 11.8 27.8 0.7 SP99 OLDS 24.4 5.7 28.1 0.8 

 FA99 OLDS 26.2 4.4 23.9 0.8 

FA92 ALEG 11.4 0.8 52.1 2.5 SP00 OLDS 51.2 11.2 25.3 0.5 

SP93 ALEG 12.8 2.2 48.2 1.8 FA00 OLDS 44.8 8.1 29.3 0.6 

FA93 ALEG 13.8 2.6 50.6 2.0 SP01 OLDS 51.6 8.1 29.8 0.5 

SP94 ALEG 15.2 1.8 49.8 1.7 FA01 OLDS 52.4 8.0 31.1 0.6 

FA94 ALEG 12.4 1.5 49.9 2.0 SP02 OLDS 44.0 7.4 29.8 0.6 

SP95 ALEG 18.0 1.1 46.4 1.7 FA02 OLDS 38.4 2.0 31.2 0.6 

FA95 ALEG 14.6 2.0 50.2 1.9 SP03 OLDS 17.8 2.4 28.0 0.9 

SP96 ALEG 14.2 1.5 48.8 1.9 FA03 OLDS 20.0 2.6 28.7 0.9 

FA96 ALEG 13.6 1.6 52.5 1.8 

SP97 ALEG 14.6 1.9 49.8 2.0 FA94 PCOV 39.4 8.4 28.7 0.7 

FA97 ALEG 13.6 2.6 49.5 2.1 SP95 PCOV 39.2 6.1 29.0 0.7 

SP98 ALEG 16.2 3.1 46.8 1.9 FA95 PCOV 32.4 5.0 30.3 0.8 

FA98 ALEG 27.4 2.4 44.2 1.5 SP96 PCOV 33.8 5.7 28.8 0.5 

SP99 ALEG 25.0 3.2 42.5 1.6 FA96 PCOV 41.6 8.6 29.3 0.7 

FA99 ALEG 28.0 2.6 40.9 1.6 SP97 PCOV 41.8 8.2 30.1 0.7 

SP00 ALEG 25.8 5.7 41.0 1.4 FA97 PCOV 35.2 7.6 32.5 0.8 

FA00 ALEG 30.4 3.6 43.5 1.1 SP98 PCOV 68.8 25.8 23.1 0.6 

SP01 ALEG 32.8 3.8 41.4 1.0 FA98 PCOV 52.4 11.2 30.2 0.6 

FA01 ALEG 28.4 3.9 44.2 1.0 SP99 PCOV 55.0 10.8 29.2 0.6 

SP02 ALEG 24.8 3.6 45.0 0.9 FA99 PCOV 74.0 15.8 27.0 0.6 

FA02 ALEG 45.2 6.6 41.0 0.7 SP00 PCOV 146.8 27.1 26.9 0.4 
 C-49 

 Count Size Count Size 

Sample Site mean se mean se Sample Site mean se mean se 

FA00 PCOV 126.4 28.7 28.4 0.5 FA96 DAPT 26.4 7.4 33.3 0.9 

SP01 PCOV 110.4 24.0 30.4 0.5 SP97 DAPT 26.0 5.5 34.9 1.0 

FA01 PCOV 96.8 20.3 28.9 0.5 FA97 DAPT 34.8 6.0 34.8 0.9 

SP02 PCOV 101.6 19.1 27.2 0.5 SP98 DAPT 29.2 5.3 40.5 1.0 

FA02 PCOV 58.4 18.0 29.9 0.6 FA98 DAPT 32.2 4.4 38.7 1.0 

SP03 PCOV 40.6 9.2 25.9 0.6 SP99 DAPT 24.8 4.4 43.5 1.1 

FA03 PCOV 47.4 15.9 27.5 0.6 FA99 DAPT 34.6 8.0 36.4 1.2 

 SP00 DAPT 27.0 6.5 38.4 1.2 

SP03 PTFM 31.2 4.3 31.8 0.7 FA00 DAPT 23.2 5.5 30.5 1.1 

FA03 PTFM 24.4 7.2 27.0 1.0 SP01 DAPT 23.2 4.5 33.5 1.1 

 FA01 DAPT 31.2 3.8 33.7 1.0 

FA96 CRCO 62.4 7.7 36.3 0.8 SP02 DAPT 32.6 4.2 32.9 1.0 

SP97 CRCO 62.6 7.6 35.2 0.6 FA02 DAPT 29.2 5.7 32.2 1.2 

FA97 CRCO 59.2 11.1 33.3 0.6 SP03 DAPT 27.6 4.5 33.2 1.1 

SP98 CRCO 52.6 6.8 35.8 0.6 FA03 DAPT 37.5 0.5 26.3 1.3 

FA98 CRCO 48.2 8.1 37.0 0.7 

SP99 CRCO 60.2 5.6 36.8 0.6 

FA99 CRCO 57.4 8.4 36.2 0.7 

SP00 CRCO 36.6 7.6 37.9 0.8 

FA00 CRCO 37.2 10.7 36.5 0.8 

SP01 CRCO 43.3 16.9 37.5 0.6 

FA01 CRCO 49.8 11.6 37.7 0.7 

SP02 CRCO 49.4 13.0 36.0 0.7 

FA02 CRCO 41.8 12.4 36.5 0.7 

SP03 CRCO 37.8 13.1 36.6 0.6 

FA03 CRCO 70.5 15.5 35.9 1.0 

 

FA96 SHCO 83.6 15.2 27.3 0.2 

SP97 SHCO 91.2 13.6 27.6 0.2 

FA97 SHCO 117.2 13.5 26.5 0.2 

SP98 SHCO 148.4 19.4 25.5 0.2 

FA98 SHCO 135.4 26.1 24.1 0.2 

SP99 SHCO 128.8 21.1 26.5 0.2 

FA99 SHCO 131.8 27.0 24.7 0.2 

SP00 SHCO 126.6 20.3 26.3 0.2 

FA00 SHCO 127.2 18.8 26.3 0.2 

SP01 SHCO 157.8 29.9 26.6 0.2 

FA01 SHCO 133.8 34.3 26.1 0.2 

SP02 SHCO 120.6 22.6 27.2 0.2 

FA02 SHCO 124.2 26.2 26.6 0.2 

SP03 SHCO 90.6 17.9 27.4 0.2 

FA03 SHCO 97.6 31.7 24.8 0.3 

 
 C-50 

Abalone 

 Size Size 

Sample Site Total mean se Sample Site Total mean se 

F95 PSN 117 80.4 2.0 SP00 Cayucos 4 87.5 9.5 

SP96 PSN 133 87.1 2.3 F00 Cayucos 7 90.0 6.9 

F96 PSN 128 80.1 2.1 SP01 Cayucos 5 88.0 12.8 

SP97 PSN 254 79.4 1.3 F01 Cayucos 0 . . 

F97 PSN 170 81.9 1.6 SP02 Cayucos 1 90.0 . 

SP98 PSN 288 91.5 1.4 F02 Cayucos 2 100.0 0.0 

F98 PSN 305 85.0 1.8 SP03 Cayucos 4 102.5 6.3 

SP99 PSN 254 78.2 2.0 F03 Cayucos 1 110.0 . 

F99 PSN 288 78.3 1.9 

SP00 PSN 339 79.2 1.7 F93 Purisima 394 98.4 1.2 

F00 PSN 362 76.2 1.7 SP94 Purisima 381 99.7 1.2 

SP01 PSN 405 80.9 1.5 F94 Purisima . . . 

F01 PSN 343 81.9 1.7 SP95 Purisima 510 100.4 1.1 

SP02 PSN 486 82.6 1.3 F95 Purisima 509 97.8 1.2 

F02 PSN 450 81.0 1.3 SP96 Purisima 535 96.4 1.2 

SP03 PSN 478 82.1 1.2 F96 Purisima 521 91.6 1.3 

F03 PSN 348 88.4 1.3 SP97 Purisima 551 84.3 1.0 

 F97 Purisima 537 88.8 1.1 

F97 Piedras 201 88.3 1.5 SP98 Purisima 492 98.7 1.3 

SP98 Piedras 197 93.6 1.7 F98 Purisima 219 88.2 1.5 

F98 Piedras 175 86.3 1.8 SP99 Purisima 71 85.6 3.2 

SP99 Piedras 165 83.8 1.9 F99 Purisima 52 91.9 3.5 

F99 Piedras 163 72.7 2.0 SP00 Purisima 49 87.3 3.0 

SP00 Piedras 169 80.5 2.2 F00 Purisima 50 90.8 3.3 

F00 Piedras 204 88.9 1.8 SP01 Purisima 33 100.9 2.7 

SP01 Piedras 217 85.3 2.0 F01 Purisima 16 106.9 7.8 

F01 Piedras 203 83.3 1.9 SP02 Purisima 22 101.8 4.5 

SP02 Piedras 219 80.3 2.0 F02 Purisima 21 107.1 5.2 

F02 Piedras 202 75.9 1.8 SP03 Purisima 14 103.6 6.8 

SP03 Piedras 196 84.9 1.7 F03 Purisima 13 113.8 9.0 

F03 Piedras 149 85.2 1.9 

 SP92 Stairs 275 76.0 1.0 

SP01 R Marino 69 78.1 2.5 F92 Stairs 414 75.3 1.0 

F01 R Marino 46 75.2 2.0 SP93 Stairs 434 75.2 0.8 

SP02 R Marino 54 78.0 2.3 F93 Stairs 318 74.8 0.9 

F02 R Marino 65 75.8 2.5 SP94 Stairs 407 81.1 0.9 

SP03 R Marino 63 80.5 2.2 F94 Stairs 494 78.0 1.1 

F03 R Marino 46 82.2 2.3 SP95 Stairs 536 76.6 1.1 

 F95 Stairs 419 70.6 1.2 

F96 Cayucos 19 80.5 5.2 SP96 Stairs 428 72.4 1.0 

SP97 Cayucos 19 66.8 4.8 F96 Stairs 204 70.9 1.6 

F97 Cayucos 16 77.5 5.0 SP97 Stairs 164 66.5 1.9 

SP98 Cayucos 14 92.1 4.7 F97 Stairs 66 77.7 2.8 

F98 Cayucos 3 73.3 17.6 SP98 Stairs 46 90.2 3.9 

SP99 Cayucos 4 100.0 10.8 F98 Stairs 17 77.1 6.0 

F99 Cayucos 3 90.0 10.0 SP99 Stairs 18 83.3 5.4 
 C-51 

 Size Size 

Sample Site Total mean se Sample Site Total mean se 

F99 Stairs 19 81.1 4.1 SP92 Govpt 87 105.2 1.9 

SP00 Stairs 26 85.8 4.7 F92 Govpt 77 90.6 2.2 

F00 Stairs 25 82.0 4.1 SP93 Govpt . . . 

SP01 Stairs 28 94.8 4.2 F93 Govpt 16 106.9 5.0 

F01 Stairs 26 92.7 4.5 SP94 Govpt 5 119.0 6.8 

SP02 Stairs 26 90.8 4.3 F94 Govpt 4 115.0 8.7 

F02 Stairs 30 88.3 3.3 SP95 Govpt 6 120.0 4.3 

SP03 Stairs 21 91.9 4.0 F95 Govpt 2 115.0 5.0 

F03 Stairs 24 101.7 4.6 SP96 Govpt 2 125.0 15.0 

 F96 Govpt 2 115.0 5.0 

SP92 Boathouse 578 81.9 1.0 SP97 Govpt 4 105.0 8.7 

F92 Boathouse 441 86.8 1.2 F97 Govpt 3 126.7 18.6 

SP93 Boathouse 407 78.0 1.2 SP98 Govpt 1 120.0 . 

F93 Boathouse 366 89.8 1.2 F98 Govpt 1 140.0 . 

SP94 Boathouse 448 87.4 1.2 SP99 Govpt 1 110.0 . 

F94 Boathouse 435 89.1 1.4 F99 Govpt . . . 

SP95 Boathouse 353 90.8 1.1 SP00 Govpt 3 106.7 23.3 

F95 Boathouse 172 91.5 2.2 F00 Govpt 1 110.0 . 

SP96 Boathouse 114 82.7 2.8 SP01 Govpt 1 120.0 . 

F96 Boathouse 78 90.5 3.2 F01 Govpt 0 . . 

SP97 Boathouse 90 82.8 3.0 SP02 Govpt 1 80.0 . 

F97 Boathouse 102 74.7 3.1 F02 Govpt 1 100.0 . 

SP98 Boathouse 40 89.0 4.7 SP03 Govpt 0 . . 

F98 Boathouse 25 69.4 5.0 F03 Govpt 0 . . 

SP99 Boathouse 12 92.5 6.3 

F99 Boathouse 20 85.7 5.8 

SP00 Boathouse 34 72.1 5.4 

F00 Boathouse 30 77.8 6.5 

SP01 Boathouse 39 80.0 4.7 

F01 Boathouse 28 103.2 5.3 

SP02 Boathouse 34 99.7 4.2 

F02 Boathouse 33 96.1 5.4 

SP03 Boathouse 24 106.7 4.5 

F03 Boathouse 26 105.0 5.4 

 
The Department of the Interior Mission

As the Nation's principal conservation agency, the Department of the Interior has responsibility for most 

of our nationally owned public lands and natural resources. This includes fostering sound use of our 

land and water resources; protecting our fish, wildlife, and biological diversity; preserving the 

environmental and cultural values of our national parks and historical places; and providing for the

enjoyment of life through outdoor recreation. The Department assesses our energy and mineral 

resources and works to ensure that their development is in the best interests of all our people by 

encouraging stewardship and citizen participation in their care. The Department also has a major 

responsibility for American Indian reservation communities and for people who live in island territories 

under U.S. administration. 

The Minerals Management Service Mission 

As a bureau of the Department of the Interior, the Minerals Management Service's (MMS) primary 

responsibilities are to manage the mineral resources located on the Nation's Outer Continental Shelf 

(OCS), collect revenue from the Federal OCS and onshore Federal and Indian lands, and distribute 

those revenues. 

Moreover, in working to meet its responsibilities, the Offshore Minerals Management Program

administers the OCS competitive leasing program and oversees the safe and environmentally sound 

exploration and production of our Nation's offshore natural gas, oil and other mineral resources. The 

MMS Royalty Management Program meets its responsibilities by ensuring the efficient, timely and 

accurate collection and disbursement of revenue from mineral leasing and production due to Indian

tribes and allottees, States and the U.S. Treasury. 

The MMS strives to fulfill its responsibilities through the general guiding principles of: (1) being 

responsive to the public's concerns and interests by maintaining a dialogue with all potentially affected 

parties and (2) carrying out its programs with an emphasis on working to enhance the quality of life for 

all Americans by lending MMS assistance and expertise to economic development and environmental 

protection.