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 

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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. 

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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