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Title A historical perspective of California recreational fisheries using a new database of "trophy" fish records (1966-2013), combined with fisheries analyses of three species in the genus Paralabrax

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Author Bellquist, Lyall F.

Publication Date 2015

Peer reviewed|Thesis/dissertation

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UNIVERSITY OF CALIFORNIA, SAN DIEGO

A historical perspective of California recreational fisheries using a new database of “trophy” fish records (1966-2013), combined with fisheries analyses of three species in the genus Paralabrax

A dissertation submitted in partial satisfaction of the requirements for the degree Doctor of Philosophy

Marine Biology

Lyall F. Bellquist

Committee in charge: Brice Semmens, Chair Richard Carson David Checkley Philip Hastings Ed Parnell

2015

Lyall F. Bellquist, 2015

The Dissertation of Lyall F. Bellquist is approved, and it is acceptable in quality and form for publication on microfilm and electronically:

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______Chair

University of California, San Diego

2015

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DEDICATION

This dissertation is dedicated to my grandfathers: Eric Cyril Bellquist (1904- 1979), who had a 50-year career in political science at the University of California, Berkeley, and who I only had the pleasure of knowing for just a few minutes on August 17, 1979; and Richard R. Ralphs (1925-1982), CEO of Ralphs Grocery Co., of whom I only have one fleeting, yet wonderful memory. I had very little time with either of them, but they have helped shape who I am today, and I know they would have been very proud of this accomplishment.

TABLE OF CONTENTS

Signature page……………………………………………………………………………iii

Dedication ...…………………………………………………………………...…………iv

Table of Contents…………………………………………………...……….…………….v

List of Figures………………………………………...………………………………….vii

List of Tables………………………………………………………………………….....xii

Acknowledgements………………………………………………………….……….….xiv

Vita…………….………………………………………………...…………………….xviii

Abstract of the Dissertation……………………………………………………..………xix

Chapter 1: Long-term changes in “trophy” sizes of pelagic and coastal pelagic fishes among California recreational fisheries (1966-2013)....…………………………………21 Abstract.………………………………………………………………………….21 Introduction……………………………………………...…………….…………22 Methods………………………….…………………...…………………….…….26 Results.…….………………………………………………………….………….30 Discussion.…………………………………………...…………………………..34 Literature Cited.…………………………………...………………………..……52

Chapter 2: Temporal and spatial dynamics of “trophy” sized demersal fishes off the California coast, 1966-2013……………………………………………………………...57 Abstract ...………………………………………….….…………………………57 Introduction…….………………………………….….………………………….58 Methods………….………………………………….……………………………62 Results ……………………………………………….…….…………………….65 Discussion………………………………………………………………………..70 Literature Cited…………………………………………..………………………82

Chapter 3: Analyses of catch-per-unit-effort (CPUE), length frequency, and post-release mortality of three species of Paralabrax relative to coastal marine protected areas (MPAs) off San Diego, California ..………...……………………………………….…..86 Abstract ……...…………………………………………………..………………86 Introduction………………….…………………………………...………………87 Methods …………………….……………………………………………………91

Results …...…………………………………………………………..…………..95 Discussion………………………………………………………………………..99 Literature Cited.……...…………………………………………………...…….123

Chapter 4: Movement patterns of three species of Paralabrax using tag-recapture and acoustic telemetry methods in the coastal marine region of San Diego, California...…126 Abstract ...………………………………………………………………………126 Introduction.……………………….……………………………………………127 Methods.….……………….…………………………………………………….130 Results…………….…………………………………………………………….133 Discussion ……………………………………………………………………...137 Literature Cited ………………………………………………………………...164

LIST OF FIGURES

Figure 1.1. Metacorrelation analysis results for pairwise comparisons between California commercial landings and recreational logbook landings (A), commercial landings and WON trophy frequency (B), and recreational logbook landings and WON trophy frequency (C). Error bars represent 95% confidence intervals….……………………….42

Figure 1.2. Time series plots of trophy fish weight showing model output (white line) and model error (grey region) overlaid onto boxplots of the raw data for Bigeye Tuna (A, n = 264), Yellowfin Tuna (B, n = 536), Pacific Bluefin Tuna (C, n = 694), and Pacific Albacore (D, n = 1526)…………………………………….…………………………….43

Figure 1.3. Time series plots of trophy fish weight showing model output (white line) and model error (grey region) overlaid onto boxplots of the raw data for Dolphinfish (dorado) (A, n = 138), Striped Marlin (B, n = 133), Opah (C, n = 45), Shortfin Mako (D, n = 120), and Common Thresher Shark (E, n = 85)………………………………..………………44

Figure 1.4. Time series plots of trophy fish weight showing model output (white line) and model error (grey region) overlaid onto boxplots for White Seabass (A, n = 4184), California Yellowtail (B, n= 5574), Pacific Bonito (C, n = 389), Pacific Barracuda (D, n = 565), Striped Bass (E, n = 1142), and Chinook Salmon (F, n = 6045)………………...45

Figure 1.5. Cross correlation results for Pacific Bluefin Tuna (black line), Yellowfin Tuna (gray line), and Pacific Albacore (dashed line). Panels show correlations with the MEI index (A), PDO index (B), NPGO index (C), SIO pier water temperature at 5m depth (D), southern California upwelling index (E), and CPFV fishing effort in angler- days (F)…………………………………………………………………………………..46

Figure 1.6. Cross correlation results for common Dolphinfish (black line), Striped Marlin (gray line), and Opah (dashed line). Panels show correlations with the MEI index (A), PDO index (B), NPGO index (C), SIO pier water temperature at 5m depth (D), southern California upwelling index (E), and CPFV fishing effort in angler-days (F)……………47

Figure 1.7. Cross correlation results for California Yellowtail, Pacific Bonito, Pacific Barracuda, and White Seabass. Panels show correlations with the MEI index (A), PDO index (B), NPGO index (C), SIO pier water temperature at 5m depth (D), southern California upwelling index (E), and CPFV fishing effort in angler-days (F) …………...48

Figure 1.8. Cross correlation results for Common Thresher (black line) and Shortfin Mako (gray line). Panels show correlations with the MEI index (A), PDO index (B), NPGO index (C), SIO pier water temperature at 5m depth (D), southern California upwelling index (E), and CPFV fishing effort in angler-days (F)……………………….49

Figure 1.9. Cross correlation results for chinook (king) salmon (black line) and Striped Bass (gray line). Panels show correlations with the MEI index (A), PDO index (B),

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NPGO index (C), SIO pier water temperature at 5m depth (D), southern California upwelling index (E), and CPFV fishing effort in angler-days (F)…………………….....50

Figure 1.10. Number of total resident annual fishing licenses sold in California (gray line), and per capita annual resident fishing licenses (x 100) sold in California (black line) from 1970 – 2013. Non-resident annual license statistics were not included due to relatively negligible values………………………………………..……………………..51

Figure 2.1. Time series plots of trophy fish weight showing model output (white line) and error (grey region) overlaid onto boxplots of the raw data for Cowcod (A, n = 2535), Vermilion Rockfish (B, n = 1083), Bocaccio (C, n = 364), Copper Rockfish (D, n = 72), Yelloweye Rockfish (E, n = 195), and all Sebastes excluding Cowcod (F, n = 1787)…..74

Figure 2.2. Time series plots of trophy fish weight showing model output (white line) and error (grey region) overlaid onto boxplots for Cabezon (A, n = 192), Lingcod (B, n = 7884), White Sturgeon (C, n = 2877), and California Halibut (D, n = 7729)…………....75

Figure 2.3. Time series plots of trophy fish weight showing model output (white line) and error (grey region) overlaid onto boxplots of the raw data for California Sheephead (A, n = 1036), Ocean Whitefish (B, n = 70), Giant Seabass (C, n = 231), Kelp Bass (D, n = 866), and Barred Sand Bass (E, n = 556)……………………….…………………..……76

Figure 2.4. Time series plots of trophy fish weight showing model output (white line) and error (grey region) overlaid onto boxplots of the raw data for Sevengill Shark (A, n = 148) and leopard Shark (B, n = 105)………………………………………………...…..77

Figure 2.5. Boxplots showing median weights of Cowcod (A, n = 1903), Vermilion Rockfish (B, n = 708), Bocaccio (C, n = 256), Yelloweye Rockfish (D, n = 153), and all Sebastes (E, n = 3104) at three distance bins from home port. Gray bars represent 1966 – 1989, and white bars represent 1990 – 2013.……………………………………………78

Figure 2.6. Boxplots showing median weights of California Sheephead (A, n = 573), Lingcod (B, n = 5652), Cabezon (C, n = 141), California Halibut (D, n = 4831), Kelp Bass (E, n = 541), and Barred Sand Bass (F, n = 353) at three distance bins from home port. Gray bars represent 1966 – 1989, and white bars represent 1990 – 2013…………79

Figure 2.7. WON trophy weights from the Northern Channel Islands before and after the 2003 implementation of the network of marine protected areas in the region. The species included are California Halibut (A, n = 1074), Lingcod (B, n = 1125), Kelp Bass (C, n = 56), California Sheephead (D, n = 211), and Ocean Whitefish (E, n = 28) ………….…80

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Figure 2.8. WON trophy weights from the Northern Channel Islands during four periods, the last of which is the most recent decade following the 2003 implementation of the network of marine protected areas in the region. The species included are Vermilion Rockfish (A, n = 95), Bocaccio (B, n = 18), and Cowcod (C, n = 317)………………....81

Figure 3.1. Map of tagging sites off San Diego, California, including MPA boundaries ………………………………………………………………………….……………….105

Figure 3.2. Kelp Bass CPUE across primary CPFV charter sites (only includes CPFV charters when Kelp Bass were the target species)……………………………………...111

Figure 3.3. Barred Sand Bass CPUE across sites…………………………………...….112

Figure 3.4. Spotted sand bass CPUE by site. Data are based on private vessel trips only ………………………………….………………………………………….……………113

Figure 3.5. Boxplots of Kelp Bass CPUE by season. Data only include CPFV charters when Kelp Bass were the target species……………………………………………..…114

Figure 3.6. California Recreational Fisheries Survey (CFRS) estimates of monthly Barred Sand Bass landings aboard CPFVs based in Los Angeles and San Diego from Jan 2004 – Dec 2014………………………………………………………………………..………115

Figure 3.7. Boxplots of spotted sand bass CPUE by season. Data only include private vessel trips in San Diego Bay and Mission Bay when spotted sand bass were the target species…………………………………………………………………………….…….116

Figure 3.8. Kelp Bass length frequency distribution at the five primary sites off San Diego……………………………………………………………………………………117

Figure 3.9. Kelp Bass length frequency by northern San Diego sites………………….118

Figure 3.10. Barred Sand Bass length frequency by site…………………………….…119

Figure 3.11. Spotted sand bass length frequency by primary sites…………….….……120

Figure 3.12. Boxplots showing size of the top five demersal bycatch species caught in the primary La Jolla sites……………………………………………………………...……121

Figure 3.13. Mean (±95% CI) standard length of Kelp Bass (A), Barred Sand Bass (B), and Spotted Sand Bass (C) across primary sites during each of the three sampling years. The gray and black dashed lines mark the original minimum size limit in 2012, and new increased minimum size limit that was implemented on March 1, 2013 ………………122

Figure 4.1. Site map of the San Diego coastal area. Gray solid lines represent MPA boundaries, specifically the Matlahuayl and South La Jolla SMRs. The gray dashed line represents the U.S.-Mexico maritime border. Black dots in the three La Jolla sites represent fixed locations of acoustic receivers attached to subsurface moorings…..….144

Figure 4.2. Kelp Bass recapture frequency. Data only include the most recent recapture for each individual……………………………………………………………….……..146

Figure 4.3. Barred Sand Bass recapture frequency. Data only include the most recent recapture for each individual…………………………………………………….……..147

Figure 4.4. Spotted sand bass recapture frequency. Data only include the most recent recapture for each individual………………………………………………………...…148

Figure 4.5. Map of the array of acoustic receivers at the three La Jolla sites. The dark gray area represents land, and the light gray polygon represents the average spatial extent of the La Jolla kelp forest. Each black dot represents the location of a Vemco VR2W acoustic receiver attached to a subsurface mooring…………………………………….149

Figure 4.6. Logistic regression results showing the probability of tag retention over time among double-tagged Kelp Bass. The solid black line represents the predicted values of tag retention probability, and the dotted lines represent upper and lower 95% confidence limits……………………………..…………………………………………..…………150

Figure 4.7. Total distance traveled versus time at liberty by Kelp Bass……….....…...154

Figure 4.8. Frequency distribution of total distance traveled by Kelp Bass……………155

Figure 4.9. Total distance traveled versus time at liberty by Barred Sand Bass….……156

Figure 4.10. Total distance traveled versus time at liberty by Spotted Sand Bass…..…157

Figure 4.11. Boxplots showing depth utilization by month for both Kelp Bass and Barred Sand Bass in the three La Jolla sites……………………………………………………158

Figure 4.12. Boxplots showing median acceleration by month for both Kelp Bass and Barred Sand Bass in the three La Jolla sites……………………………………………159

Figure 4.14. The proportion of detections at each of the three La Jolla Sites for Kelp Bass that were initially tagged in the Matlahuayl SMR (a), in La Jolla (b), and in the South La Jolla SMR (c).….……………………………………………………………………….161

Figure 4.15. The proportion of detections at each of the three La Jolla sites for Barred Sand Bass that were initially tagged in the Matlahuayl SMR (a), in La Jolla (b), and in the South La Jolla SMR (c)…………………………………………………..……………..162

LIST OF TABLES

Table 1.1. All fifteen pelagic and coastal pelagic species ranked by sample size (N) within the 1966-2013 study period. Records from long-range fishing trips to central and southern Baja California are not included……………………………………………….41

Table 3.1. Number of CPFV tagging charters by site and sampling season. Sites are separated by location inside or outside of MPAs, which include Matlahuayl SMR (La Jolla Cove), South La Jolla SMR, and Swami’s SMCA (Encinitas)………………...…106

Table 3.2. Total number of fish tagged for each species at each site, excluding initial mortalities. Total numbers of recaptured individuals by species are also included…….107

Table 3.4. Initial post-release mortality rates for each of the three bass species as well as a combined estimate both for all other bycatch species and for all species. The total tagged- and-released represents the number of initial mortalities subtracted from the number caught………………………………………………………….….……………110

Table 4.1. Tagging summary data for each species by tagging site. Sites include Long Beach (LB), Newport Bay (NB), north San Diego County (NC), Matlahuayl SMR (M), La Jolla (LJ), South La Jolla SMR (SLJ), Mission Beach (MB), Mission Bay (MBy), Point Loma (PL), San Diego Bay (SDB), and Imperial Beach (IB) ……………...……145

Table 4.2. Kelp Bass recaptures for the following sites: Long Beach (L), San Clemente (S), north San Diego County (N), Matlahuayl SMR (M), La Jolla (LJ), South La Jolla SMR (SLJ), Mission Beach (MB), Mission Bay (MBy), Point Loma (PL), San Diego Bay (SDB), Imperial Beach (IB), and unknown location (U)…………………….…………151

Table 4.3. Barred Sand Bass recapture frequencies for the following sites: Long Beach (LB), Dana Point (DP), north San Diego County (NC), Matlahuayl SMR (M), La Jolla (LJ), South La Jolla SMR (SLJ), Mission Beach (MB), Mission Bay (MBy), Point Loma (PL), San Diego Bay (SDB), Imperial Beach (IB), and unknown location (Unk).….…152

Table 4.4. Spotted Sand Bass recapture frequencies for the following tagging and recapture sites: Newport Bay (NB), Mission Bay (MBy), and San Diego Bay (SDB). No individuals were recorded traveling between these sites……………….………………153

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Table 4.5. The total number of detections and percentage of detections for each species both inside and outside the two MPAs in La Jolla………………………….………..…163

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ACKNOWLEDGEMENTS

I thank my original advisor, Jeffrey B. Graham (1941-2011), for believing in me, and for taking a chance on me when conventional metrics (e.g., GRE and GPA) advised otherwise. His support and his guidance helped shape my research ideas into a community-based program that has bridged gaps between stakeholders, created a foundation for future collaborative research, and provided valuable results for California fisheries managers. I also thank the Graham family for their continued support for my research. Jeff loved his students like he loved his family, and we will all miss him forever.

I would like to thank my current advisor, Brice Semmens, who also took a chance by accepting me as a graduate student with little information to work with other than a single recommendation and a brief meeting. Brice provided an incredible amount of support during my transition from the Graham Laboratory, and his guidance since has led to the overwhelming success of our collaborative research program. I have improved in so many ways because of Brice, and I sincerely appreciate everything he has done along the way. I thank my committee members for their insight and ideas that have helped to significantly improve the quality of my research. Their doors were always open for me to ask questions and share ideas, and I am incredibly grateful for their guidance and advice.

My research would also not have been possible without the overwhelming support that I have received over the years from the California recreational fishing community.

Todd Keller (1963- 2003), a naturalist, fisherman, deckhand, and friend, first introduced me to this community when I was only 12 years old, and in doing so, he showed me my first steps on a path that I am still walking today. I also thank Fred Hall (1919-2000), Bart

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Hall, Mike Lum, Tim Baker, and the Hall family for their friendship and support for over two decades; Ken Franke (President, Sportfishing Association of California) for his friendship and incredibly extensive support for our research; Pete Nelson and

Collaborative Fisheries Research-West for funding our research program, the Coastal

Angler Tagging Cooperative; the Cooperative Institute for Marine Ecosystems and

Climate (CIMEC); the Graham family, Jerome and Miriam Katzin, Ralph Lewin, Edna

Bailey-Sussman, and Richard Rosenblatt for creating fellowships that helped to support me along the way; Dave Rudie, Samantha Harrod, John Valencia, and the San Diego

Oceans Foundation; James Nelson and Kelvin Nettleton for devoting an incredible amount of time and energy to the fieldwork; Tommy Gomes, for helping to facilitate the beginning of the tagging fieldwork; Tom Mason, Erica Jarvis, Heather Gliniak, Chuck

Valle, and Otis Horning (California Department of Fish and Wildlife) for their advice from a management perspective; Craig Heberer and the NOAA SWFSC; the Pfleger

Institute of Environmental Research (PIER) for their summer funding and research support; San Diego Anglers; San Diego Rod n Reel Club; San Diego Dive Club;

Oceanside Anglers; Oceanside Senior Anglers; Balboa Angling Club; American

Sportfishing Association (Mike Leonard and Gordon Robertson); the San Diego Port

Tenants Association; the Port of San Diego; the Coastal Conservation Association;

Friends of Rollo; Michael Fowlkes and Inside Sportfishing; Shimano; AFTCO (Bill

Shedd); United Anglers of Southern California (Dave Elm); Saltwater Bass Anglers;

Saltwater Bass Series; Ali Hussainy (BD Outdoors); Pat McDonnell (WON); Eric

Landesfeind; Eric Bent; Jeff Barr; Noah Ben-Aderet; Dwayne Patenaude; Pete Gray and

Rick Maxa; John Law (Wild West); Corey Sanden; Phil Friedman; Doug Kern; Vaughn

Podmore; Ron Ashby; John DeLaurentis; Howard Hada; Keith Patterson; John Nakatani;

Walter Heim; Dennis Burlason; Ed Corbin; Kim Burkhart; AA and Optimum Baits; Big

Hammer Swimbaits; Seaforth Landing; H&M Sportfishing; Point Loma Sportfishing;

Pierpoint Landing; Rick Oefinger (Marina Del Rey Sportfishing), John Klein (Qualifier

105), Joe Cacciola (Sea Star Charters), Greg Ewart (Ventura Sportfishing); Fred Huber and Steve Peterson (Daily Double, Point Loma, and Mission Belle); Kris Karpow (Sea

Watch); Ron Baker (Point Loma); Paul Fischer (Outer limits); Billy Treviranus; Eric

McCully; Dave Mandagie; Gerry Mahieu; Bill Hoksted; Chris Barrick; Harry Orgovan;

Dave Valadez; Ed Smith; John Beerling; Jon Kuch; Josh Dunlap; Karl Erbacher;

Mike Ryba; captains and crews of the Outer Limits, Sea Watch, Daily Double, Point

Loma, Chubasco II, Mission Belle, Sea Star, New Seaforth, Dolphin II, Premier,

Fisherman III, and Toronado; and hundreds of other volunteer anglers!

My family and friends have given me the encouragement and support that has helped me progress every day, and my appreciation for their love could never be fully expressed in words. In addition, my Scripps family (labmates, fellow students, faculty, and staff) has been unbelievably helpful during my time at SIO. My volunteer assistants,

Amanda Barker, Jason Ho, Ashley Rankin, Tom Wooten, Ryan Woo, Elizabeth Cerny-

Chipman, Andrea Hatlen, Stephen Huerta, Bianca Andrade, Emily Botts, and Jenny Dao, all spent hundreds of hours to help create the historical database. I sincerely appreciate all of my dive assistants, especially Rachel Morrison (1987-2014), who always volunteered to help regardless of her own schedule.

I would never have entered the Scripps community without one very specific thing, which was a letter of recommendation from Stuart Sandin to my former advisor,

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Jeff Graham. Jeff was undecided about accepting me as a graduate student, and Stuart’s letter tipped the balance in my favor. Stuart Sandin and Jen Smith gave me a tremendous amount of encouragement and insight, and I will always be grateful for their help.

Finally, Alayna Siddall not only spent more time helping with my cooperative research program than anyone else, she has given me complete, unconditional love and support. I am forever hers.

Chapter 1, in part is currently being prepared for submission for publication of the material. Barker, Amanda; Ho, Jason; and Semmens, Brice. The dissertation author was the primary investigator and author of this material.

Chapter 2, in part is currently being prepared for submission for publication of the material. Semmens, Brice. The dissertation author was the primary investigator and author of this material.

Chapter 3, in part is currently being prepared for submission for publication of the material. Semmens, Brice. The dissertation author was the primary investigator and author of this material.

Chapter 4, in part is currently being prepared for submission for publication of the material. Semmens, Brice. The dissertation author was the primary investigator and author of this material.

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VITA

2002 Bachelor of Science, University of California Santa Barbara

2006 Master of Science, California State University Long Beach

2015 Doctor of Philosophy, University of California San Diego

PUBLICATIONS

Lowe, C.G., K.M. Anthony, E.T. Jarvis, L.F. Bellquist, and M.S. Love. 2006. Site fidelity of characteristic fish species at offshore petroleum platforms in the Santa Barbara Channel. MMS OCS Study 2004-2007 Annual Report. California State University Long Beach, Long Beach, California. MMS Cooperative Agreement Number 1435-01-04-CA- 34196.

Bellquist, L.F., C.G. Lowe, and J.E. Caselle. 2008. Fine-scale movement patterns, site fidelity, and habitat selection of Ocean Whitefish (Caulolatilus princeps). Fisheries Research 91(2-3): 325-335.

Lowe, C.G., K.M. Anthony, E.T. Jarvis, L.F. Bellquist, and M.S. Love. 2009. Site fidelity and movement patterns of groundfish associated with offshore petroleum platforms in the Santa Barbara Channel. Marine and Coastal Fisheries: Dynamics, Management, and Ecosystem Science 1: 71-89.

FIELDS OF STUDY

Major Field: Marine Biology

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ABSTRACT OF THE DISSERTATION

A historical perspective of California recreational fisheries using a new database of “trophy” fish records (1966-2013), combined with analyses of three species in the genus Paralabrax

Lyall F. Bellquist

Doctor of Philosophy in Marine Biology

University of California, San Diego 2015

Professor Brice Semmens, Chair

Recreational fishing in the United States promotes economic growth, cultural traditions, and environmental stewardship, but can also negatively impact fish stocks when fisheries are mismanaged. The scale of recreational fisheries is often underestimated due to difficulties in monitoring their complex, dynamic, and often expansive nature. In California, the documented history of recreational fishing dates to the late 1800s, and the industry has grown since. In 2013, California marine anglers took

5.3 million fishing trips, landed 8.4 million fish, and contributed $2.8 billion to the economy. Despite this extensive history and the well developed fisheries of the present day, we have limited quantitative historical fisheries data as well as measurements of current fish population dynamics. We developed a new historical database of “trophy”

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fish catch records (1966-2013) that could provide early signs of fisheries overexploitation, given that these large size classes are generally the most vulnerable to the impacts of fishing. We also founded the Coastal Angler Tagging Cooperative in partnership with the recreational fishing community to conduct demographic analyses of three of the most important recreationally targeted inshore species in the genus

Paralabrax. Historical analyses indicate that “trophy” sizes of pelagic and coastal pelagic species are generally showing little signs of decline, while most decreasing trends are exhibited by demersal species. Rockfishes in particular (Sebastes sp) have shown both temporal and spatial declines in trophy size. However, the demersal species are also the most likely to benefit from the recently implemented statewide network of Marine

Protected Areas (MPAs), and many rockfishes have begun to show positive trends since new management policies were adopted in 2001. Our analyses of Paralabrax populations indicate a positive response to the recently implemented increase in minimum size limits, as well as spatial and seasonal differences in both length frequency and catch-per-unit- effort, depending on the species. We documented a new spawning aggregation site for

Barred Sand Bass (P. nebulifer), and we highlight the virtual absence of traditional spawning behavior for this species in both 2013 and 2014. Collaborations between the scientific and fishing communities can be highly successful, and we encourage this in future research efforts.

CHAPTER 1: Long-term changes in “trophy” sizes of pelagic and coastal pelagic fishes among California recreational fisheries (1966-2013)

Abstract

The biological and economic importance of recreational fisheries is being increasingly recognized in the United States. In California, recreational fisheries contribute a significant amount to coastal economies, and two of the primary drivers of fisheries economics in the state are pelagic and coastal pelagic species during summer and fall. While traditional catch frequency (landings) datasets exist for these species, size specific information is limited, especially for the largest size classes that are the most sensitive to exploitation. One previously overlooked data source, Western Outdoor News, is a popular weekly fishing and hunting newspaper that has been published in California since 1953. Each weekly issue includes a list of the largest fish caught at each sportfishing landing along the California coast each week. We developed a database that contains all of these “trophy” catch records from the period of 1966-2013. The database contains 48,750 individual trophy records of 94 total fish species spanning the region from the Revillagigedos Islands, Mexico to southern Oregon. Species were grouped into demersal, pelagic, and coastal pelagic categories. Coastal pelagic species were defined as highly mobile species that maintain an affinity for coastal waters, whereas pelagics were considered highly mobile while primarily inhabiting offshore waters. Time series analyses show long-term declines in trophy size in only two of the fifteen pelagic and coastal pelagic species (Yellowfin Tuna and Striped Bass). Three species showed long-

term increases in size (Pacific Bluefin Tuna, White Seabass, and California Yellowtail), and pelagic species were generally more variable while coastal pelagics were more stable. Our results show that a combination of oceanographic processes, natural history characteristics, recreational fishing, and fisheries regulations all play a role in trophy size dynamics, but the relative influences of each vary between species.

Introduction

The historical record of recreational fishing in California dates back to the late

1800s, and the recreational fishing industry developed significantly throughout the twentieth century. Fisheries perspectives have also evolved, from the idea that fisheries are an “inexhaustible” resource (Huxley 1884) to the early recognition of fishing impacts

(Garstang 1900), and more recently into the widespread documentation of declines on multiple scales (e.g., Pauly et al. 1998; Jackson et al. 2001; Myers and Worm 2003;

Pandolfi et al. 2003; Sala et al. 2004; Pauly and Palomares 2005; Sáenz-Arroyo et al.

2005; McGinnis 2006; McClenachan 2009a,b). While commercial fishing is often the primary focus in fisheries impact analyses, the importance of recreational fisheries is being increasingly recognized in terms of both overall landings (Schroeder and Love

2002; Dotson and Charter 2003; Coleman et al. 2004; Cooke and Cowx 2004; Arlinghaus and Cooke 2005; Arlinghaus et al. 2007) and contribution to local and state economies

(NMFS 2011). In 2013 alone, California recreational anglers took 5.3 million recreational fishing trips in marine waters, contributing approximately $2.8 billion to the economy, and 70% of this occurred in the disproportionately small region of southern California

(CDFW 2014). Clearly, recreational fishing in California is highly important from both biological and socioeconomic perspectives.

The primary economic drivers of marine recreational fisheries in California are pelagic and coastal pelagic fishes from May to October (Hill & Barnes 1998, Hill &

Schneider 1999). Catch rates for these species strongly influence angler behavior (Doston

& Charter 2003), especially when total fishing effort peaks during the summer. Because these species are highly sought after and primarily caught offshore, they not only drive both the industry economics and offshore fishing effort, but they can also shift fishing effort away from inshore demersal species. Thus, pelagic and coastal pelagic fishes in

California are extremely important as top predators in a complex ecosystem, drivers of fishing economies, and the health of their populations supports the stability of demersal fishes that are more susceptible to overfishing.

Pelagic and coastal pelagic species are generally thought to be more resilient to fishing than demersal species due to differences in life history characteristics. However, some pelagic and coastal pelagic species have shown signs of decline during the last several decades. For example, Ragen 1990 (later summarized in Dayton and MacCall

1992) found that historical trophy record weights of two coastal pelagic species in southern California, White Seabass (Atractoscion nobilis) and California Yellowtail

(Seriola lalandi), declined between 1884 and 1940 as the fisheries for these species developed. In addition, commercial fishing for Striped Bass (Morone saxatilis) in the San

Francisco Bay and Sacramento River Delta was banned in 1935 due to signs of population decline (Stevens et al. 1985), and White Seabass were commercially

overfished in 1970s and 1980s (Vojkovich and Reed 1983). More recently, Maunder et al. (2014) concluded that Pacific Bluefin Tuna have been severely overfished, which has led to proposed changes in California recreational fishing regulations for this species.

Clearly, even species with favorable life history characteristics can be impacted, but data specific to trophy sizes that can allow early identification of these impacts are rare.

Throughout the history of recreational fisheries in California, size-selective fishing has been the common model, and the impacts of size-selection in general have been well documented (e.g., Ricker 1981, Stokes et al. 1993, Hutchings 2000, Law 2000,

Palumbi 2001, Hamilton et al. 2007). However, the largest size classes within a population are the most sensitive to size selective exploitation due to their higher recreational value, yet slower growth rates. The initial impact of sustained size-selective fishing is thus a decrease in the abundance of the largest individuals (Pandolfi et al.

2003), yielding a decrease in the size structure of a population (Pauly et al. 2002, Froese

2004). If this trend continues within a targeted population, the size structure can become impacted to the point at which fishers shift to the next largest species. This systematic process of fishing from higher to lower trophic levels over time can ultimately result in the frequently cited process of fishing down the food web (Pauly et al. 1998). While this process has remained controversial on a global scale in terms of both trophic dynamics

(Essington et al. 2006, Branch et al. 2010) and price/value of higher trophic levels (Sethi et al. 2010), evidence of fishing down food webs has been documented on local and regional scales (e.g., Stergiou & Koulouris 2000, Sala et al. 2004; McClenachan 2009a,b;

Mumby et al. 2012). Because this process begins with a decrease in the abundance of the

largest individuals, data that focus specifically on the dynamics of these largest size classes, i.e. trophy fishes (e.g., Powers et al. 2013), can be essential for gaining early indications of fishery over-exploitation (Yunne-Jai Shinn 2005).

We developed a new long-term database of recreational trophy fish catch in

California based on a popular publication called Western Outdoor News (WON).

Published weekly since 1953, this newspaper reports information specific to recreational fishing and hunting on the west coast of the United States, primarily California.

Beginning in 1966, the “Whopper of the Week” was published in each weekly issue. This is a list of the largest fish caught at each sportfishing landing each week along the

California coast. Each record contains the species, size (usually weight), approximate date and location caught, vessel, port, and angler information. Other existing datasets, such as the Pacific Fisheries Environmental Laboratory (PFEL) commercial fisheries database or the California Commercial Passenger Fishing Vessel (CPFV) logbook database, do not contain information specific to individual fish size. Both the Marine

Recreational Fisheries Statistics Survey (MRFSS) and the California Recreational

Fisheries Survey (CRFS) database contain size-specific information, but these two databases are not statistically comparable due to changes in data collection methodology, and the newer CRFS database only dates back to 2003. The WON database (1966-2013) thus represents the only source of data that could potentially be used as an early indicator of fisheries impacts for most of the primary targeted fishes in California. Our goal was to quantify long-term trends in trophy sizes of fishes in the context of both oceanographic conditions and fishing effort, with the null hypotheses that 1) no changes in trophy size

have occurred over time, and 2) that trophy fish sizes show no correlation with environmental or anthropogenic factors. Pelagic and coastal pelagic species are the focus of this study, and demersal species are the subject of an ongoing study.

Methods

Data collection

We acquired weekly issues of WON from 1966-2013 using the lead author’s personal collection as well as partial collections from other subscribers. In 2013, a digital edition of WON was created for subscribers, but this new format still requires manual data transcription. We collected data from all WON issues by taking photographs of the page containing the “Whopper of the Week” list for each WON issue, and we then transcribed each list into a database. This database contains fish species, length/weight, catch location, date, vessel name, and home port. We further subdivided these data into vessel categories (private, Commercial Passenger Fishing Vessel, and 6-pack charter boats), catch location categories (embayment, mainland coast, island, offshore bank, county, north/south of both the Mexico border and Point Conception), and species groups

(elasmobranch or teleost, followed by pelagic, coastal pelagic, or demersal). However, pelagic and coastal pelagic species are the primary focus of this paper. We cross- referenced the records with the original photographs to ensure data accuracy. Catch records from long range fishing trips (defined as >100 miles from home port), virtually all of which were in Mexican waters, were not included in the analyses because these were not considered to be representative of California coastal fisheries.

Database Validation

We first measured the level of agreement between this dataset and other data sources that are currently accepted by fisheries managers. We conducted a correlation meta-analysis using the ‘metacor’ package (Laliberté 2015) within the statistical platform

R (R Development Core Team 2008) to compare the percent occurrence in the WON database with two other landings databases: the California Department of Fish and

Wildlife recreational logbook data and National Oceanic and Atmospheric

Administration (NOAA) Pacific Fisheries Environmental Laboratory (PFEL) commercial landings data. The R package ‘metacor’ calculates an individual correlation coefficient and 95% confidence interval for each species comparison, and also calculates one global correlation coefficient for all comparisons. The species are then ranked and plotted relative to a zero-correlation line. We used this approach for the years encompassed by all three datasets (1966-2008), which is likely a conservative approach given that overall landings trends are often similar between different datasets, but the exact annual timing may differ slightly, yielding non-significant correlations in some cases. The three pairwise comparisons (Logbook – PFEL, PFEL – WON, Logbook – WON) did not always include the same species because not all species in this study are commercially fished.

Statistical Analyses

We used a Bayesian approach to analyze temporal trends in trophy fish sizes within the statistical platform R. Models were run using JAGS (Plummer 2003), which

was called from R using the package ‘R2jags’ (Su & Yajima 2012), and model convergence was assessed with trace plots and the Gelman—Ruben R statistic (Gelman

& Rubin 1992). We first plotted temporal trends in fish size using boxplots to illustrate the distribution of the raw data. We then used a Bayesian state-space model to describe the overall trend as well as to estimate process (population stochasticity) and observation

(estimation) errors. Both the process and observation models given here represent the complete model used to describe temporal trends:

Process:

Observation:

where represents the mean size (weight) of species i in year t, and represents the normally distributed process error. For the observation model, denotes the observed size (weight) for species i in year t, and denotes the observation error, which is assumed to be normally distributed around 0 with a variance matrix . The Monte Carlo

Markov chain (MCMC) method was applied with 25,000 iterations and a 10,000 iteration

burn-in period, and convergence was assessed using the Gelman-Rubin diagnostic

(Spiegelhalter et al. 2003). This model allows for multiple observations in a time series as well as missing observations.

After fitting the long-term trends in trophy size for each species, we compared these trends to multiple environmental processes that are known to influence the

California coastal region. We used cross-correlation analysis in R to compare mean annual trophy size with five different environmental variables: the Multivariate El Niño-

Southern Oscillation (ENSO) index, the Pacific Decadal Oscillation (PDO) index

(Mantua and Hare 2002), the North Pacific Gyre Oscillation (NPGO) index (Di Lorenzo et al. 2008), the Scripps Institution of Oceanography (SIO) pier water temperature data

(5m depth), and the PFEL southern California upwelling index (33oN, 119oW: http://www.pfeg.noaa.gov/products/PFELData/upwell/daily/p11dayac.all). We also used this approach to compare mean annual trophy size with fishing effort (angler-days) obtained from California Department of Fish and Wildlife recreational Commercial

Passenger Fishing Vessel (CPFV) logbook records. Alpha levels were changed using a

Bonferroni adjustment to α = 0.01. The cross correlation analysis tests for potential lags in correlations between fish size and each environmental variable. While these processes have been shown to influence numerous local fish populations (Allen et al. 2003, Jarvis et al. 2004), it remains unclear how these factors might influence the largest size classes of fishes. We pooled the data for each species across the study region to provide overall statewide estimates.

Results

Database description

The WON database contains trophy fish records from 1966-2013, and is composed of 48,750 records of 94 total species caught between the Revillagigedos

Islands, Mexico and the California-Oregon border. However, 499 records from long- range fishing trips into waters off southern Baja California were not considered to be representative of California fisheries, and thus we did not include them in the analyses.

The study area encompasses the entire California coastline, although 63% of the data were from the Southern California Bight. The remaining records were primarily from the

San Francisco Bay area. We separated species into broad habitat-based groups, but the pelagic and coastal pelagic species are the focus of this paper.

Database validation

The two primary sources for long-term landings in the state are the California

Dept. of Fish and Wildlife recreational CPFV logbook database and the PFEL commercial landings database. Comparisons between these two databases and the WON database showed evidence for positive correlations across the majority of the focal species (Fig. 1). We conducted analyses on fifteen pelagic and coastal pelagic species, but each pairwise comparison did not necessarily contain all 15 species due to lack of coverage in one of the three databases used. For example, Striped Marlin are not commercially targeted, so we did not include this species in pairwise comparisons that contained the commercial database. Logbook-commercial comparisons showed positive

correlations for the majority (8 of 13) of species (Fig. 1a), WON-commercial showed significant positive correlations for only 3 of 13 species (Fig. 1b), and WON-logbook comparisons were significant for 12 of 15 species (Fig. 1c, although one of these species,

Striped Bass, showed a significant negative correlation). The commercial and recreational logbook databases, which have a 62% agreement in the number of species showing significant positive correlations, are the two primary sources of landings data used by fisheries analysts. The WON database has a 73% agreement (Striped Bass not included) with the logbook database. This suggests that the new WON database is a comparable or better representation of recreational landings in California than other data sources that are currently accepted by fisheries managers.

Temporal Trends

Pelagic and coastal pelagic fishes and their respective sample sizes and temporal coverage are shown in Table 1. Pelagic teleosts exhibited a high degree of variability in mean weight over time (Fig. 2 and 3). Records for Yellowfin Tuna (Thunnus Albacores),

Pacific Bluefin Tuna (T. orientalis), Bigeye Tuna (T. obesus), Striped Marlin (Kajikia audax), Common Dolphinfish or Common Dolphinfish (Coryphaena hippurus), and

Opah (Lampris guttatus) were primarily from south of Point Conception, whereas Pacific

Albacore (Thunnus alalunga) records were more evenly distributed between the southern and northern regions. These seven species constituted 98% of the data for all pelagic teleosts combined. Most of the pelagic teleosts showed variable but stable trends in size over time, although some species were not always present in the landings record. Bigeye

Tuna (Fig. 2a) were only caught between the early 1980s and 2000 (this is supported in the CDFW logbook data), with no apparent trend in weight over time. However,

Yellowfin Tuna approximately halved in size over four years in the late 1980s from approximately 40kg to 20kg, and have remained at this level since the decline (Fig. 2b).

Meanwhile, Bluefin Tuna trophy weight doubled in size from approximately 10kg to

20kg during the same time period, and has since remained at this level (Fig. 2c). Pacific

Albacore showed annual variation in size, but generally appear stable at approximately

15kg during the entire study period (Fig. 2d). Striped Marlin, Dolphinfish, and Opah showed high variability with no trends in weight over time (Fig. 3a-c). Other pelagic fishes in the database, such as wahoo (Acanthocybium solandri), bullet tuna (Auxis rochei), Pacific skipjack (Katsuwonus pelamis), and shortbill spearfish (Tetrapterus angustirostris) each had too few records for robust analyses.

Records of pelagic elasmobranchs, which consisted of Shortfin Mako (Isurus oxyrhincus) and Common Thresher Sharks (Alopias vulpinus), were mostly from the

Southern California Bight (SCB), and average sizes each year were highly variable for both species (Fig. 3d,e). The recreational fishery for Common Thresher Sharks is relatively young, so most of the data for this species began in the early 1990s, whereas records for Shortfin Mako were more consistent throughout the study period. Mako and thresher Sharks both showed high variability in weight over time, with no consistent trend apparent in the data.

Coastal pelagic teleosts consisted of White Seabass (Atractoscion nobilis),

California Yellowtail (Seriola landali), Pacific Barracuda (Sphyreana argentea), and

Pacific Bonito (Sarda chiliensis) (Fig. 4a-d) in the Southern California Bight. Striped

Bass (Morone saxatilis) and Chinook Salmon (Oncorhynchus tshawytscha) were caught primarily north of Point Conception (Fig. 4e,f). Among these six coastal pelagic species, only one (Striped Bass) showed a sustained decline in size over time. White Seabass showed a decrease in size during the 1970s, but have increased consistently since 1985.

California Yellowtail show a similar pattern with sizes also decreasing in the 1970s, and subsequently increasing during the last three decades (Fig. 5b). Pacific Barracuda, Pacific

Bonito, and Chinook Salmon do not show any signs of long-term declines in size.

Chinook Salmon seem to show four distinct domains within the time series, each progressively larger than the previous, except the last period from 2009-2013, which begins low but increases consistently each year after 2009. Striped Bass declined in size by almost half during the 1990s, and then again in 2012-2013.

Correlations with environmental indices and fishing effort

We conducted cross correlation analyses between trophy size and five different environmental factors (ENSO, PDO, NPGO, SIO pier water temperature at 5m depth, and southern California upwelling) as well as one anthropogenic factor (fishing effort in terms of CPFV angler-days), for each of the 15 species. Time lags of up to 13 years were included in the analysis. Figures 5-9 show cross correlation results for each of the fifteen species. Each figure consists of 6 panels (a-f), which represent correlations with the MEI index (a), PDO index (b), NPGO index (c), water temperature at SIO pier at 5m depth

(d), southern California upwelling index (e), and recreational fishing effort (f). Alpha

levels were adjusted using a Bonferroni correction for each species (α = 0.01), and gray horizontal lines above and below zero represent confidence intervals. Significant correlations with at least two environmental factors were found for every one of the 15 species, and 14 of these species showed significant correlations with at least three of the environmental factors. Figures 5-9a show significant correlations within at least one lag step with the MEI index for 11 out of 15 species. Figures 5-9b show significant correlations within at least one lag step with the PDO index for 13 out of 15 species.

Figures 5-9c show significant correlations within at least one lag step with the NPGO index for 13 out of 15 species. Figures 5-9d show significant correlations within at least one lag step with the SIO pier water temperature for 12 out of 15 species. Finally, figures

5-9e show significant correlations within at least one lag step with the southern California upwelling index for 12 out of 15 species. In addition to oceanographic time series, annual recreational CPFV fishing effort show correlations within at least one lag step for 12 out of 13 species (Opah and Common Thresher were not included because defining effort for these species is somewhat ambiguous).

Discussion

The size structure dynamics of trophy fishes in California are likely influenced by a combination of physical, biological, and anthropogenic factors. The pelagic and coastal pelagic environments in this region are characterized by diverse community assemblages with a broad range of life history traits, dynamic recreational and commercial fisheries, fisheries management efforts that continually adapt to new biological and economic

information, and highly complex oceanographic processes. As a result, we see a range of trends among the fifteen species included in this study: decreases in trophy size were exhibited by Yellowfin Tuna and Striped Bass, while increases were measured for

Bluefin Tuna, California Yellowtail, and White Seabass. The remaining ten species showed no long-term trends but the pelagic species were more highly variable, whereas the coastal pelagics were more stable.

The natural history of each species plays a significant role in the dynamics of trophy fishes. Differences in growth rates, reproductive behavior, recruitment patterns, early life history, age at maturity, fecundity, and movement patterns all are known to affect population dynamics of marine fishes. The 15 focal species in this study have relatively fast growth rates, early ages at maturity, high fecundity (with the exception of the elasmobranchs), and are either highly migratory or highly mobile, with areas of movement extending far beyond California waters in many cases. These characteristics suggest a relatively low susceptibility to localized recreational fisheries impacts, which may explain the limited signs of decline measured in this study. However, the differences in variance between the pelagic and coastal pelagic species could be attributed to both sample size and species behavior, specifically movement patterns. The pelagic species generally had lower sample sizes in the database, which likely accounts for some of the increased variability in this group. The pelagic species are also generally highly migratory, some of which have ranges that extend across the entire north Pacific (e.g.,

Bayliff 1994, Kimura et al. 1997). The coastal pelagics are highly mobile, but have a more restricted range, primarily from southern California to central Baja California (e.g.,

Collins & MacCall 1977, Squire 1987). Pelagic fishes are thus likely influenced by a broader range of external processes, which may contribute to the more variable trophy sizes measured in this study.

The fisheries for pelagic and coastal pelagic species are highly dynamic. Most species are targeted both commercially and recreationally, but some (Striped Marlin,

Common Dolphinfish, Pacific Barracuda, and Striped Bass) are only targeted recreationally. Past studies have attributed signs of population decline to overfishing in only four of the 15 species focal species in this study (White Seabass, Pacific Bluefin

Tuna, Striped Bass, and Chinook Salmon), and these were primarily linked to commercial rather than recreational fisheries: White Seabass populations declined from the 1950s through the 1970s (Vojkovich and Reed 1983), but the population has been in recovery since the ban on inshore commercial gill nets in 1994 (Allen et al. 2007,

Pondella & Allen 2008); Pacific Bluefin Tuna declines are primarily attributed to commercial overfishing by western Pacific fleets (Maunder et al. 2014); Striped Bass declines were attributed to both commercial overfishing in the 1920s, which was banned in 1935, as well as diversion of water from the Delta in more recent years (Stevens et al.

1985, Kohlhorst 1999); and Chinook Salmon declines have been attributed to a combination of commercial overfishing, loss of spawning habitat, and anomalously poor oceanographic conditions in some years (Yoshiyama et al. 1998, Lindley et al. 2009).

Both Aalbers et al. 2004 and Aalbers 2008 have suggested a potential for recreational impacts on White Seabass, especially given that recreational landings are sometimes greater than commercial landings for this species (Crooke 2006), and landings peak

during the spawning season at known spawning sites. This is a situation that should be monitored closely because White Seabass have already shown long-term impacts of fishing in the past.

Oceanography is highly variable across multiple spatial and temporal scales in the

California pelagic environment (McClatchie 2014). While our study focuses on trophy fish size, Jarvis et al. (2004) focused on overall recreational landings (numbers of fish), and found significant correlations with environmental variables for 27 out of 45 species

(36 total significant correlations). The environmental variables ranked by order of importance were upwelling (53%), PDO (33%), offshore sea surface temperature (11%), and the ENSO index (3%). Our study yields somewhat similar rankings, with PDO,

NPGO, and upwelling showing the highest number of species correlations (each index showed significant correlations for 13/15 species), followed by water temperature (12/15 species), and lastly the ENSO index (11/15 species). Two species that showed generally opposite trends were Yellowfin Tuna and Pacific Bluefin Tuna, likely due to their different offshore habitats: yellowfin inhabit warmer waters (~17-26oC) while bluefin tend to prefer slightly cooler waters (~12-22oC), which yields a latitudinal separation in area use between the two species, with an area of overlap from the Southern California

Bight to central Baja California (Block et al. 2011). Peak years of yellowfin landings tend to coincide with peak years in water temperature, while peak years in bluefin landings tend to coincide with years of lower water temperature (although 2014 was an anomalous year with high water temperatures and high catches of both yellowfin and Bluefin Tuna).

In the WON database, yellowfin decreased in trophy size by approximately 50% from

1978-1987, but bluefin doubled in size during this same approximate time period. These two species also showed virtually opposite correlations with all five oceanographic indices. The 1978-1987 period represents a known warming event, and it is possible that larger individuals that inhabit cooler waters (e.g., Pacific Bluefin Tuna) may favor these warmer conditions while larger individuals that already inhabit warmer waters (e.g.,

Yellowfin Tuna) may not favor additional warming. This may explain the pattern observed in the WON database, and suggests that oceanography plays an important role in the size structure of yellowfin and Bluefin Tuna on a local scale. Striped Bass, the only coastal pelagic species that showed a decline during the study period, showed the strongest correlation with PDO, and almost no correlation with recreational fishing effort.

This species is primarily caught in San Francisco Bay, and is largely influenced by physical conditions in their spawning areas of the Sacramento-San Joaquin River Delta.

Decreases for this species coincide with decreases in freshwater input via the Delta

(Stevens et al. 1985), which suggests that river dynamics, water diversion, and long-term environmental processes (e.g., PDO) may impact the species more than recreational fishing. White Seabass and California Yellowtail each showed similar patterns of correlation with oceanographic indices, both showing a negative correlation between trophy size and temperature. However, mean annual water temperature at Scripps pier has not shown any consistent trend since the late 1990s, but sizes of both Yellowtail and

White Seabass have increased during that time, which suggests other fisheries interactions might be at play. Also beginning in the late 1990s, braided fishing line became popular within the recreational fishing industry. This may have allowed anglers

to land larger fish in kelp forest habitats than was previously possible because braided line cuts through kelp stipes much more easily than monofilament, meaning fewer trophy-sized fish become entangled and lost in the kelp.

While this study only focuses on species that are more resilient to fishing impacts, these fifteen species are among the most heavily targeted species in California, and they do not appear to be following the systematic and widespread declines that are often documented in the primary literature (Pauly et al. 1998, 2005; Jackson et al. 2001; Myers and Worm 2003; Pandolfi et al. 2003; Sala et al. 2004). One possible explanation in addition to natural history characteristics may be related to fishing effort in California. It is generally assumed that fishing effort has increased over decadal time scales simply because the human population has grown. However, some studies show a more stable trend in total fishing effort during the study period (Hill and Schneider 1999, Dotson and

Charter 2003). Records from the California Department of Fish and Wildlife indicate that state per capita annual fishing license sales have steadily declined by approximately 72% since 1970 (Fig. 10). The CRFS database and CDFW fisheries forum reports do not show any consistent trends in total annual effort during the last decade. There is thus a possibility that recreational fishing effort may not be increasing over time as is often assumed. In addition, while our study does span a period of almost 5 decades, it is likely that some of our focal species were already impacted before 1966, the first year in the database (e.g., Ragen 1990, Dayton and MacCall 1992, Dayton et al. 1998). Clearly, natural history, oceanography, and fisheries dynamics all play important roles in influencing the trophy sizes of pelagic and coastal pelagic fishes in California, and the

WON database is a useful new source of information that can increase our understanding of California fish population dynamics. Demersal species in the WON database are the subject of another study that will focus on trophy dynamics among a species guild with life history characteristics that are different from pelagic and coastal pelagic species.

Acknowledgements

I would like to acknowledge my co-authors, Brice Semmens, Amanda Barker, and

Jason Ho for their assistance with building the WON database.

Species Scientific name Species group Years N Oncorhynchus Chinook Salmon tshawytscha coastal pelagic 1966-2013 6045 CA Yellowtail Seriola landali coastal pelagic 1966-2013 5574 White Seabass Atractoscion nobilis coastal pelagic 1966-2013 4184 Albacore Thunnus alalunga pelagic 1968-2013 1526 Striped Bass Morone saxatilis coastal pelagic 1968-2013 1142 Bluefin Tuna Thunnus orientalis pelagic 1968-2013 694 Pacific Barracuda Sphyraena argentea coastal pelagic 1966-2012 565 Yellowfin Tuna Thunnus albacares pelagic 1968-2013 536 Pacific Bonito Sarda chiliensis coastal pelagic 1966-2013 389 Bigeye Tuna Thunnus obesus pelagic 1969-2005 264 Common Dolphinfish Coryphaena hippurus pelagic 1968-2013 138 Striped Marlin Kajikia audax pelagic 1968-2013 133 Shortfin Mako Isurus oxyrinchus pelagic 1974-2010 120 Common Thresher Alopias vulpinus pelagic 1971-2013 85 Opah Lampris guttatus pelagic 1968-2013 45 Total 21,440

Figure 1.3. Time series plots of trophy fish weight showing model output (white line) and model error (grey region) overlaid onto boxplots of the raw data for Dolphinfish (dorado) (A, n = 138), Striped Marlin (B, n = 133), Opah (C, n = 45), Shortfin Mako Shark (D, n = 120), and Common Thresher Shark (E, n = 85).

Figure 1.4. Time series plots of trophy fish weight showing model output (white line) and model error (grey region) overlaid onto boxplots of the raw data for White Seabass (A, n = 4184), California Yellowtail (B, n= 5574), Pacific Bonito (C, n = 389), Pacific Barracuda (D, n = 565), Striped Bass (E, n = 1142), and Chinook Salmon (F, n = 6045).

Figure 1.5. Cross correlation results for three tuna species: Pacific Bluefin Tuna (black line), Yellowfin Tuna (gray line), and Pacific Albacore (dashed line). Bigeye Tuna were not included due to limited years of catch (primarily only 1981-2000). Reference lines are shown at a correlation value of zero as well as confidence limits with an adjusted alpha level of 0.01. Panels show correlations with the MEI index (A), PDO index (B), NPGO index (C), SIO pier water temperature at 5m depth (D), southern California upwelling index (E), and CPFV fishing effort in angler-days (F).

Figure 1.6. Cross correlation results for three other pelagic species: Common Dolphinfish (black line), Striped Marlin (gray line), and Opah (dashed line). Reference lines are shown at a correlation value of zero as well as confidence limits with an adjusted alpha level of 0.01. Panels show correlations with the MEI index (A), PDO index (B), NPGO index (C), SIO pier water temperature at 5m depth (D), southern California upwelling index (E), and CPFV fishing effort in angler-days (F).

Figure 1.7. Cross correlation results for four coastal pelagic species: California Yellowtail (black line), Pacific Bonito (gray line), Pacific Barracuda (black dashed line), and White Seabass (gray dashed line). Reference lines are shown at a correlation value of zero as well as confidence limits with an adjusted alpha level of 0.01. Panels show correlations with the MEI index (A), PDO index (B), NPGO index (C), SIO pier water temperature at 5m depth (D), southern California upwelling index (E), and CPFV fishing effort in angler-days (F).

Figure 1.8. Cross correlation results for two pelagic elasmobranch species: Common Thresher (black line) and Shortfin Mako (gray line). Reference lines are shown at a correlation value of zero as well as confidence limits with an adjusted alpha level of 0.01. Panels show correlations with the MEI index (A), PDO index (B), NPGO index (C), SIO pier water temperature at 5m depth (D), southern California upwelling index (E), and CPFV fishing effort in angler-days (F).

Figure 1.9. Cross correlation results for two coastal pelagic species from central and northern California: Chinook (king) Salmon (black line) and Striped Bass (gray line). Reference lines are shown at a correlation value of zero as well as confidence limits with an adjusted alpha level of 0.01. Panels show correlations with the MEI index (A), PDO index (B), NPGO index (C), SIO pier water temperature at 5m depth (D), southern California upwelling index (E), and CPFV fishing effort in angler-days (F).

Figure 1.10. Number of total resident annual fishing licenses sold in California (gray line), and per capita annual resident fishing licenses (x 100) sold in California (black line) from 1970 – 2013. Non-resident annual license statistics were not included due to relatively negligible values. Short-term license statistics were also not included due to inconsistent availability of several different license types since 1970.

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Chapter 2: Temporal and spatial dynamics of “trophy” sized demersal fishes off the

California coast, 1966-2013

Abstract California recreational fisheries are biologically and economically important, and depend heavily on demersal fish species, especially during winter and spring months.

While many of these demersal fishes have shown signs of population decline on decadal scales, the size-selective nature of fishing generally impacts population size structure before these overall population declines are evident. However, long-term size-specific information, especially for the largest size classes, is rare for most California fishes. Our ability to gain an early warning of fishery overexploitation is thus limited. We developed the Western Outdoor News (WON) database, which consists of a long-term (1966-2013) time series of trophy-sized fishes caught along the entire California coast and Baja

California, Mexico. Sixteen demersal species, with a combined total of 25,943 records, were the focus of this study for both temporal and spatial analyses. Twelve of the 16 species showed signs of long-term decline in trophy size, but 9 of these 12 showed signs of transition to either stabilization or initial recovery in the most recent years. The more progressive state fisheries management actions adopted since 2001 has likely contributed to this apparent shift in trends, and the new coastal Marine Protected Area (MPA) network are likely to benefit these populations even more in the near future. Trophy size increased with distance from home port, and the distance required to reach maximum trophy sizes increased over time, especially for the rockfishes. Trophy sizes of fishes caught specifically within the Northern Channel Islands region showed long-term

declines through the early 2000s, but also show a period of stabilization or recovery for some species since the 2003 implementation of the Northern Channel Islands network of

MPAs.

Introduction

Marine recreational fisheries in California contribute significantly to the state economy (NMFS 2011), support coastal communities and cultural heritage (Scholz et al.

2004), and account for a significant portion of total landings for a number of fish species

(Hill and Schneider 1999, Schroeder and Love 2002, Dotson and Charter 2003). These fisheries are highly dynamic, with a primary focus on pelagic species during the summer months. However, fishing economies are sustained during the rest of the year by demersal species, which have recently undergone extensive management changes. Prior to 2001, recreational fisheries regulations for demersal fishes primarily consisted of individual species bag limits, size limits for some species, and seasons for only a few species. This was also a period characterized by signs of population decline for many fished species (Dayton et al. 1998, Love et al. 1998, Mason 1998). In 2001, several rockfishes, especially Cowcod (Butler et al. 2003), were declared overfished. This led to an unprecedented shift to more progressive regulatory policies that were fully underway by 2003. This shift began with the implementation of the Cowcod Conservation Areas, and later included broad fishing depth restrictions, seasonal and depth closures, and networks of Marine Protected Areas (MPAs) along the mainland coast as well as the northern and southern Channel Islands. While these new policies were contentious among

marine stakeholders, fisheries adapted quickly after the implementation of these management changes, and positive signs have emerged since. For example, Cowcod were initially projected to be rebuilt by 2068 (Dick et al. 2007), but a more recent report indicates that the population has increased more rapidly than expected, and may be rebuilt as soon as 2020 (Dick & MacCall 2013). In addition, Field et al. (2010) showed that Bocaccio (Sebastes paucispinis) are in a rebuilding state following an earlier conclusion that stocks were severely depleted (MacCall et al. 2002), and Hamel et al.

(2009) found that the spawning biomass of Lingcod (Ophiodon elongatus) has been in recent recovery. Just over a decade after a more conservation-oriented shift in management policy, combined with some years of strong recruitment (Love et al. 2012), some rockfish and other groundfish populations are beginning to show signs of rebuilding

(Miller et al. 2014, Sewell et al. 2013).

It is clear that declining trends among demersal species in California can be reversed if the necessary temporal and spatial data are available to assess stock status and inform new management decisions. The most important information required for a stock assessment includes catch and effort data, abundance data, and data specific to the biology of the species, such as size structure, growth, fecundity, maturity, and movement characteristics (NOAA 2012). Long-term and spatially explicit information yields much more informative assessments, yet species are often considered “data poor,” lacking some or much of the necessary data. For example, long-term datasets for California species often lack size-specific information or spatial characterizations, and are limited only to total landings estimates. This provides an incomplete snapshot of population information,

and ultimately limits our ability to make informative conclusions about the status of the stock.

Spatial information specific to population demographics is not only important for stock assessments, but can also be used to understand the influence of fishing effort on population characteristics. For example, a state-wide MPA network was recently implemented along the entire California coast, including the local islands. The state of

California implemented this network on a regional basis, first in the Northern Channel

Islands (effective April 9, 2003), followed by the central marine region (Sept 21, 2007), north-central marine region (May 1, 2010), southern marine region (Jan 1, 2012), and lastly the northern marine region (Dec 19, 2012). Spatial information specific to coastal fisheries allows measurements of how these MPAs function under varying ages and levels of protection, and studies have recently been implemented to quantify baseline metrics for these MPAs (e.g., Hovel et al. 2015). Hamilton et al. 2010 found that commonly targeted fish species showed increased densities and biomass inside the boundaries of the Northern Channel Islands MPAs after five years of protection, although signs of improved fishing outside the MPA boundaries have still not been measured.

Spatial information can also be used to measure the impacts of fishing because more isolated areas tend to receive less fishing pressure, thus allowing demographic analyses across a gradient of fishing pressure (e.g., DeMartini et al. 2008, Weijerman et al. 2013).

The California Recreational Fisheries Survey (CRFS) was created to fill the critical gaps in data collection, but this program began only recently in 2003, and the data are not statistically comparable to the preceding program, the Marine Recreational Fisheries

Statistics Survey (MRFSS). As a result, continuous long-term time-series metrics other than total landings, such as size specific information, are rare. This is especially important in California fisheries due to their generally size-selective nature.

Impacts of size selective fishing on fish populations are broad, and have been the subject of numerous studies during the last several decades (since Beverton and Holt

1957). Many of these impacts occur after sustained size-selective fishing over long time periods. For example, changes in life history characteristics of California Sheephead

(Semicossyphus pulcher) were found to be largely dependent on levels of site-specific fishing pressure over a 2-3 decade period (Hamilton et al. 2007). However, some impacts can be measured on short-term scales. Demographic effects, such as decreased age and size structure (e.g., Schroeder and Love 2002), are often the first sign of fishing impacts

(Laugen et al. 2014). Such decreases in the abundance of both large individuals and large species (e.g., McClenachan 2009a,b; Powers et al. 2013) can occur even before declines in total catch frequencies (Froese 2004). Size structure information may thus allow us to recognize early signs of the impacts of fishing (Yunne-Jai Shin 2005), because a population lacking large individuals may thus be the product of recent over-exploitation

(Gislason and Rice 1998, Pauly et al. 2002, DeMartini et al. 2008, Sandin et al. 2008).

However, this is only detectable if data specific to the largest size classes are available.

These trophy fishes represent an especially important component of the overall spawning stock biomass because they have greater fecundity with higher quality eggs (Berkeley et al. 2004, Calduch-Verdiell 2014), and they maintain genetics for large individual size within the population (Fenberg and Roy 2008). The California Recreational Fisheries

Survey (CRFS) database does contain size structure data, but only a small fraction of the data represents the largest size classes because these individuals are rarely caught.

We generated a new trophy size specific database from a California fishing newspaper, Western Outdoor News (WON). On a weekly basis, this publication lists all the largest fish caught at each sportfishing landing along the California coast from 1966 to present. We entered all of these data into a central database, and the purpose of this study is to measure temporal and spatial changes of trophy size dynamics for demersal species in California, with the null hypotheses that 1) no changes in trophy size of demersal species have occurred over time and 2) trophy size of demersal fishes does not change relative to distance from fishing port or proximity to established MPA networks.

Methods

Demersal species were the focal group for this study, with records from 1966-

2013 ranging from northern Baja California to the California-Oregon border. Data from long-range fishing trips were not included in the analysis because they are not considered representative of California coastal populations. Almost all records contained at least a general catch location, and most contained specific catch locations. Specific catch locations were those that could be assigned latitude-longitude coordinates within a 3km confidence radius. For example, we could assign a single set of coordinates for a White

Sturgeon with a “Roe Island” catch location because this is a specific place in the San

Joaquin-Sacramento Delta where White Sturgeon are sometimes caught. We can also classify this into a generalized catch location of Suisun Bay within Solano County.

Conversely, a catch location reported as “Catalina Island” is too broad for specific coordinates due to the size of the island, but the record could still be used in coarse scale spatial analysis. Distance from home port to specific catch locations could then be calculated for those records that were specific enough to be assigned latitude-longitude coordinates.

Temporal changes in trophy size were measured using the same methods described in chapter 1 for pelagic and coastal pelagic species. A Bayesian approach was used with the package ‘R2jags’ (Su & Yajima 2012), which uses JAGS (Plummer 2003) within the statistical program R. A Bayesian hierarchical model was used to measure temporal trends, including process and observation errors. Annual boxplots were used to show the distribution of the raw data within the time series, and the state-space model output was overlaid onto the boxplots to describe the trend over time.

Spatial patterns to trophy size were first analyzed with respect to the degree of isolation of catch locations relative to vessel home port. These data were also arbitrarily split into two time periods within the database (1966-1989 and 1990-2013) to test for an influence of time on isolation effects. A two-way ANOVA and Tukey’s post hoc tests for subsequent pairwise comparisons were used to test for effects of both time and distance on trophy size. The relationship between fish size and distance from port was plotted using boxplots at three distance intervals (0-25km, 26-50km, and 51-100km) as well as the two time intervals. This range of distances generally encompasses the overall range of

CPFVs and private vessels that operate locally for demersal species, which combined represent the majority of California recreational landings for demersal species. For the

purposes of this study, this includes ½-day, ¾-day, overnight, and 1 ½-day CPFV trips as well as most private vessel trips. Some of the species had too few catch location coordinates assigned for robust analysis, and were thus not included in the results.

Spatial patterns in trophy size were also measured before and after the implementation of the first network of marine protected areas (MPAs) in California. This network was implemented in 2003 within the Northern Channel Islands National Marine

Sanctuary off the coast of Santa Barbara and Ventura counties. One of the original goals of this MPA network was to improve fisheries for the entire Northern Channel Islands region as a whole. WON data from this region were thus treated as such, and pooled for analyses. Trophy size data for demersal species within this region were then split into four time periods. Three of these are approximately equal periods (~12 years each) before the 2003 MPA network implementation, and the last period represents the period after implementation (11 years). A one-way ANOVA and Tukey’s post hoc for pairwise comparisons were used to test for differences in size during the four periods.

Unfortunately, a true BACI design is likely not realistic even though data are available for adjacent areas, such as the Santa Barbara mainland coast, the Ventura mainland coast, or southern Channel Islands. While these regions are geographically close to the study region, they are often characterized by very different and confounding oceanographic features, meaning that defining a true control region is likely unrealistic. Species with too few records within the Northern Channel Islands region were not included in the results.

A new network of MPAs has also been implemented along the mainland California coast,

but these are likely too young to show reserve effects on regional populations, so analyses with respect to these MPA networks will be the subject of a future study.

Results

Temporal Trends

The WON database consists of 48,750 records total, but this study focuses on the top 16 demersal species (14 teleosts and 2 elasmobranchs), which account for 25,943 records, or 53.2% of the database (excluding long-range catch records from Mexico that were not included in the study). The trophy record ranges temporally from 1966-2013 and spatially from northern Baja California to the California-Oregon border. The

Southern California Bight represents 60.9% of the data while the majority of the remaining data are from the San Francisco Bay area. Other demersal species are represented in the database, but their frequencies are too low for robust time-series or spatial analysis.

Temporal trends show declines in trophy weight for most of the species during the study period, although most of these long-term declines show a period of stabilization in the most recent years (since the early 2000s). This is especially true for the groundfishes in this study, including most of the rockfishes in the genus Sebastes. Cowcod (S. levis:

Fig. 1a) show a consistent decline in size until the fishery was closed in 2001, after which time almost no records of catch were reported. Vermilion Rockfish (S. miniatus: Fig. 1b) show a decline during the 1990s, but the decline stabilized after approximately 2002.

Bocaccio (S. paucispinus: Fig. 1c) are slightly more variable, but show a gradual decline

during the study period that appears to stabilize beginning around 2004. Copper Rockfish

(S. caurinus: Fig. 1d) were reported at a much lower frequency than other species but this frequency has increased over time, showing a decline in trophy size since the mid-1990s.

Yelloweye Rockfish (S. ruberrimus: Fig. 1e) were only reported from the late 1970s until the fishery was closed in 2001, but the limited data show a slight decrease in trophy size.

Figure 1f shows the Sebastes complex combined, but Cowcod are excluded from this do to the significant influence of such a large species on the overall outcome. Even with

Cowcod excluded, the Sebastes complex shows a significant decline by approximately

50% until the early 2000s. During the last decade, the Sebastes complex trophy size has stabilized.

Temporal trends in other demersal nearshore species are shown in figure 2a-d for

Cabezon (Scorpaenichthys marmoratus), Lingcod (Ophiodon elongatus), White Sturgeon

(Ascipenser transmontanus), and California Halibut (Paralichthys californicus), respectively. Records for Cabezon (Fig. 2a) show a slight decline in trophy size over time, followed by a significant decrease in 2012 and 2013. Lingcod (Fig. 2b) show a consistent decline in size, but the last two years show a significant increase for the first time in over a decade. White Sturgeon (Fig. 2c), which were recorded almost entirely from the San Francisco Bay area, show a significant decrease in size around 1990-1991, but this is an artifact of the implementation of “slot” size limits for this species. After the slot limit adoption, this species has shown a gradual decline in mean trophy size from approximately 28kg to 21kg, even though larger outliers begin to appear more in recent years. California Halibut (Fig. 2d) showed a significant decline until 1980, after which

time the trophy size stabilized until 2012-2013 when size increased to levels representative of the late 1960s and early 1970s.

Species more associated with kelp forest communities (Fig. 3a-e) also showed signs of decline during the study period. California Sheephead (Semicossyphus pulcher:

Fig. 3a) declined in the 1970s, again in the 1990s, and have stabilized during approximately the last decade. Ocean Whitefish (Caulolatilus princeps: Fig. 3b) show a relatively consistent decline in trophy size, but sample sizes are quite low for the time series. The recreational fishery for Giant Seabass (Stereolepis gigas: Fig. 3c) was closed in 1981, so data after that time are limited. While the early portion of the database shows no overall trend in size of Giant Seabass from 1966-1983, we do see a loss of the largest outliers by the mid-1970s. The few data points in more recent years show that relatively large individuals are beginning to be caught (and released) again, but data are extremely limited. Kelp Bass (Paralabrax clathratus: Fig. 3d) show a significant decline in trophy size throughout the study period, and this decline gains steepness during the most recent

5-6 years in the database. Barred Sand Bass (P. nebulifer: Fig. 3e) show a stable trend in trophy size until the late 1990s, and size has decreased significantly since that time.

Demersal elasmobranchs in the database primarily consisted of Broadnose

Sevengill (Notorynchus cepedianus: Fig. 4a) and Leopard Sharks (Triakis semifasciatus:

Fig. 4b), both of which are primarily recorded from the San Francisco Bay area. Both species are absent from the database prior to 1984, and Leopard Sharks do not show a clear trend in size since that time. However, Sevengill Sharks show a significant increase in trophy size beginning in the mid-1990s, and continuing through 2013.

Spatial patterns

The relationship between trophy size and distance from port was measured for 10 of the 16 focal species in this study. Omitted species were due to an insufficient number of records containing latitude-longitude coordinates for catch locations, which precluded measurements of distance from home port to the catch location. Rockfishes showed significant patterns in trophy size with respect to catch distance, and there was also a significant temporal component to this pattern (F = 45.50, df = 2, p < 0.001, Fig. 5a-e). In all three distance categories from home port, the Sebastes complex exhibited trophy sizes that were significantly less in the 1990-2013 period than in the 1966-1989 period (p <

0.01, Fig. 5e). In addition, during the 1966-1989 period, the Sebastes complex shows lower trophy sizes within 0-25km from home port, and higher catch sizes at distances greater than 25km from port (p < 0.01, Fig. 5e). During the 1990-2013 period, this pattern becomes stronger, with trophy sizes remaining lower to an increased distance of

50km from home port, and increasing only at distances greater than 50km from home port. However, these larger sizes from greater than 50km away are not significantly different from the trophy sizes that were recorded from less than 25km away during the

1966-1989 period (p = 0.065) . In other words, not only did anglers travel farther distances to catch larger fish in both periods, the distance required increased over time. In addition, the trophy sizes at the greatest distances ultimately decreased to a point that was comparable to trophy sizes at the closest distances from port during the earliest years.

Anglers now have to travel greater than 50km to catch the same sized rockfishes that they used to catch next to home port. A similar pattern, although not as strong, can also be

seen for California Sheephead, Lingcod, Cabezon, and California Halibut (Fig. 6a-d), and

Tukey’s post hoc results are reported in each panel. Kelp Bass show significantly decreased trophy sizes between the two time periods at all three distances from port (F =

19.39, df = 5, p < 0.001), but there is no relationship between trophy size and catch distance itself. Barred Sand Bass show no clear pattern in size with distance from home port, although sample sizes in the farthest distance bin are very low (less than 10 individuals) because most are caught at spawning aggregations that are relatively close to each home port.

Changes in trophy size over time in the Northern Channel Islands region may be associated with the implementation of the 2003 network of MPAs. Trophy size data from only the Northern Channel Islands were analyzed for 8 species during the study period

(the remaining focal species in this study were not recorded in this region with a large enough sample size for spatial analysis). Data for each species was divided temporally into four approximately equal time steps within the database, the last of which (2003-

2013) represents the period after the MPA network was implemented. California Halibut

(Fig. 7a) showed a significant decrease in size during the 37-year period before the MPA network, and a significant increase during the last ten years after the network had been implemented. For comparison, California Halibut along the adjacent Santa Barbara mainland coast do not show the same increase in size during the period after MPA implementation. Lingcod (Fig. 7b) and Kelp Bass (Fig. 7c) both showed a significant decrease before the MPAs, and this decrease has stabilized for both species since the

MPAs have been in place. Kelp Bass along the Santa Barbara mainland coast show

continuous declines in trophy size during throughout the full time series. California

Sheephead (Fig. 7d) decreased in trophy size before the MPAs were adopted, and this decrease has since continued even after the MPAs. Ocean Whitefish (Fig. 7e) may have decreased before the MPAs, reaching a point of stabilization afterwards, although the earliest time steps lack a sufficient sample size to reach a definitive conclusion. In the genus Sebastes, Vermilion Rockfish (Fig. 8a) show a consistent decrease in size both before and after the MPA network, although the earliest time step does not contain any records of this species for the Northern Channel Islands region. Bocaccio (Fig. 8b) show some variation over time, but no change directly before and after the MPAs. However, this species has a very low sample size for the region overall. Cowcod (Fig. 8c) show a consistent and significant decrease over time before the MPA network, but no data are available following the MPAs due to the 2001 closure on this fishery.

Discussion

Temporal analyses of trophy sizes among demersal species show a range of trends, but the majority of cases illustrate an overall pattern of decline, followed by a more recent period of stabilization in some cases. Twelve of the sixteen demersal species included in this study exhibited long-term declines in trophy size in California from

1966-2013. In contrast to the pelagic and coastal pelagic species (chapter 1), demersal species are generally longer-lived, slower growing, less fecund, mature later, exhibit more localized movement patterns, and are more regularly accessible to a broader segment of the fishing community. Demersal species thus have a greater likelihood of

being impacted by fishing, and this may explain the broad declines exhibited in the WON database. Some of the focal species in this study are targeted in both commercial and recreational fisheries (e.g., Vermilion Rockfish and California Sheephead), and others are targeted only recreationally (e.g., Kelp Bass and Barred Sand Bass), but species in both groups exhibited long-term declines in trophy size. While the impacts of fishing are often associated with commercial fisheries, it is clear that California fish populations can be impacted by recreational fisheries as well (e.g., Schroeder & Love 2002, Jarvis et al.

2014).

The rockfishes show the strongest signs of decline among the species in this study. These declines come not only in temporal form during the 1966-2013 period, but spatial declines as well. During the early half of the database (1966-1989), pooled analyses for the genus Sebastes show rockfishes weighing approximately 8kg could be caught close to home port. During the more recent half of the database (1990-2013), similar sized fish could only be found greater than 50km from home port, primarily at offshore banks, the more isolated islands, and Mexican waters. Anglers now have to travel over twice the distance to catch the same sized rockfish from earlier decades, and the largest outliers are seldom caught at all, at any distance from port (this is true even with Cowcod excluded). A similar pattern can be seen for other demersal species in the

WON database, such as California Sheephead and Lingcod, but it is particularly evident with fish in the genus Sebastes. Even within the Northern Channel Islands region, the

WON trophy data show that Vermilion Rockfish continue to decline in size despite the

2003 MPA network implementation. Evidence of serial depletion is rare in California

recreational fisheries, although Karpov et al. 2000 found sequential spatial depletion of red, pink, and black abalone (Haliotis sp) populations in coastal California waters.

Despite the overall pattern of decline, there are two points that may suggest a more positive future for California demersal species. First, of the twelve species that showed long-term declines during the study period, nine of these have more recently transitioned into a state of either stabilization or initial recovery in trophy size. Declining trends for these species may thus be in various states of population rebuilding due to the more progressive fisheries management policies adopted since 2001. These policies included fishing depth restrictions, gear restrictions (reduced number of hooks allowed), seasonal restrictions, and implementation of the Cowcod Conservation Areas. Signs of stocks rebuilding have recently been measured for a number of different groundfish species (Miller et al. 2011, Sewell et al. 2013), so it is plausible that trophy sizes of some of these species may be recovering as well. Second, an extensive state-wide network of

MPAs was recently implemented in a spatially phased process from 2003-2012. This network consists of 119 MPAs, 15 special closures, and 5 marine managed areas covering approximately 2200 km2, or ~16% of California state waters, across all of the most abundant benthic habitat types (Pope 2014). Because of the life history characteristics stated above, demersal species are the most likely group to benefit from these new MPAs. Some of these species are already exhibiting initial signs of benefit within the Northern Channel Islands MPA network that was recently implemented

(Hamilton et al. 2010, Karpov et al. 2012), and WON trophy data show some signs that are consistent with this pattern. For example, California Halibut have increased in size

since the network was implemented, and both Lingcod and Kelp Bass have stabilized following declines that were occurring prior to the MPA network.

The four species that do not follow the general pattern of decline are Yelloweye

Rockfish (Sebastes ruberrimus), Giant Seabass (Stereolepis gigas), Sevengill Shark

(Notorynchus cepedianus), and Leopard Shark (Triakis semifasciatus). However, the time series available for Yelloweye Rockfish and Giant Seabass are very short, due to population declines and subsequent closures of these fisheries in 2001 and 1983, respectively. In addition, Giant Seabass were already heavily impacted long before the beginning of the time series in this study (Ragen 1990, Dayton and MacCall 1992). Since the closure for Giant Seabass, a few larger individuals were reported in catch-and-release records. These records suggest a recovery in trophy size may be in progress, and recent signs of population recovery for this species have been documented recently (Pondella and Allen 2008). Broadnose Sevengill and Leopard Sharks may simply be younger fisheries, or they were not reported prior to the mid-1980s. Sevengills show an increase in size during approximately the last fifteen years, likely because anglers have learned more recently how to target and catch this species.

Overall patterns show declines in trophy size for most of the demersal species, but more progressive management actions may be helping to reverse this trend.

Acknowledgements

I would like to acknowledge my co-author, Brice Semmens, for assisting with the analyses of the WON time series data.

Figure 2.2. Time series plots of trophy fish weight showing model output (white line) and error (grey region) overlaid onto boxplots of the raw data for Cabezon (A, n = 192), Lingcod (B, n = 7884), White Sturgeon (C, n = 2877), and California Halibut (D, n = 7729).

Figure 2.4. Time series plots of trophy fish weight showing model output (white line) and error (grey region) overlaid onto boxplots of the raw data for Broadnose Sevengill Shark (A, n = 148) and Leopard Shark (B, n = 105).

Figure 2.5. Boxplots showing median weights of Cowcod (A, n = 1903), Vermilion Rockfish (B, n = 708), Bocaccio (C, n = 256), Yelloweye Rockfish (D, n = 153), and all Sebastes (E, n = 3104) at three distance bins from home port. Gray bars represent the temporal half of the database (1966 – 1989), and white bars represent the second half (1990 – 2013). Numbers above each boxplot represent groupings from Tukey’s pairwise comparisons.

Figure 2.6. Boxplots showing median weights of California Sheephead (A, n = 573), Lingcod (B, n = 5652), Cabezon (C, n = 141), California Halibut (D, n = 4831), Kelp Bass (E, n = 541), and Barred Sand Bass (F, n = 353) at three distance bins from home port. Gray bars represent the temporal half of the database (1966 – 1989), and white bars represent the second half (1990 – 2013). Numbers above each boxplot represent groupings from Tukey’s pairwise comparisons.

Figure 2.7. WON trophy weights from the Northern Channel Islands during four periods, the last of which is the most recent decade following the 2003 implementation of the network of marine protected areas in the region. The species included are California Halibut (A, n = 1074), Lingcod (B, n = 1125), Kelp Bass (C, n = 56), California Sheephead (D, n = 211), and Ocean Whitefish (E, n = 28). Numbers above each boxplot represent groupings from Tukey’s pairwise comparisons.

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CHAPTER 3: Analyses of catch-per-unit-effort (CPUE), length frequency, and post- release mortality of three species of Paralabrax relative to coastal marine protected areas

(MPAs) off San Diego, California

Abstract

The genus Paralabrax includes three of the most important species in California recreational fisheries, yet signs of population declines have become evident in some cases. Both Kelp Bass (P. clathratus) and Barred Sand Bass (P. nebulifer) show long- term negative impacts from fishing and oceanography, while Spotted Sand Bass (P. maculatofasciatus) are more data limited. However, fundamental population dynamics are still not fully understood for all three species. We implemented a collaborative study in partnership with the recreational fishing community in San Diego, California in order to estimate population parameters that are essential to conduct future stock assessments on these three species. We achieved high catch rates for both Kelp Bass and Spotted Sand

Bass, both during and outside their respective spawning seasons. Low catch rates of

Barred Sand Bass were due to recent changes in spawning aggregation behavior in their traditional spawning grounds. Catch-per-unit-effort (CPUE) did not differ between sites for any of the three species, likely because numerous factors contributed to high variability in CPUE estimates. Kelp Bass showed significantly larger size classes in

Imperial Beach, as well as both State Marine Reserves (SMRs) in La Jolla. Spotted Sand

Bass showed significantly larger size classes in Mission Bay than San Diego Bay.

Increased sizes following the 2013 regulatory increases in minimum size limits were

documented in 2014 for both Kelp Bass and Spotted Sand Bass. Kelp Bass exhibited the greatest population response in areas open to fishing than within MPA boundaries.

Barred Sand Bass, however, showed decreased sizes between 2013 and 2014, possibly due to recent changes in spawning behaviors. Overall, the two MPAs in this study exhibited positive effects in terms of size structure rather than CPUE. While Matlahuayl

SMR has been in place for several decades, it is relatively small, which may limit its overall effectiveness. The success of this project was largely due to the extensive collaborative relationships we developed between the fishing, scientific, and fisheries management communities, and we encourage this approach for future studies.

Introduction

Marine recreational fishing plays a vital role in the ecological, social, and economic constitution of California. In 2006, California recreational anglers generated the highest sales in the nation (by state) at an estimated $3 billion on retail goods and services (Gentner & Steinback 2008). In 2013, California recreational anglers contributed

$2.8 billion to the state economy, and spent over 5.3 million angler-days fishing in marine waters (CADFW 2014 Annual Forum Report). Given both the value and impact of recreational fisheries in the state, there is a critical need for detailed and quantitative information specific to exploited fish populations. However, even species with extensive fisheries histories that are relatively well studied have nonetheless exhibited signs of long-term decline, and still lack up-to-date stock status information. One such group of species is in the genus Paralabrax, which in southern California includes the Kelp Bass

(P.clathratus), Barred Sand Bass (P. nebulifer), and Spotted Sand Bass (P. maculatofasciatus). These species have fisheries data that date back to the 1930s, with population analyses dating back several decades (e.g., Clark 1933, Collyer 1949, Collyer and Young 1953, Young 1963), yet signs of population declines have been shown for two of the three species (Sweetnam 2009, Jarvis et al. 2010, Erisman et al. 2011, Jarvis et al.

2014). These two species, Kelp Bass and Barred Sand Bass, consistently rank among the top recreational fisheries by landings. Pooled CPFV logbook data for all targeted fishes in

California from 1936-2008 indicate that Kelp Bass ranked 1st in number of fish kept for a single species (excluding “unidentified rockfishes”), with approximately 20.6 million fish landed during the 67-year period. Barred Sand Bass ranked 4th overall (excluding Pacific mackerel and “unidentified rockfishes”) with 12.7 million landed. However, both species were often pooled into a “rock bass” category (ranked 5th overall) prior to the mid-1970s, which accounts for an additional 11.9 million fish (Jarvis et al. 2014). Thus, Kelp Bass and Barred Sand Bass likely represent the top two species (excluding Pacific mackerel and “unidentified rockfishes”) landed on CPFVs throughout the recorded history of recreational fishing in California. However, signs of their long-term declines have become evident (Erisman et al. 2011), which has been due to a combination of fishing and oceanographic factors (Jarvis et al. 2014).

Population declines of Kelp Bass and Barred Sand Bass were documented in the

1940s due to commercial overfishing, which ultimately led to a ban on commercial fishing for these species in 1953 (Young 1963). While recreational fishing is considered to have lesser impacts than fishing on commercial scales, the popularity of these bass

species among the recreational fleets has resulted in over 45 million bass landed aboard

CPFVs alone in California waters since 1936 (CDFW logbook data). Overfishing is thus a concern for these species, not only because of the sheer numbers landed, but also due to the social and economic importance of the species. In addition, the regulatory history for these species has been dynamic throughout the history of the fishery (Jarvis et al. 2014), and the California Department of Fish and Wildlife recently implemented new individual bag limits and size limits for the basses, as well as new MPAs for all species. An understanding of the response of the bass fisheries to these harvest control rules and spatial regulations can help to establish management actions that ultimately facilitate a productive and sustainable fishery.

Susceptibility to overfishing is largely influenced by species behavior and biology. Barred Sand Bass are of particular concern because this species forms large summer spawning aggregations that are highly predictable in time and space (Hovey et al. 2002, Jarvis et al. 2010), and thus easy to target. Erisman & Allen (2006) suggest that

Kelp Bass also form spawning aggregations, but they also state that these aggregations are spatially and temporally unpredictable, which implies that the susceptibility of this species to overfishing is lower than that of Barred Sand Bass. Jarvis et al. (2014) recently concluded that a combination of fishing impacts, fisheries regulations, and environmental processes have collectively led to long-term declines in both Kelp Bass and Barred Sand

Bass landings. However, current estimates of population statistics for these species are needed, especially given that some data sources are decades old (e.g., Young 1963).

Clearly, an intensive study on temporal and spatial dynamics in catch-per-unit-effort

(CPUE) and length frequency analyses would not only lend insight into the status of the populations, but would also allow measurements of the effectiveness of recent regulatory changes. These changes include coastal MPAs that were recently implemented, as well as increased minimum size limits and decreased individual bag limits that were adopted in

2013 for all of the bass species. In addition, there has been limited research focusing on the third species of Paralabrax, Spotted Sand Bass (P. maculatofasciatus) in general.

This species inhabits bays and estuaries almost exclusively, and therefore is primarily targeted by private boaters and shore anglers because CPFVs mostly operate along the open coast. Spotted Sand Bass are also targeted primarily for catch-and-release among recreational anglers (Hovey and Allen 2000). This stark difference in recreational use patterns between the bass congeners provides an opportunity to compare differences in the influence of primarily consumptive versus primarily non-consumptive recreational fishing pressure on population dynamics.

Given the popularity of catch-and-release fishing in southern California, especially for the three saltwater basses, there is a need for post-release mortality estimates of each of the three species. Data from the California Recreational Fisheries

Survey (CRFS) indicate total catch-and-release rates for Kelp Bass, Barred Sand Bass, and Spotted Sand Bass are 73%, 55%, and 95%, respectively (CalCOFI 2014). Such high catch-and-release rates are likely due to a combination of minimum size limits and the perception among recreational anglers that catch-and-release promotes conservation due to high post-release survival. If post-release survival is much lower than perceived, then these catch-and-release rates would equate to significantly greater overall take from

recreational fisheries, which should be factored into population analyses. Lowe et al.

2003, McKinzie et al. 2014, and Freedman et al. 2015 each measured high post release survival of Kelp Bass, Barred Sand Bass, and Spotted Sand Bass, respectively, but generally only fish that appeared healthy were actually tagged in these studies because acoustic transmitters are costly.

The goal of our project was to work cooperatively with the recreational fishing community, the California Dept. of Fish and Wildlife, and non-profit organizations to conduct a quantitative analysis of CPUE, length frequency, and post-release mortality for all three bass species in the San Diego county coastal area, both inside and outside local marine protected areas (MPAs). This will inform future stock assessments of these species. There are primarily two MPA types located in our study area. These include

State Marine Reserves (SMRs), which prohibit all extractive activities, and State Marine

Conservation Areas (SMCAs), which are primarily no-take, but include some allowances for recreational take only. This information will shed light on functionality of both old and new MPAs, responses of each species to recently increased minimum size limits, and the often overlooked impacts of catch-and-release fishing.

Methods

Study Area

Sampling trips were conducted from Long Beach, California south to the US-

Mexico border, although the primary research sites consisted of north San Diego County,

La Jolla, Mission Bay, Point Loma, San Diego Bay, and Imperial Beach off the San

Diego County coast (Fig. 1). The north San Diego County area was divided into two sites, which consisted of the Swami’s State Marine Conservation Area (SMCA) established in 2012, and the adjacent Encinitas area that has always been open to fishing.

The La Jolla area was divided into three sites, which consisted of 1) the Matlahuayl State

Marine Reserve (SMR), which was expanded and renamed in 2012 from the former La

Jolla Ecological Reserve implemented in 1971, 2) La Jolla (has always been open to fishing), and 3) the South La Jolla SMR, which was recently established in 2012. Sites were chosen based on suitable habitat and known popular fishing locations for each of the three species. Adult Kelp Bass generally are targeted along open coast rocky reef and kelp bed habitat, Barred Sand Bass are targeted in bay and coastal sand flats near rocky reefs, and Spotted Sand Bass are fished in primarily soft bottom bay habitats. As such, primary Kelp Bass sites included La Jolla and Point Loma kelp beds, Barred Sand Bass sites included Imperial Beach and north San Diego County, and Spotted Sand Bass sites were located in Mission Bay and San Diego Bay.

Field Methods

Two approaches were used to implement tagging efforts for the three bass species. The first consisted of ¾-day charters aboard Commercial Passenger Fishing

Vessels (CPFVs) in collaboration with the San Diego Oceans Foundation, a local 501(c)3 non-profit organization. These CPFV charters focused on sampling coastal sites only (not sites inside bays), and the primary target species for these trips were both Kelp Bass and

Barred Sand Bass. The second sampling approach was with private vessels in the two bay

sites, Mission Bay and San Diego Bay. These trips focused primarily on tagging the third species of Paralabrax, Spotted Sand Bass.

The CPFV tagging charters were separated into sampling periods based on season during the two-year study (Table 1). Seasons consisted of fall 2012 (Sep-Nov), summer

2013 (Jun-Aug), fall 2013 (Sep-Nov), and summer 2014 (Jun-Aug). Each individual charter focused specifically on one site and one target species. This yielded site-specific

CPUE for both the target species as well as all bycatch that was associated with each target species. Volunteer recreational anglers were invited aboard each of these charters to assist with the fishing, and SIO researchers were aboard for catch data collection. All fish caught were measured, recorded, and released immediately after capture. The information collected for each individual fish caught consisted of the species, standard length (mm), latitude-longitude, time, depth, and physical condition of the fish.

Catch-and-release mortality was measured using two methods. First, mortality events were observed visually from the boat, yielding an initial mortality estimate.

However, initial post-release mortality rates are likely to be underestimates of overall catch-and-release mortality given that they represent only immediate mortality events that were observable from the boat. Thus, the second method consisted of fish being held in moored net pens located in San Diego Bay for a 10-day assessment period, yielding a short-term post-release mortality estimate. Fish from a local annual fishing tournament in both 2013 and 2014 were released into these net pens, and then all fish were released after the temporary holding period. All mortality events during the holding period were recorded.

Data Analysis

Measurements for both CPUE and length frequency were grouped by species and site, while short-term post release mortality was grouped by species and pooled across all sites. Species-specific CPUE estimates were only generated for those trips where the species being estimated was the primary target species (e.g. we only estimated trip- specific Kelp Bass CPUE when Kelp Bass was the primary target species for the trip).

Catch per unit effort was used as a measure of relative abundance, and was defined in terms of the number of fish caught per angler-hour. This metric was compared for each species between sites using a one-way ANOVA and post hoc Tukey’s test for pairwise comparisons. Length-frequency data were compared between sites also using a one-way

ANOVA and post hoc Tukey’s test for pairwise comparisons. Individual growth rates based on tag-recapture data are the subject of a separate study comparing current growth estimates with past studies (Allen et al. 1995, Love et al. 1996). However, in response to signs of long-term population decline among both Kelp Bass and Barred Sand Bass, the

California Department of Fish and Wildlife implemented regulatory changes for all three species of Paralabrax effective March 1, 2013 (Jarvis et al. 2014). These regulations included an increased minimum size limit from 305 to 356 mm (12 to 14 inches) total length, as well as a decreased individual daily bag limit from 10 to 5 fish. This allowed us to measure changes in the size frequency of each species following the release of fishing pressure on 12-14-inch fish. We tested for changes in the overall size distribution of each species of Paralabrax between the 2012, 2013, and 2014 sampling years at the primary

tagging sites with a two-way ANOVA and post hoc Tukey’s test for pairwise comparisons.

Results

Realized tagging effort

Our tagging efforts consisted of a total of 51 individual ¾-day tagging charters aboard local CPFVs (Table 1), and 151 private vessel trips conducted from Sep 30, 2012

- Sep 17, 2014. Tags were deployed at 11 sites between Long Beach, California and the

U.S.-Mexico border, including two State Marine Reserve sites (Matlahuayl SMR and

South La Jolla SMR) and one State Marine Conservation Area (Swami’s SMCA). A total of 19,327 fish from 71 species were caught-and-released during the trips, although

Pacific Jack Mackerel (Trachurus symmetricus), Chub Mackerel (Scomber japonicus), and Jacksmelt (Atherinopsis californiensis) were not recorded because catch rates for these species were sometimes high enough to interfere with data collection for the primary target species. A total of 12,581 Kelp Bass, 1079 Barred Sand Bass, and 2353

Spotted Sand Bass were caught and released during this period (Table 2). The basses represent 84.2% of the total records in the catch dataset, and the remaining 15.8% bycatch consisted primarily of Pacific Barracuda (2.45%), Brown Rockfish (1.76%),

Kelp Rockfish (1.51%), Bonefish (1.36%), Yellowfin Croaker (1.06%), and 60 additional species with less frequent catches (Table 3).

Catch Per Unit Effort Across Species and Sites

Neither Kelp Bass (Fig. 2, F = 0.93, df = 6, p= 0.44) nor Barred Sand Bass (Fig.

3, F = 0.08, df = 1, p= 0.78) showed significant differences in CPUE between survey sites. There were also no significant differences in Spotted Sand Bass CPUE between sites (Fig. 4, F = 0.069, df = 1, p= 0.794). However, there were significant differences between sampling seasons in Kelp Bass CPUE (Fig. 5, F= 16.14, df = 3, p = 0.0002), with the Tukey test showing higher values during the summer seasons (p < 0.01), and no differences in the same seasons between years (p = 0.85). Barred Sand Bass were primarily only caught during summer months, which precluded seasonal analyses.

California Recreational Fisheries Survey (CRFS) estimates of recreational landings from

Los Angeles and San Diego CPFVs show very few fish caught at all in 2013, and only slightly more in 2014 (Fig. 6); in nearly all years since 2004, CPUE was highest during the spawning season for this species (summer months). Spotted Sand Bass showed significant differences in CPUE by season (Fig. 7, F= 6.32, df = 3, p = 0.001). The

Tukey’s post hoc test showed higher CPUE in winter months (p = 0.01), and no differences between the other seasons (p = 0.64), although only summer and fall had very low sample sizes.

Length frequency Across Species and Sites

All three basses showed significant spatial differences in length-frequency distributions. Kelp Bass showed significant differences between sites (Fig. 8, F = 60.42, df = 9, p < 0.001), with the post hoc test showing the largest mean bass size in Imperial

Beach (283mm SL ± 2mm 95% CI, p < 0.001), followed by Matlahuayl SMR (267mm

SL ± 2mm, p < 0.001), South La Jolla SMR (260mm ± 2mm, p < 0.001), La Jolla

(254mm ± 1mm, p < 0.001), and lastly Point Loma (251mm SL ± 2mm, p = 0.049). Kelp

Bass did not show significant differences inside and outside of the Swami’s SMCA (Fig.

9), but these data were only generated from a single tagging trip at each site inside and outside the SMCA. Barred Sand Bass were larger at Imperial Beach than all other sites

(Fig. 10, F = 10.24, df = 3, p < 0.001), but sampling was limited at sites other than

Imperial Beach due to low catch rates. Spotted Sand Bass showed significantly larger sizes in Mission Bay than San Diego Bay (Fig. 11, F = 171.07, df = 1, p < 0.001).

Inside/outside reserve comparisons do not apply to Spotted Sand Bass in this study because they generally only live inside bays and estuaries where there have been no

MPAs implemented (there are several de-facto reserves in these areas, such as naval zones, but these sites were not sampled). We also compared the size distributions of the top five demersal bycatch species in the La Jolla sites (La Jolla Cove, La Jolla, and South

La Jolla; Fig. 12). These consisted of Brown Rockfish (Sebastes auriculatus), California

Sheephead (Semicossyphus pulcher), Kelp Rockfish (S. atrovirens), Gopher Rockfish (S. carnatus), and Treefish (S. serriceps). California Sheephead were the only species that showed significant differences between sites (F = 16.31, df = 2, p < 0.001), with the

Tukey’s post hoc test showing larger sizes inside the two SMRs than in the adjacent outside area of La Jolla (p < 0.001), but no differences between the two SMRs (p = 0.63).

Kelp Bass showed significant effects of both site (F = 74.21, df = 4, p < 0.001) and year (F = 12.36, df = 2, p < 0.001) on mean size. Significant increases in size were found from 2013 to 2014 (p < 0.001), with the greatest increases occurring in La Jolla

and Point Loma, the two areas open to fishing. No significant changes in size were measured from 2012 to 2013 (p = 0.022). Barred Sand Bass showed a significant decline in size from 2013 to 2014 (F = 10.47, df = 1, p = 0.001), but this is largely driven by fish from Imperial Beach. Barred Sand Bass from Imperial Beach were significantly larger sized than in San Diego Bay (F = 61.21, df = 1, p < 0.001). Spotted Sand Bass showed significant effects of both site (F = 188.68, df = 1, p < 0.001) and year (F = 11.79, df = 2, p < 0.001), with increases in size from 2012-2014 in both San Diego Bay and Mission

Bay. Individual growth rates are the focus of a separate ongoing study that compares historical analyses to current estimates of Paralabrax demographics.

Post-release mortality

Initial post-release mortality for all species combined was 1.8%, sources of which primarily included fishing-related trauma (64.8%), predation from California sea lions

(25.4%), predation from seabirds (7.8%), predation from harbor seals (2.0%), and predation from other fish (0.6%). California sea lions consumed most of the common species caught, but showed clear preferential behavior towards Pacific Barracuda, Pacific

Bonito, chub mackerel, and White Seabass. Initial post-release mortality for the basses alone were 1.87%, 0.92%, and 0.63% for Kelp Bass, Barred Sand Bass, and Spotted Sand

Bass, respectively (Table 4). Fish held in net pens over a ten day period exhibited a 3.1% mortality rate during this trial period prior to release. Note however, that these fish were

Floy tagged prior to placement inside the holding pens for a separate mark-recapture

study. Thus, this mortality rate likely represents an over-estimate of the extended-term post release mortality.

Discussion

As expected, Kelp Bass CPUE was highest during the summer spawning months

(Jun-Aug), although even the relatively lower catch rates incurred during the fall sampling season (Sep-Nov) translated to several hundred fish caught per trip in many cases. However, there were no site-specific differences in Kelp Bass CPUE, which was largely driven by a few days of exceptionally high catch rates outside MPA boundaries in

La Jolla. In general, CPUE is impacted by numerous factors, such as season, small-scale oceanographic conditions, and angler skill level, that can result in high variability in estimates. In our case, catch rates were potentially influenced by angler skill level, bait availability (e.g., anchovy, sardine, squid), and physical conditions, all of which translated into highly variable CPUE. Catch-per-unit-effort may not be a particularly sensitive metric for the assessment of abundance differences among sites (Beverton and

Holt 1957, Hilborn & Walters 1992), and as such may not be sufficient to detect modest differences in abundance. We measured no significant spatial differences in CPUE for the basses relative to MPA boundaries, although CPUE may not be powerful enough to detect differences at such small spatial scales. The South La Jolla SMR is likely still too young to show a significant MPA effect; however, the Matlahuayl SMR has been protected for over four decades, yet no MPA effect was detected relative to CPUE.

Parnell et al. 2005 found limited signs of an effect within this same MPA, and

highlighted the limited value of small MPAs. Hastings et al. 2014 also found similar fish communities inside and outside this MPA. The extent of the Matlahuayl SMR was expanded in 2012, and this increase in size may yield increased fish abundance in the future.

Catch rates for Barred Sand Bass were far lower than expected, despite focusing our sampling efforts at known spawning locations during spawning season (Jun-Aug).

Based on both landings records and personal communication with CPFV captains, Barred

Sand Bass spawning aggregations did not form as expected at their traditional southern

California sites, which normally include Imperial Beach, San Onofre, Huntington Beach flats, and Ventura flats. Such broad-scale alteration of traditional spawning behavior had never been documented since the aggregations were first discovered decades earlier. In the San Diego area, the 2013 and 2014 spawning seasons consisted of much smaller isolated groups of spawning Barred Sand Bass that were found on isolated rocky reefs off

Imperial Beach, and in the Point Loma kelp forest, and these smaller “satellite” aggregations were targeted heavily by local CPFVs. Barred Sand Bass landings do tend to be lower during warmer years, and 2014 was characterized by anomalously warm sea surface temperatures (Bond et al. in press, Kintisch 2015), with an annual mean value that was surpassed only once in 1997 by temperatures resulting from El Niño (Scripps

Institution of Oceanography Pier water temperature data). The warm waters in 2014 brought offshore species within range of ½-day CPFVs, so most of these vessels were targeting these offshore species when they would normally target Barred Sand Bass.

However, 2013 was much closer to average temperatures extending back to the late

1970s, yet Barred Sand Bass spawning aggregations were virtually absent during both

2013 and 2014, and at all other traditional southern California aggregation sites during both years. Thus, it is likely that factors in addition to warm temperatures have impacted the species.

Differences in the mean size of fishes inside and outside La Jolla reserve sites indicate that reserves are affording some degree of protection for larger fish. A comparison of mean Kelp Bass size between the La Jolla sites indicated that fish were significantly larger inside both reserves, with the largest fish inside the oldest reserve.

California Sheephead also showed significantly larger sizes inside the Matlahuayl SMR.

This species has been heavily targeted for decades by both recreational and commercial fishers (Hamilton et al. 2007); whereas most other fishes in the kelp forests are targeted only by recreational anglers. This higher fishing pressure on California Sheephead outside the MPA may reflect the differences in size between the areas inside and outside of the MPA boundaries. Interestingly, across all sites studied, Kelp Bass were largest in

Imperial Beach, which has always been open to fishing. We believe this finding reflects the relatively limited fishing pressure this site faces relative to Point Loma and La Jolla, combined with the fact that fish densities in the La Jolla reserve sites are at least partially mediated by fishing pressure due to spill-over (the movement of legal fish from inside reserves to open areas). Barred Sand Bass were also relatively large off Imperial Beach, but this is likely because this site is a historical spawning ground for this species. While spawning aggregations were not found, the small numbers of fish we did find were

generally larger, and mature individuals often exhibit fidelity to their spawning grounds

(Jarvis et al. 2010).

Spotted Sand Bass showed a greater abundance of smaller size classes in San

Diego Bay relative to Mission Bay. Most of the smaller individuals from San Diego Bay were caught in the southern half of the bay, which contains a very expansive, shallow eelgrass (Zostera marina) flat where the species generally recruits (Allen 1985, Allen et al. 1995). Mission Bay also has extensive eelgrass habitat, but virtually all of the shallow flats were completely re-shaped by massive dredging and filling operations in the late

1940s. If shallower eelgrass flats are the preferred recruitment habitat for this species, then this could potentially translate to differences in smaller size classes between the two bays.

In addition to spatial differences in size classes, differences in mean size were found between the three sampling years. The sites that have historically been open to fishing, northwest La Jolla and Point Loma, both showed the greatest increases in Kelp

Bass size structure from 2013 to 2014. Point Loma showed the greatest increase in sizes between the two sites, possibly due to its closer proximity to San Diego’s primary fishing port, San Diego Bay, and thus historically higher levels of fishing effort in the area.

Parnell et al. 2010 compared estimates of fishing effort between virtually the same sites as our study, and found greater effort in Point Loma for both recreational private vessel and CPFV fleets. Such higher levels of fishing effort would imply a greater release from fishing pressure for smaller size classes following more conservative regulations, and ultimately an increase in the number of fish at previously legal sizes. Spotted Sand Bass

also showed significant increases in size in both Mission Bay and San Diego Bay following increases in size limits, with a more consistent pattern apparent in San Diego

Bay. Taken together, these findings suggest that anglers are adhering to the new regulations and that these fisheries have responded rapidly to the increased minimum size limits.

Understanding the mortality rates associated with catch-and-release angling is a necessary part of any effort to assess cumulative fisheries impacts. All three of the basses are targeted by a significant portion of the recreational fishing community for catch-and- release purposes only. California Recreational Fisheries Survey data from 2004-2013 indicate that 73% of all Kelp Bass, 55% of all Barred Sand Bass, and 95% of all Spotted

Sand Bass are released by recreational anglers (Holder et al. 2014). We found that both initial and short term mortalities associated with catch and release are surprisingly low at

1.8% and 3.1%, respectively. The CDFW currently assumes a 10% mortality rate associated with catch-and-release. While this may be high relative to the measurements in this study, it also represents a conservative approach for long-term management of the stocks. Our estimates of mortality may under-estimate the total long-term mortality associated with catch-and-release. However, given that catch-and-release mortality usually occurs within just a few days following catch-and-release (Aalbers et al. 2004,

Lowe et al. 2009), it seems unlikely that extending the duration of our holding study would influence our findings. Additionally, the stress of catching and holding the fish in the net pen after an all-day fishing tournament likely imposes additional stress to the fish,

with a concurrent increase in the risk of mortality. As such, we believe our estimates provide an accurate estimate of initial and short-term catch-and-release mortality.

Spatial and temporal analyses of CPUE and length frequency, as well as estimates of initial and short term post-release mortality will ultimately help to inform future stock assessments of the three bass species. The extensive stakeholder collaborations we developed as part of this study were vital to our research success. The scientific community, southern California fishing community, the San Diego Oceans Foundation, and California Department of Fish and Wildlife biologists all combined their knowledge, expertise, and labor to address specific needs for local fisheries management. Hanan and

Curry (2012) had similar success with the recreational fleet in Southern California, and tagged 32,366 rockfish during a 4-year period in southern California. Additionally, the success of the California Reef Check Program demonstrates that recreational enthusiasts can serve as citizen-scientists that provide useful data for subtidal MPA monitoring (e.g.,

Hodgson 2000). The recreational fishing community is a valuable source of information fisheries studies, and future research efforts would significantly benefit from such collaborative partnerships not only with respect to data collection, but by also gaining increased research validation through stakeholder participation.

Acknowledgements

I would like to acknowledge my co-author, Brice Semmens, for his assistance with all aspects of this project.

Figure 3.1. Map of sampling sites off San Diego, California, including MPA boundaries. Dark gray dashed lines represent State Marine Reserve (SMR) boundaries, and light gray dashed lines represent State Marine Conservation Area (SMCA) boundaries.

Fall Summer Fall Sites Inside / Outside 2012 2013 2013 Summer 2014 Matlahuayl SMR inside 3 2 3 3 La Jolla outside 3 2 3 2 South La Jolla SMR inside 3 2 3 2 Point Loma outside 3 2 3 2 Imperial Beach outside 3 3 Long Beach outside 1 Mission Beach outside 1 Encinitas outside 1 Swami’s SMCA inside 1

Table 3.2. Total number of fish caught-and-released for each of the three Paralabrax species at each site (including tournaments), excluding initial mortalities. Total numbers of recaptured individuals by species are also included.

Kelp Barred Sand Spotted Sand Site Bass Bass Bass Total La Jolla 3523 13 3536 Matlahuayl SMR 2766 25 2791 South La Jolla SMR 2670 35 2705 San Diego Bay 71 577 1968 2616 Point Loma 2300 40 2340 Imperial Beach 528 266 794 Encinitas (in/out) 452 70 522 Mission Bay 4 1 363 368 Mission Beach 185 15 200 Long Beach 82 37 119 Newport Bay 22 22

Table 3.3. Total number of records and size range of bycatch species. Size ranges are shown in standard length (mm), but a * denotes total length (mm) and ** denotes fork length (mm). Species with only a single size value only had a single measurement, and some fishes were not measured at all when tagging was too busy. The last row shows 43 individual fish that were not identified because anglers accidentally released them before they were seen by the project scientist. Anglers could usually identify them to rockfishes, croakers, and surfperches, but complete species identification was not possible. Std. Len. Species Scientific Name N (mm) Pacific Barracuda Sphyraena argentea 474 275-765 Brown Rockfish Sebastes auriculatus 341 48-315 Kelp Rockfish Sebastes atrovirens 293 150-308 Bonefish Albula vulpes 264 310 Yellowfin Croaker Umbrina roncador 206 257-293 California Sheephead Semicossyphus pulcher 177 155-565 Gopher Rockfish Sebastes carnatus 134 112-241 Olive Rockfish Sebastes serranoides 110 163-285 White Seabass Atractoscion nobilis 104 301-722 California Halibut Paralichthys californicus 81 204-900 Treefish Sebastes serriceps 68 150-295 Smoothhound spp Mustelus californicus 62 * 514-650 California Scorpionfish Scorpaena guttata 61 170-277 Cabezon Scorpaenichthys marmoratus 54 192-395 Sargo Anisotremus davidsonii 49 220-360 California Lizardfish Synodus lucioceps 49 183-350 California Bat Ray Myliobatis californica 43 no data Vermilion Rockfish Sebastes miniatus 39 140-330 Lingcod Ophiodon elongatus 38 389-765 Round Stingray Urobatis halleri 36 no data Black Surfperch Embiotoca jacksoni 35 143-267 Shovelnose Guitarfish Rhinobatos productus 29 * 1219 Calico Rockfish Sebastes dalli 27 114-277 Copper Rockfish Sebastes caurinus 25 165-298 Giant Kelpfish Heterostichus rostratus 21 215-393 Pacific Bonito Sarda chiliensis lineolata 21 ** 289-378 White Croaker Genyonemus lineatus 16 173-255 Black Croaker Cheilotrema saturnum 13 no data Shortfin Corvina Cynoscion parvipinnis 11 202-435 Butterfly Ray Gymnura marmorata 9 no data

Table 3.3 Continued. Std. Len. Species Scientific Name N (mm) Diamond Turbot Hypsopsetta guttulata 8 no data Sharpnose Seaperch Phanerodon atripes 8 178-220 Grass Rockfish Sebastes rastrelliger 8 175-272 Ocean Whitefish Caulolatilus princeps 7 325-437 Blacksmith Chromis punctipinnis 7 143-178 Opaleye Girella nigricans 7 263-315 Blue Rockfish Sebastes mystinus 7 170-271 Senorita Oxyjulis californica 6 135-183 California Moray Gymnothorax mordax 5 * 560-1000 Horn Shark Heterodontus francisci 5 * 558-777 Black and Yellow Rockfish Sebastes chrysomelas 5 193-214 Leopard Shark Triakis semifasciata 5 * 1270 Halfmoon Medialuna californiensis 3 265-278 White Surfperch Phanerodon furcatus 3 210-230 Rubberlip Surfperch Rhacochilus toxotes 3 275-311 Bocaccio Sebastes paucispinis 3 177 Honeycomb Rockfish Sebastes umbrosus 3 142-173 Rainbow Surfperch Hypsurus caryi 2 188 Starry Rockfish Sebastes constellatus 2 210-235 Squarespot Rockfish Sebastes hopkinsi 2 122-210 Queenfish Seriphus politus 2 no data Spiny Dogfish Squalus acanthias 2 * 960-1080 Fantail Sole Xystreurys liolepis 2 265-343 Common Thresher Shark Alopias vulpinus 1 ** 820 Pacific Sanddab Citharichthys sordidus 1 207 Longfin Sanddab Citharichthys xanthostigma 1 219 Garibaldi Hypsypops rubicundus 1 188 Painted Greenling Oxylebius pictus 1 143 Plainfin Midshipman Porichthys notatus 1 no data Spotfin Croaker Roncador stearnsii 1 no data California Yellowtail Seriola landalii 1 ** 480 Californian Needlefish Strongylura exilis 1 no data Banded Guitarfish Zapteryx exasperata 1 * 943 TOTAL 3005

Initial Initial mortality Tagged-and- Species Caught mortalities (%) released Kelp Bass 12821 240 1.87 12581 Barred Sand Bass 1089 10 0.92 1079 Spotted Sand Bass 2368 15 0.63 2353 Bycatch (other spp) 3049 91 2.98 0 Total 19327 356 1.84 16013

Figure 3.2. Kelp Bass CPUE across primary CPFV charter sites (only includes CPFV charters when Kelp Bass were the target species). Raw data points are overlaid onto boxplots at each site, which include Matlahuayl SMR (n = 11), La Jolla (n = 10), South La Jolla SMR (n = 10), and Point Loma (n = 11).

Figure 3.3. Barred Sand Bass CPUE across sites, which include Long Beach (n =1), Encinitas (n = 1), Swami’s SMCA (n = 1), and Imperial Beach (n = 6).

Figure 3.4. Spotted Sand Bass CPUE by site, which include San Diego Bay (n = 26) and Mission Bay (n = 16). Data are based on private vessel trips when Spotted Sand Bass were the target species.

Figure 3.5. Boxplots of Kelp Bass CPUE by season, which includes fall 2012 (n = 11), summer 2013 (n = 8), fall 2013 (n = 12), and summer 2014 (n = 10). Fall trips were within Sept-Nov, and summer trips were within Jun-Aug periods. Numbers above each maximum represent groupings that resulted from Tukey’s pairwise comparisons. Data only include CPFV charters when Kelp Bass were the target species.

200,000

180,000 160,000 140,000 120,000 100,000 80,000 60,000 40,000 20,000

04 05 06 07 08 09 10 12 13 14

------

CPFV Landings CPFVLandings fish) (no.

Los Angeles Angeles and San Diego Los

Jan

Jan Jan Jan Jan Jan Jan Jan Jan Jan Jan Month - Year

Figure 3.6. California Recreational Fisheries Survey (CFRS) estimates of monthly Barred Sand Bass landings aboard CPFVs based in Los Angeles and San Diego from Jan 2004 – Dec 2014.

Figure 3.7. Boxplots of Spotted Sand Bass CPUE by season, spring (n = 13), summer (n = 2), fall (n = 4), and winter (n = 23). Numbers above each maximum represent groupings that resulted from Tukey’s pairwise comparisons. Data only include private vessel trips in San Diego Bay and Mission Bay when Spotted Sand Bass were the target species.

Figure 3.8. Kelp Bass length frequency distribution at the five primary sites off San Diego.

Figure 3.9. Kelp Bass length frequency by northern San Diego sites.

Figure 3.10. Barred Sand Bass length frequency by site.

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Figure 3.11. Spotted Sand Bass length frequency by primary sites.

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Figure 3.12. Boxplots showing size of the top five demersal bycatch species caught in the primary La Jolla sites. Numbers above each maximum represent groupings within each species that resulted from Tukey’s pairwise comparisons.

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Figure 3.13. Mean (±95% CI) standard length of Kelp Bass (A), Barred Sand Bass (B), and Spotted Sand Bass (C) across primary sites during each of the three sampling years. The gray dashed lines mark the original minimum size limit in 2012 converted to standard length (12 inches total length), and the black dashed line represents the new increased minimum size limit (14 inches total length) that was implemented on March 1, 2013.

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Hastings, P. A., M.T. Craig, B.E. Erisman, J.R. Hyde, and H.J. Walker. 2014. Fishes of Marine Protected Areas Near La Jolla, California. Bulletin, Southern California Academy of Sciences, 113(3): 200-231.

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CHAPTER 4: Movement patterns of three species of Paralabrax using both tag- recapture and acoustic telemetry methods in the coastal marine region of San Diego,

California

Abstract

The effectiveness of fisheries management actions is in part, dependent upon accounting for fish behavior. Spatial management approaches have become common, but they require an understanding of species movement patterns, and harvest control rules can ultimately prove ineffective if these regulations are implemented without the knowledge of spatially mediated life history characteristics (e.g., spawning sites). The state of California recently implemented an extensive network of coastal marine protected areas (MPAs). The degree to which species (and ultimately fisheries) benefit from these closures depends largely on the movement behavior of targeted species.

Understanding the movements and behaviors of fishes relative to reserve boundaries affords the opportunity to quantify the extent of fishing protections (or conversely, spillover) associated with an MPA or a network of MPAs. This can inform both the design and assessment of MPAs. We conducted a fishing community-based tagging study, and integrated both tag-recapture and acoustic telemetry methods to measure movement patterns of three species of Paralabrax on multiple spatial scales. We externally tagged a total of 12,581 Kelp Bass (P. clathratus), 1079 Barred Sand Bass (P. nebulifer), and 2353

Spotted Sand Bass (P. maculatofasciatus), with recapture rates of 10.8% (1362), 4.5%

(49), and 3.8% (90), respectively. Floy tag retention was approximately 81% after 1 year,

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and 50% after 1.75 years for Kelp Bass. Times at liberty ranged from ~1 min - 733 days, and linear distances travelled ranged from 0m - 95km. Most (77%) recaptures were within 0.5 km of the initial tagging site. We also tagged 41 Kelp Bass and 25 Barred

Sand Bass with Vemco coded acoustic transmitters to measure more detailed behavioral patterns. An array of 43 acoustic receivers was used to measure the movements of these individuals throughout the La Jolla kelp bed. Both species showed clear vertical migrations and elevated activity levels during the summer spawning season, and Barred

Sand Bass showed a southward migration to a newly documented aggregation site off south La Jolla. Only 12% of acoustically tagged Kelp Bass were regularly detected across multiple La Jolla sites, while the remaining 88% were primarily detected at a single site.

However, 48% of Barred Sand Bass were regularly detected at two or all three La Jolla sites, and the apparent spawning site for this species spans the southern boundary of the

South La Jolla State Marine Reserve. Increased spatial protection of this spawning aggregation could be achieved by shifting the southern boundary farther south. However, much greater conservational benefits for Barred Sand Bass would likely be achieved through seasonal protection during the June-August peak spawning season.

Introduction

Scientifically informed spatial management initiatives can be successful for recovering and maintaining healthy fish stocks. A classic example of this is the well documented Sockeye Salmon (Oncorhynchus nerka) fishery in Bristol Bay, Alaska where record returns have recently been the product of both favorable environmental conditions

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and management policies informed by spatially explicit stock structure information

(Hilborn et al. 2003). In California, spatial management policies were recently implemented in the form of a statewide network of marine protected areas (MPAs) from

2007-2012. This development process was based largely on lessons learned from the very first MPA network implementation process in California, which focused on the Northern

Channel Islands in 2003 (Botsford et al. 2013, Gleason et al. 2013). Hamilton et al. 2010 conducted an evaluation of this MPA network, and found significantly greater densities and biomass, primarily of popularly fished species, inside the boundaries after only 5 years of protection. This is especially important given the social, economic, and ecological importance of fishing in California (NMFS 2011).

California has a 120-year documented history of recreational and commercial fishing (Ries 1997) with approximately 70% of the annual fishing effort occurring in southern California (CDFW 2014), yet the necessary science to facilitate local stock assessments is often limited, and the population status of some species is still unresolved.

For example, recent debates regarding the population status of two species of Paralabrax in southern California have highlighted data deficiencies that still exist even among these extremely popular fisheries (Erisman et al. 2011, Jarvis et al. 2014a). These species represent ideal examples of the importance fish behavior in influencing fisheries management because they are heavily targeted and exhibit strong site fidelity (e.g., Lowe et al. 2003, Mason & Lowe 2010), and are thus among the most likely candidates to benefit from the spatial protection afforded by MPAs (Gell & Roberts 2003). Despite the importance of understanding fish movements, this information is still a primary source of

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uncertainty in assessing fish populations (Grüss et al. 2011). Data specific to movement patterns and habitat use of coastal demersal species in southern California is still limited to only a few species, frequently at only one or two sites (e.g., Lowe et al. 2003, Topping et al. 2005, Bellquist et al. 2008, Mason & Lowe 2010, McKinzie et al. 2014). Our limited knowledge of fish movements thus leads to inefficiencies in fisheries management, with an incomplete understanding of the effectiveness of MPAs and a failure to account for spawning habitats in fisheries management plans. All three locally common species in the genus Paralabrax, Kelp Bass (P. clathratus), Barred Sand Bass

(P. nebulifer), and Spotted Sand Bass (P. maculatofasciatus) are ecologically, economically, and socially important in southern California and Mexico. California

Commercial Passenger Fishing Vessel (CPFV) logbooks indicate that Kelp Bass and

Barred Sand Bass are likely among the top three most frequently landed species during the entire documented history of California recreational fishing (Hill and Schneider

1999), with approximately 45.3 million fish landed from 1936-2008. Jarvis et al. 2014a note that in recent years, private boaters landed an additional 31% of annual landings for all three species. In total, recreational angling and harvest of these species have a broad range of biological and economic impacts in California. The angling community thus cares deeply about gaining the necessary data to ensure the long-term health of the stocks, and being a part of the fisheries management process (Sholz et al. 2004).

Our goal was to measure movement patterns of all three species of Paralabrax in the San Diego coastal area within a network of MPAs, in contrast to most studies that more typically focus on movement within individual MPAs. To accomplish this, we used

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tag-recapture methods to assess coarse-scale movements for all three species, and acoustic telemetry to generate a more detailed spatial and temporal record of the behaviors of both Kelp Bass and Barred Sand Bass. Acoustic telemetry was not used for

Spotted Sand Bass because this species generally inhabits bays and estuaries, which are separate from the coastal rocky reefs inhabited by Kelp Bass and Barred Sand Bass.

Using the telemetry data, we assessed the movement and behavioral patterns of the two species in the context of reproductive season and spatial management in Southern

California, and provide estimates of the proportional use or reserves, and frequency of spillover. Spillover in this case is defined as any individuals crossing from MPAs to areas open to fishing, independent of the influences of conspecific densities. Using both tag- recapture and acoustic telemetry, we provide an estimate of the extent of protections afforded by reserves off of La Jolla, and characterize the temporal extent and spatial location of behaviors that are consistent with known Barred Sand Bass spawning patterns.

Methods

Study Sites

Our study sites were located primarily along coastal San Diego County, one of the three primary centers of recreational fishing in California, which also include Los

Angeles and San Francisco. While we tagged a small number of individuals off Los

Angeles and Orange Counties, our primary tagging sites from north to south included

Matlahuayl State Marine Reserve (SMR), La Jolla, South La Jolla SMR, Mission Bay,

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Point Loma, San Diego Bay, and Imperial Beach (Fig. 1). Matlahuayl SMR has been protected from all fishing since 1971, while the South La Jolla SMR was implemented in

January 2012. The La Jolla, Point Loma, Mission Bay, and San Diego Bay sites have always been open to recreational fishing. Mission Bay was historically a tideland that was dredged and reshaped into a recreation area in the late 1940s. Similarly, San Diego Bay has undergone substantial anthropogenic and urban modifications, with 37% of the shoreline owned and 19.8% of the total land and water area occupied by the U.S. Navy

(San Diego Bay Integrated Natural Resources Management Plan, 2013, pg. 221).

Traditionally, Barred Sand Bass spawning aggregations occur at the Imperial Beach site from June-August (Jarvis et al. 2010). Collectively, these sites represent a broad range of angler-targeted habitats and locations with differing spatial protections. Tagging Methods

Movement patterns were measured using both tag-recapture methods and acoustic telemetry. Our tag-recapture methods consisted of standard Floy® T-Bar Anchor tags applied with Avery Dennison Mark III tagging guns. In total, we tagged 16,013 individuals of the three bass species combined over a 2 year period. Of these, we double- tagged 673 Kelp Bass in order to measure tag loss rates. Each tag was inserted in the dorsal musculature between the pterygiophores beneath the anterior portion of the dorsal fin. Printed on each tag was the individual ID number as well as a phone number for reporting recaptures. We also created a website (www.cooperativefishtagging.org) and a freely available smartphone application (CatchReporter) for anglers to report tagged fish.

The acoustic telemetry approach consisted of Vemco model V13 coded acoustic transmitters, which only measure presence/absence near a receiver, and V13AP coded

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acoustic transmitters, which measure presence/absence, acceleration, and depth at the time of signal transmission. Transmitters reported randomly within 100-200sec intervals, yielding an approximate battery life of 440-480 days after correcting for time of deployment. Fish were surgically fitted with either the V13 or V13AP transmitters using methods that mirrored other local telemetry studies (e.g., Lowe et al. 2003, Topping et al.

2005, Bellquist et al. 2008, Mason & Lowe 2010, McKinzie et al. 2013) under approved

IACUC protocol S12116. After release, each tagged fish was detected by an array of 44

Vemco VR2W acoustic receivers attached to subsurface moorings at fixed locations (Fig.

2, www.LaJollaArray.org). Detections from coded transmitters provided receiver-specific presence/absence data, and for a subset of tags, depth and tag acceleration profiles at 2- minute intervals.Data Analysis

Tag-recapture data were collected for all three bass species directly aboard the

CPFV tagging charters and private boat trips, as well as from tag reports by recreational anglers. Anglers had the choice of reporting tags by calling the phone number printed on the tag, entering the tag information on the project website, or by submitting the information with a new smartphone application that we developed (CatchReporter).

Recapture data were used to measure coarse-scale movement and recapture rates between sites both within the La Jolla kelp bed (encompassed by the array) and outside the La

Jolla sites. We used linear regression to analyze the log of total distance traveled with respect to time at liberty and individual size at the time of tagging. Measurements of distance traveled were only possible when recapture locations had reliable latitude- longitude coordinates associated with them. Frequency distributions of time at liberty,

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distance traveled, and tag-recapture locations were generated, and logistic regression was used to estimate Floy® Tag loss rates over time based on recaptures of double-tagged individuals.

Acoustic telemetry data were used to measure seasonal movements of both Kelp

Bass and Barred Sand Bass throughout the La Jolla kelp bed. All acoustic data were download managed in the Vemco NUE software. Analysis of variance and Tukey’s post hoc tests were used to test for seasonal effects on depth and acceleration. The percent of detections at each receiver were plotted using a Geographic Information System (GIS), and split between seasons to show spatial changes in detection patterns for each species.

The proportion of detections for each individual was also measured as a proxy for time spent inside/outside MPA boundaries.

Results

A total of 12,581 Kelp Bass, 1,079 Barred Sand Bass, and 2,353 Spotted Sand

Bass were Floy tagged from Oct 2012 – Sep 2014 at 11 sites from Long Beach,

California south to the U.S. - Mexico border. However, the majority of individuals were tagged at the primary sites off San Diego County (Fig. 1). A total of 1362 Kelp Bass, 49

Barred Sand Bass, and 90 Spotted Sand Bass recaptures were recorded during this period, with site-specific recapture rates ranging from 0-17.5% (Table 1). Among the Kelp Bass,

1,207 fish were recaptured once, 132 were recaptured twice, 22 were recaptured three times, and 1 individual was recaptured 4 times during the study period. Times at liberty among recaptures ranged from ~1 minute to 733 days (Figures 2-4), with 60 individuals

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recaptured on the same day they were tagged. In addition, a total of 41 Kelp Bass and 26

Barred Sand Bass were tagged with coded acoustic transmitters at the three La Jolla sites, which contained an array of 43 underwater acoustic receivers attached to fixed subsurface moorings (Fig. 5).

Tag Loss Rates

A total of 673 Kelp Bass were double tagged across multiple sites, and 113 of these were recaptured with one or both tags still attached. Logistic regression revealed that the probability of Floy tag loss was related to time at liberty (p < 0.001). After one year at liberty, the probability of tag retention was approximately 80.8%, and decreased to 62% after 1.5 years, and 50% after 1.75 years (Fig. 6). Tag retention rates were not estimated for Barred Sand Bass or Spotted Sand Bass.

Movement based on tag-recapture

Tag recapture analyses show limited movements for all three bass species overall, with the majority of individuals recaptured at the site of original tagging (Tables 2-4).

However, several instances of movement between sites were detected. For example, among the Kelp Bass recaptures that were originally tagged inside the Matlahuayl SMR, approximately 14.8% (71/481) were recaptured outside the SMR boundaries (3 of these were recaptured as far away as San Clemente, California). In comparison, among the 517 recaptures that were originally tagged in La Jolla (open to fishing), only 1.9% (10/517) were recaptured inside the adjacent Matlahuayl SMR boundaries. However, there is a

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much greater probability of fishes being recaptured outside the SMR boundaries because the area is open to fishing, and is subject to legal recaptures from the fishing community.

Thus, if only data from tagging charters are taken into account (i.e. if we standardize for fishing effort), then 3.1% of fish that were tagged inside the SMR were recaptured outside, while 5.8% that were tagged outside the SMR were recaptured inside the boundaries aboard subsequent tagging charters. The farthest distance recorded among all three species was a Barred Sand Bass that was tagged at Shelter Island in San Diego Bay, and recaptured 95.4 km away off Dana Point (Table 3). All Spotted Sand Bass were recaptured in the same bay they were originally tagged (Table 4). Kelp Bass showed no relationship between time at liberty and either total distance traveled (Fig. 7, F = 2.47, df

= 1, p = 0.12) or standard length (F= 2.89, df = 1, p = 0.09), and almost 86.2% of individuals were recaptured within 1km of their tagging location, regardless of time at liberty (Fig. 8). Barred Sand Bass also showed no relationship between distance traveled and either time at liberty (F = 3.20, df = 1, p = 0.08) or standard length (F = 2.15, df = 1, p = 0.15), although relatively few recaptures were reported with reliable spatial information that would allow travel distance calculations (Fig. 9). Spotted Sand Bass showed no significant relationship between distance traveled and either time at liberty

(Fig. 10, F= 3.03, df = 1, p = 0.09) or standard length (F = 3.17, df= 1, p = 0.08), and no individuals traveled between San Diego Bay and Mission Bay. Only 42.6% (640) of our total recaptures were recorded directly on CPFV tagging charters, while the remaining

57.4% (861) were reported by the recreational fishing community. Of the tags reported

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by anglers, 13.1% of these were reported directly to the SIO researcher via community contacts that we developed during the course of the project.

Movement and behavior based on acoustic telemetry

Acoustic telemetry results provide additional detail to movement analyses for both Kelp Bass and Barred Sand Bass. Both species showed distinct seasonal differences in depth utilization (Fig. 11), with vertical movements in the water column during warmer summer months. Kelp Bass used shallower depths from Jun-Aug in both 2013 and 2014 relative to winter months, although these vertical summer movements were shallower in 2013 than 2014. Barred Sand Bass also showed shallower depth use with greater variability during the summer, but the time period of these vertical movements ranged from Apr-Aug, and was generally deeper than for Kelp Bass. In addition, from

Sep 2014 – Jan 2015 (outside spawning season), Barred Sand Bass depth of occurrence remained relatively shallow, but decreased in monthly variability. Seasonal differences were also apparent for both species with respect to acceleration (Fig. 12), which we assume to reflect the general activity levels of individuals. Both species showed greater activity during the summer months of both 2013 and 2014, and Kelp Bass showed slightly greater activity than Barred Sand Bass for almost every month. Variability in activity was generally similar for both species across all months.

In addition to vertical and behavioral measurements, seasonal horizontal movement was measured for both species between the three La Jolla sites. Kelp Bass did not show strong seasonal differences in detections at specific receivers. However, Barred

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Sand Bass showed southward movements during late spring and early summer to the southern end of the South La Jolla SMR, which is an area dominated by broad sand flats adjacent to kelp forest habitat (Fig. 13). Movement patterns for two individuals are shown in Fig. 13. One migrated from northern La Jolla so the South La Jolla SMR (Fig.

13a), and another from central La Jolla to the South La Jolla SMR (Fig. 13b). Upon reaching this broad sand flat, both individuals began vertical movements (and greater acceleration) that are consistent with known spawning behaviors (McKinzie et al. 2014).

One individual only showed vertical behavior for a brief period of time (Fig. 13a), likely because this fish was primarily using an area just outside the detection range of the receivers. However, the other individual showed strong vertical movements from early

June to late August, which implies an approximate 3-month overall spawning duration for this individual. This duration is consistent with spawning estimates from batch fecundity analyses (Jarvis et al. 2014b). Detections for both species decreased at the end of 2014 presumably because this was the projected end date of the transmitter batteries.

Kelp Bass spent the majority of time within the sites where they were originally tagged (Fig. 14), while Barred Sand Bass exhibited more regular movements between the

La Jolla Sites (Fig. 15). Eleven of fifteen Kelp Bass tagged inside the Matlahuayl SMR were detected over 90% of the time inside the SMR (Fig. 14a). Only 1/40 Kelp Bass traveled between all three sites in La Jolla, while 3/24 Barred Sand Bass traveled between all three sites. Barred Sand Bass that were tagged inside the South La Jolla SMR showed a much broader range of percent detections inside vs. outside the SMR (Fig. 15c). While spatial patterns in detections are largely dependent upon where individuals were initially

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tagged, we achieved relatively even spatial distribution of tags deployed (Fig. 5a).

Overall, Kelp Bass and Barred Sand Bass showed 72.7% and 15.7% of all detection inside the SMRs, respectively (Table 5).

Discussion

Understanding fish movement patterns is essential for fisheries management, especially given the relatively recent development of spatially explicit stock assessment models that are beginning to inform new marine spatial planning initiatives (Hilborn

2012). Past stock assessments for demersal species in southern California often assumed no movement by or no variation in movement among individuals (e.g., Key et al. 2005), but studies are showing that some species travel much farther than originally thought. For example, Hanan & Curry 2012 found that nine species of rockfish (Sebastes sp.) were recaptured greater than 50km from their initial tagging site in southern California. Such information provides a more complete knowledge base of fish behaviors that can be used to inform fisheries management more effectively.

While fine-scale movement patterns have been studied for Kelp Bass (Lowe et al.

2003) and Barred Sand Bass (Mason and Lowe 2010), these studies were both conducted at the same individual MPA site at Santa Catalina Island. A benefit of conducting similar research along the mainland coast was that it allowed us to measure movement patterns relative to a network of MPAs rather than a single MPA. In addition, habitat and bathymetric characteristics differ between the islands and the mainland, so movement patterns of these two species are likely to be different between the two regions. Barred

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Sand Bass are also generally more abundant along the mainland coast, so research focusing on mainland movements represents additionally valuable information about the stock. Jarvis et al. (2010) used tag-recapture data from both the 1960s and 1990s to document seasonal spawning aggregations of Barred Sand Bass along the mainland. In addition McKinzie et al. (2013) used acoustic telemetry to measure fine-scale spawning and non-spawning behaviors of Barred Sand Bass at one of these aggregations sites off

Huntington Beach, California. However, the data from these two studies were collected while the Barred Sand Bass were still following traditional spawning behavior at historically consistent aggregation sites. In both 2013 and 2014 there was a virtual absence of spawning aggregations at the traditional sites in southern California. Our results indicate that this has resulted in a shift from the traditional large aggregations on broad sand/mud flat habitat to much smaller aggregations within kelp forest habitat.

Our tag-recapture results varied by species and by site, with the greatest success being achieved largely with Kelp Bass in the La Jolla and Point Loma sites, and with

Spotted Sand Basses in San Diego Bay. Overall recapture rates were 10.83% for Kelp

Bass, 4.54% for Barred Sand Bass, and 3.82% for Spotted Sand Bass, and these rates continue to grow because we are still collecting new recapture reports from anglers. Floy tag retention rates were sufficiently high to measure movements over relatively long time periods for Kelp Bass (almost 2 years), but retention rates were not measured for the other two congeners. Given that 60 Kelp Bass were recaptured on the same day they were tagged, and that time at liberty was as short as ~1 minute, it appears that catch-and- release fishing imposes surprisingly low stress on this species. On the other hand, at least

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some individuals are adversely impacted, with 1.7% suffering initial mortality upon capture, and an additional 3.1% succumbing over an extended period of time.

The vast majority of Kelp Bass were recaptured within 1km of their tagging site regardless of time at liberty. Collyer and Young (1953) reported the first known tag- recapture results for Kelp Bass, and found similarly limited movements among the majority of recaptured individuals. Young (1963) also found similar results, with 364/458 individuals recaptured at the same site as initial tagging. In our study, movements relative to the Matlahuayl SMR boundaries showed similarly low rates of between-site movement for Kelp Bass. While most results a show limited scale of movement, several individuals in our study traveled up to 80km from the initial tagging site. Kelp Bass are not known to migrate such distances for spawning purposes, and there was no relationship between individual size and distance traveled, so the reasons behind these rare but distant movements remain unclear.

Annual spawning migrations are exhibited by Barred Sand Bass, and their spawning aggregation sites are highly predictable, well-documented, and heavily targeted by recreational fishing fleets (Turner et al. 1969; Feder et al. 1974; Love et al. 1996a,b;

Jarvis et al. 2010). The primary aggregation site off San Diego is the broad sand flat off the coast of Imperial Beach and Tijuana, Mexico (Jarvis et al. 2010). Our sampling effort was designed to take advantage of this aggregation site, and tag a large number of individuals with relatively few trips during the spawning season. However, Barred Sand

Bass exhibited a virtual absence of traditional spawning behavior during the summer in

2013 and 2014, not only in this aggregation site, but in all known southern California

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sites (e.g., Imperial Beach, San Onofre, Huntington Beach flats, Santa Monica Bay, and

Ventura Flats). As a result, the majority of Barred Sand Bass in our study were tagged at isolated rocky reefs off Imperial Beach during the spawning season (summer), and in San

Diego Bay outside the spawning season (winter). Our tag-recapture results did show the majority of movement occurring between Imperial Beach, San Diego Bay, and Point

Loma, so it is possible that individuals that make up the spawning aggregation at Imperial

Beach generally come from the three sites combined. However, sample sizes are too low for a definitive conclusion. This is cause for serious concern given not only the susceptibility of aggregating species to overfishing, but also that fishing has already caused the depletion of other fishes that exhibit similarly predictable aggregating behaviors in other places (Sadovy & Eklund 1999, Sala et al. 2001, Whaylen et al. 2004).

While tag-recapture methods were not as successful as we hoped for quantifying movements at the Imperial Beach spawning site, acoustic telemetry methods did shed light on potential spawning behaviors in southern La Jolla. Seventy percent of Barred

Sand Bass tagged with acoustic transmitters showed clear affinity for an area just south of the South La Jolla SMR during their known spawning season (Jarvis et al. 2014b). This area is a broad sand flat adjacent to kelp forest habitat, which is consistent with the depth and habitat at other known spawning aggregation sites. Some of these fish traveled from as far away as Matlahuayl SMR directly to this area, and upon arrival, all individuals performed daily vertical migrations in the water column with greater acceleration rates than at other times. All of these behaviors are consistent with known spawning behaviors at other known spawning sites (McKinzie et al. 2014). Kelp Bass also exhibited vertical

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migrations, but this occurred at areas around the entire La Jolla kelp bed. In addition, there were clear differences between the two species in vertical behaviors. Kelp Bass ascended high in the water column during the spawning season in June and July, with a consistent vertical separation from the benthic substrate during these months. This was based on the depth of the tagged individual relative to the known depths surrounding each receiver where fish were detected. Barred Sand Bass also rose vertically in the water column during their spawning period but consistently returned to the bottom, i.e. they did not remain high in the water column for extended periods of time as did the Kelp Bass. In winter of 2014-2015, it appears as though Barred Sand Bass remain high in the water column, but these individuals simply moved to shallower areas while staying near the bottom (Fig.11). If Barred Sand Bass were spawning in this area, then this is the first quantitative documentation to our knowledge of this spawning aggregation site. This site is partially protected by the boundaries of the South La Jolla SMR, but the areas open to fishing in south La Jolla are characterized by high fishing effort among both CPFV and private vessel fleets (Parnell et al. 2010). Parnell et al. 2006 conducted analyses of physical and biological habitat within the entire La Jolla kelp forest, and suggested specific northern and southern boundaries for this SMR (32o 49.5’N and 32o 48.0’N, respectively). The boundaries that were ultimately adopted encompass an area slightly greater in size (32o 49.573’N and 32o 47.945’N), but our telemetry data suggest that a

500m southward shift of the southern boundary would encompass a much greater portion of the Barred Sand Bass spawning site. However, specific site protection within the study area may produce the best conservational value by focusing on the primary aggregation

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site off Imperial Beach, California and Tijuana, Mexico. Recreational CPFV logbook data show that 60% of Barred Sand Bass landed in this area in 2013 and 2014 were from

Mexican waters off Tijuana, meaning that the traditional spawning site off Imperial

Beach has recently either shrunk or moved to Mexican waters. A joint management initiative between both countries would thus be required if spawning site/season protection became a priority for fisheries managers.

Despite our findings of a potentially new spawning aggregation site, Barred Sand

Bass did not form their traditional aggregation off Imperial Beach during our sampling period. Rather, smaller aggregations were located by the CPFV fleet and heavily targeted in the Point Loma and Imperial Beach kelp forests. The aggregation site in south La Jolla may be another of these smaller aggregation sites that apparently occurred in the absence of the larger primary aggregation site. Aburto-Oropeza et al. 2008 note that Barred Sand

Bass are a commercially important species targeted along the Baja California coast, and it is essential to understand whether other aggregation sites in Baja are exhibiting declines similar to those in southern California.

The array of acoustic receivers distributed throughout the three La Jolla sites allowed us to assess the protections afforded individuals by the existing MPA network.

Kelp Bass exhibited very limited movement between the three La Jolla sites, with most individuals being detected almost entirely at the site where they were initially tagged.

Barred Sand Bass exhibited greater rates of exchange between protected and unprotected sites, suggesting that the MPA network in La Jolla affords less protection for Barred Sand

Bass than for Kelp Bass. This is especially true given that much of the spawning

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aggregation behaviors observed for Barred Sand Bass occurred just outside of the South

La Jolla SMR. The trans-boundary movements of individuals documented in this study may or may not represent instances of density-dependent spillover, but fishes of both species are clearly leaving the MPA boundaries for varying periods of time. Future analyses of movements will hopefully include integration with high resolution habitat maps that were recently generated for the La Jolla and Point Loma areas (Parnell et al.

2015). Spotted Sand Bass exhibit complex spawning behaviors and spatial variation in protogynous hermaphroditism (Hastings 1989, Allen et al. 1995, Hovey & Allen 2000).

Allen et al. 1995 suggest that Spotted Sand Bass form spawning aggregations at the entrance of bays and estuaries during late spring and summer, although their sources are primarily anecdotal. While we did find areas within San Diego Bay that were characterized by exceptionally high catch rates, these areas were sampled outside the summer spawning season. In San Diego Bay, tagged fish from the southern portion of the bay were generally smaller and caught at shallower depths, while larger individuals were caught in the northern portion and in deeper channels. This segregation of size classes may reflect an ontogenetic shift from shallow eelgrass (Zostera marina) flats to deeper channel habitats. A more prolonged and intensive mark-recapture study would be required to definitively demonstrate this shift.

The success of this project was largely due to our collaborative partnerships with the southern California fishing community, the California Department of Fish and

Wildlife, and the San Diego Oceans Foundation. The level of cooperation between these stakeholder groups yielded significant benefits, especially with respect to data quality and

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quantity, and established strong collaborative relationships for future projects. Our final recapture rates show that 57.4% of our total recaptures were reported by the recreational fishing community. The support we received from the recreational fishing community with respect to tagging efforts and recapture reporting was thus fundamental to our success, and the Sportfishing Association of California was a strong facilitator of this collaborative progress.

Acknowledgements

I would like to acknowledge my co-author, Brice Semmens, for his assistance with all aspects of this project.

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Figure 4.1. Site map of the San Diego coastal area. Dark gray solid lines represent Matlahuayl and South La Jolla SMR boundaries, and light gray lines represent SMCA boundaries. The gray dashed line represents the U.S.-Mexico maritime border. Black dots in the three La Jolla sites represent fixed locations of Vemco VR2 acoustic receivers attached to subsurface moorings.

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Table 4.1. Total number of individuals tagged and percent recapture rates for each of the three species by tagging site. Sites include Long Beach (LB), Newport Bay (NB), north San Diego County (NC), Matlahuayl SMR (M), La Jolla (LJ), South La Jolla SMR (SLJ), Mission Beach (MB), Mission Bay (MBy), Point Loma (PL), San Diego Bay (SDB), and Imperial Beach (IB).

Kelp Bass Barred Sand Bass Spotted Sand Bass Number Recapture Number Recapture Number Recapture Site Tagged Rate (%) Tagged Rate (%) Tagged Rate (%)

LB 82 4.88 37 2.70

NB 22 9.09

NC 452 8.41 70 2.86

M 2766 15.18 25 4.00

LJ 3523 15.38 13 7.69

SLJ 2670 1.80 35 2.86

MB 185 9.19 15 6.67

MBy 4 0 1 0 363 6.34 PL 2300 9.83 40 17.50

SDB 71 4.23 577 3.99 1968 3.30 IB 528 4.17 266 4.14

Total 12581 10.83 1079 4.54 2353 3.82

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350

300

250

200

150

100

Recapture Recapture frequency 50

100 150 200 250 300 350 400 450 500 550 600 650 700 Time at liberty (days)

Figure 4.2. Kelp Bass recapture frequency. Data only include the most recent recapture for each individual.

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16 14 12 10 8 6 4 2

Recapture Recapture frequency 0

400 150 200 250 300 350 450 500 550

100

------

351 101 151 201 251 301 401 451 501 Time at liberty (days)

Figure 4.3. Barred Sand Bass recapture frequency. Data only include the most recent recapture for each individual (excluding tournaments).

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45 40 35 30 25 20 15 10

Recapture Recapture frequency 5

200 500 150 250 300 350 400 450 550 600 650 700

100

------

> 700 >

151 451 101 201 251 301 351 401 501 551 601 651 Time at liberty (days)

Figure 4.4. Spotted sand bass recapture frequency. Data only include the most recent recapture for each individual (excluding tournaments).

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Figure 4.5. Map of acoustic transmitter deployment locations for Kelp Bass and Barred Sand Bass (a), and map of the array of acoustic receivers at the three La Jolla sites (b). The dark gray area represents land, and the light gray polygon represents the average spatial extent of the La Jolla kelp forest. In panel A, the black and gray dots represent acoustic tagging locations of Kelp Bass and Barred Sand Bass, respectively. In panel B, each black dot represents the location of a Vemco VR2W acoustic receiver attached to a subsurface mooring.

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Figure 4.6. Logistic regression results showing the probability of tag retention over time among double-tagged Kelp Bass. The solid black line represents the predicted values of tag retention probability, and the dotted lines represent upper and lower 95% confidence limits. Black points along the top and bottom are shown relative to the secondary (right) Y-axis, and represent double-tagged individuals that were recaptured with both tags and only one tag remaining, respectively.

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Table 4.2. Kelp Bass recapture frequencies for the following tagging and recapture sites: Long Beach (L), San Clemente (S), north San Diego County (N), Matlahuayl SMR (M), La Jolla (LJ), South La Jolla SMR (SLJ), Mission Beach (MB), Mission Bay (MBy), Point Loma (PL), San Diego Bay (SDB), Imperial Beach (IB), and unknown location (U).

Recapture Sites Site L S N M LJ SLJ MB MBy PL SDB IB U Total LB 4 4 SC 0 NC 33 2 35 M 3 4 409 60 1 1 2 1 481 LJ 1 10 465 7 3 3 3 25 517 SLJ 1 14 40 2 1 1 2 61 MB 1 5 1 7

Tagging Sites MBy 0 PL 1 1 2 214 1 7 226 SDB 3 3 IB 5 21 2 28 Total 4 5 38 420 542 48 11 6 226 3 22 37 1362

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Table 4.3. Barred Sand Bass recapture frequencies for the following tagging and recapture sites: Long Beach (LB), Dana Point (DP), north San Diego County (NC), Matlahuayl SMR (M), La Jolla (LJ), South La Jolla SMR (SLJ), Mission Beach (MB), Mission Bay (MBy), Point Loma (PL), San Diego Bay (SDB), Imperial Beach (IB), and unknown location (Unk).

Recapture Site Site LB DP NC M LJ SLJ MB Mby PL SDB IB Unk Total LB 1 1 DP NC 1 1 M 1 1 LJ 1 1 SLJ 1 1 MB 1 1

Tagging Site Mby PL SDB 1 6 20 5 1 33 IB 1 3 6 10 Total 1 1 1 1 1 1 1 7 23 11 1 49

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Recapture Site Site NB MBy SDB Total

NB 2 2

MBy 22 22 Site

Tagging SDB 65 65 Total 2 22 65 89

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70 60 50 40 30

20 Distance traveled Distance traveled (km) 10 0 0 100 200 300 400 500 600 700 800 Time at liberty (days)

Figure 4.7. Total distance traveled versus time at liberty by Kelp Bass.

137

300

250

200

150

Frequency 100

2.1 2.4 2.7 0.0 0.3 0.6 0.9 1.2 1.5 1.8 3.0 3.3 3.6 3.9 4.2 4.5 4.8 5.1 5.4 5.7 6.0 6.3 6.6 6.9 Distance traveled (km)

Figure 4.8. Frequency distribution of total distance traveled by Kelp Bass.

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16 14 12 10 8 6

4 Distance traveled Distance traveled (km) 2 0 0 100 200 300 400 500 600 Time at liberty (days)

Figure 4.9. Total distance traveled versus time at liberty by Barred Sand Bass.

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Distance traveled (km) traveledDistance 2

0 0 100 200 300 400 500 Time at liberty (days)

Figure 4.10. Total distance traveled versus time at liberty by Spotted Sand Bass.

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Figure 4.11. Boxplots showing depth utilization by month for both Kelp Bass and Barred Sand Bass in the three La Jolla sites.

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Figure 4.12. Boxplots showing median acceleration by month for both Kelp Bass and Barred Sand Bass in the three La Jolla sites.

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Figure 4.13. Horizontal movement, depth use, and acceleration of two individual Barred Sand Bass. Individual 1 is represented by panels A, C, and E, and individual 2 is represented by panels B, D, and F. The top panels show horizontal movement between receiver locations from north to south, the middle panels represent individual depth detections at each day/time, and the bottom panels represent individual acceleration detections at each day/time.

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Figure 4.14. The proportion of detections at each of the three La Jolla Sites for Kelp Bass that were initially tagged in the Matlahuayl SMR (a), in La Jolla (b), and in the South La Jolla SMR (c).

144

145

Table 4.5. The total number of detections and percentage of detections for each species both inside and outside the two MPAs in La Jolla.

Total Detections Percent Detections Kelp Barred Sand Kelp Barred Sand Species Bass Bass Bass Bass Inside 250,797 24,169 72.7 15.7 Outside 94,394 129,447 27.3 84.3 Total 345,191 153,616

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