Jarvisetal bsb movements_01_mar10

SPAWNING-RELATED MOVEMENTS OF BARRED SAND BASS, PARALABRAX 1

NEBULIFER, IN SOUTHERN CALIFORNIA: INTERPRETATIONS FROM TWO DECADES 2

OF HISTORICAL TAG AND RECAPTURE DATA. 3

E.T. Jarvis, C. Linardich, and C.F. Valle. California Department of Fish and Game, Marine 4

Region, Los Alamitos, CA, 90720. 5

2

Abstract 6

During the 1960s and 1990s, the California Department of Fish and Game tagged 8,634 7

barred sand bass in southern California. Tag and recapture data suggest migrant individuals reside 8

at spawning grounds for approximately one month. Although 64% of non-spawning season 9

recaptures occurred within 1 km of the spawning location, the average (± SD) recapture distance of 10

migrant fish was 13 ± 8 km. Fish size (TL) was weakly correlated with “migration” distance from 11

spawning locations (rs(57) = 0.31, p = 0.02). Spatial and temporal trends in recaptures suggest a 12

high degree of spawning and non-spawning season site fidelity. 13

3

Introduction 14

Barred sand bass, Paralabrax nebulifer, continues to be one of the most sought-after sport fish 15

in southern California. In the early 1900s, barred sand bass were landed in both the commercial 16

and recreational fisheries; however, due to limited demand in the commercial fishery and scarcity 17

of the resource during the 1950s, commercial take was banned in 1953 and a 12 in (30.5 cm) 18

minimum size limit was implemented in 1959 (Collyer 1949; Young 1969). Since the 1960s, 19

barred sand bass have ranked among the top 10 sport fish caught by commercial passenger fishing 20

vessels (CPFVs = party boats) in southern California, with total annual catches averaging nearly 21

two million fish per year between 1980 and 2003* (Allen and Hovey 2001; PSMFC 2010a). 22

Although no commercial fishery currently exists for barred sand bass in California, “heavy annual 23

landings” of barred sand bass have been reported in the commercial fisheries of Baja California, 24

Mexico, and the Gulf of Mexico in recent years (Aburto-Oropeza et al. 2008). 25

In southern California, fishing effort for barred sand bass by both party and private boaters is 26

highest during peak spawning season (June to August). In recent years, approximately 77% of the 27

June, July, and August barred sand bass harvest was taken by party/charter boats (PSMFC 2010b). 28

Well-known spawning aggregation sites include the Ventura Flats, inner Santa Monica Bay, 29

Huntington Beach Flats, San Onofre, and Silver Strand in San Diego. According to California 30

Department of Fish and Game (CDFG) CPFV logbook data, Huntington Beach Flats ranks the 31

highest in numbers of barred sand bass caught in the past 40 years (CDFG unpublished data). 32

From 1961 to 2008, barred sand bass CPFV catch-per-unit-effort (CPUE = barred sand bass 33

kept/angler*hr) increased to a high in 2000, and has recently decreased to the lowest CPUE since 34

*Data reported were collected by the Marine Recreational Statistics Survey (MRFSS), which was replaced in 2004 by the California Recreational Fisheries Survey (CRFS). Catch estimates after 2003 were not included because data from these two surveys are not comparable.

4

1985 (CDFG unpublished data), causing concern regarding the vulnerability of the population to 35

future harvest impacts. 36

Fish species that are targeted during their spawning aggregations are especially susceptible to 37

overexploitation because harvest effects may not be immediately evident (Sadovy and Domeier 38

2005). Furthermore, overexploitation of an aggregate spawner can result in the complete absence 39

of spawning aggregation formations at historic sites (Domeier and Colin 1997; Sadovy and 40

Domeier 2005). Consequences of this occurring in the barred sand bass fishery might include 41

negative ecological impacts, as well as severe economic impacts to the sport fishing industry in 42

southern California. 43

Barred sand bass is one of three temperate serranine fishes (sea basses), including kelp 44

bass, Paralabrax clathratus, and spotted sand bass, Paralabrax maculatofasciatus, that participate 45

to varying degrees in aggregate spawning behavior (Hovey and Allen 2000; Erisman and Allen 46

2006; Miller and Allen 2006). Barred sand bass breeding aggregations are reported to occur over 47

sand flats in depths of 20 to 40 m (Turner 1969; Feder et al. 1974; Love et al. 1996), and based on 48

the exceptionally high landings of barred sand bass during the summer, it is possible that these 49

aggregations consist of thousands of fish. However, underwater video documentation of these 50

aggregations has never been reported. Following peak spawning, considerably fewer barred sand 51

bass are caught over the sand flats and catches typically resume inshore in bays or near low relief 52

natural or artificial reefs, but not in such high numbers (Love et al. 1996). These catch trends may 53

suggest transient spawning aggregation behavior, in which large aggregations form at specific, 54

predictable locations at higher than average densities for a period of several weeks to months 55

(Domeier and Colin 1997). Transient spawning aggregations are characterized by having 56

individuals that migrate relatively longer distances to breed than “resident” aggregate spawners, 57

5

but it is still unclear to what degree this may occur with barred sand bass. Knowledge of the 58

origins and destinations of barred sand bass spawning migrations, and understanding whether all or 59

some of the adult population participates in spawning aggregations will be important for effective 60

management of this species. 61

In the 1960s and 1990s, biologists with the California Department of Fish and Game (CDFG) 62

tagged nearly 9,000 barred sand bass in southern California. The recapture information from these 63

two time periods enables us to document the historical spawning-related movements of barred 64

sand bass for the first time. Specifically, our objectives of this study are to examine these 65

historical data for trends in 1) residency at spawning locations, 2) movement to and from spawning 66

locations, and 3) breeding site fidelity. 67

68

Methods 69

Tagging Effort 70

During the 1960s and 1990s, barred sand bass were tagged along the coast of southern 71

California and at one location in Baja California, Mexico (Figure 1). Tagging locations included 72

sand flats, reefs, and bay habitat. During both tagging periods, fish were captured by hook-and-73

line, measured to the nearest mm total length (TL), externally tagged with spaghetti or T-bar tags, 74

and released. In the 1990s, fish were also captured by bottom trawl, and upon release, tagged fish 75

suffering from barotrauma were recompressed to depth using milk crates. Loran or GPS 76

coordinates of the tagging sites were recorded; otherwise, a site name or geographic landmark was 77

provided. In addition, depth (m) and release condition were recorded for some but not all fish. 78

Rewards for recaptures of tagged fish were offered during both tagging periods. Recapture 79

6

information included date, location, TL (mm), and tag ID number. In the 1990s, recapture depth 80

(m) and Loran or GPS coordinates were also provided when available. 81

Analysis 82

All historical barred sand bass tag and recapture data were archived into a relational 83

database for analysis. To standardize tagging effort across the two tagging periods, locations for 84

all records were assigned a fishing site code based on historical southern California CPFV sport 85

fish surveys (Ally et al. 1990). Spawning season codes were also assigned to each tag and 86

recapture record based on capture month (Nov.–Mar. = non-spawning season, Apr.–May = early 87

spawning season, Jun.–Aug. = peak spawning season, Sept.–Oct. = late spawning season). Days at 88

liberty, recapture distance (estimated or actual km), and general direction of movement were 89

calculated and incorporated into the database. Recapture distances were measured as linear 90

distances between approximate or actual tagging and recapture locations. In this paper we report 91

recapture rates and return rates. Recapture rates refer to the number of fish recaptured at a given 92

site divided by the total number fish recaptures. Return rates refer to the number of fish recaptured 93

at a given site divided by the total number of fish tagged at that site. 94

Spawning season residency 95

To investigate the residence time of individuals at spawning locations, we selected fish 96

tagged at Huntington Beach Flats during peak spawning season and recaptured at Huntington 97

Beach Flats within the same year (peak spawning season through December). This location was 98

chosen because of the high return rate and due to the relatively high historical catch numbers of 99

barred sand bass. For each group of fish tagged in June, July, and August, we plotted the 100

cumulative number of tag returns over time (days at liberty), as well as the frequency of recaptures 101

over time. Each trend line was examined for the point in time after which the rate of tag returns 102

7

decreased; this point was assumed to represent the minimum residence time. We also report the 103

locations and recapture distances of fish that were recaptured away from Huntington Beach Flats 104

during the same peak spawning season. 105

Movement to non-spawning season locations 106

Movement from peak spawning season tagging locations to non-spawning season recapture 107

locations was assumed to be movement from spawning grounds to non-spawning season 108

residences. To estimate the proximity of non-spawning season residences to spawning grounds, 109

we grouped non-spawning season recapture distances for fish tagged during peak spawning season 110

into 5 km bins. We then calculated the average non-spawning season recapture distances from 111

each tagging location to determine whether non-spawning season “migration” distances varied by 112

spawning location. Finally, we tested for a relationship between fish length (TL) and “migration” 113

distance to non-spawning season residences using a Spearman Rho rank test. 114

Movement to spawning locations 115

We examined peak spawning season recaptures of fish tagged in Newport Bay during the 116

non-spawning season to identify if and where Newport Bay residents migrate to spawn. Spawning 117

“migration” distances from Newport Bay to spawning grounds were reported and also tested for a 118

relationship with fish length (TL) using a Spearman Rho rank test. Lastly, we looked for seasonal 119

patterns in site fidelity to Newport Bay by creating a recapture plot of fish tagged in Newport Bay 120

(Nov.-May) for the years 1964 to 1973. 121

Spawning and non-spawning season site fidelity 122

To investigate annual site fidelity of barred sand bass to specific peak spawning season 123

tagging locations (i.e., presumed spawning grounds) we considered fish that were only tagged 124

during peak spawning season and recaptured during subsequent peak spawning seasons. We 125

8

constructed a matrix of the number of fish recaptured by tagging location and recapture location, 126

with tag and recapture locations arranged from N to S. A higher number of recaptures that occur 127

along a series of corresponding tag/recapture locations within the matrix (i.e., where recapture 128

location = tag location) indicates a higher degree of spawning site fidelity than an arrangement of 129

non-corresponding tag/recapture locations or few corresponding tag/recapture locations within the 130

matrix. 131

132

Results 133

Tagging Effort 134

From 1962 to 1976 barred sand bass were tagged from Santa Barbara to San Diego Bay (Table 135

1, Figure 2a). Most of this tagging effort occurred in the 1960s. Thirty-eight percent were tagged 136

at Huntington Beach Flats during peak spawning season (Jun.–Aug.), while 21% were tagged in 137

Newport Bay during the non-spawning season and early spawning season (Nov.–May). Tagging 138

at other locations primarily occurred during peak spawning season. Between 1989 and 1999, 139

barred sand bass were tagged from Santa Barbara to Baja California, including Santa Catalina 140

Island (Table 1, Figure 2b); 20 of these locations had also been visited in the 1960s. In the 1990s, 141

50% of fish were tagged at Huntington Beach Flats (32%) and Horseshoe Kelp, mostly during 142

peak spawning season. Most fish were captured by hook-and-line, while others were captured by 143

trawl (Table 1). Tagging effort (= average number of fish tagged per day and average number of 144

tagging months per year) was similar between the two tagging periods (Table 1), although in the 145

1990s, most of the tags were out within the first four years, versus six years in the 1960s. 146

9

Demographics of tagged barred sand bass 147

Length frequency distributions of tagged fish were similar at locations with the highest 148

numbers of tagged fish (Manhattan Reef, Horseshoe Kelp, Huntington Beach Flats, and Newport 149

Bay, Figure 3). On average, fish tagged in the 1990s were larger than fish tagged in the 1960s 150

(Table 1), although this trend was likely influenced by the larger fish tagged in Ventura and 151

Mexico in the 1990s (Figure 3). The majority of fish tagged in Ventura were captured by trawl, 152

with no difference in capture depth between line-caught and trawl-caught barred sand bass (Table 153

1). There was a weak but significant positive linear relationship between capture depth and TL (r2 154

= 0.14, p = 0.001). 155

Most tagged fish were greater than the size at 100% maturity (~ 270 mm); however, the 156

proportion of legal sized (= 305 mm) barred sand bass was higher in the 1990s than the 1960s 157

(Table 1). In the 1990s, the proportion of tagged adults decreased from 88% in the non-spawning 158

season to 59% in the late spawning season; likewise, the proportion of tagged legal fish decreased 159

from 65% to 38%. This trend was also apparent for fish tagged in the 1960s, but was less 160

pronounced. 161

Recaptures 162

Despite similar tagging effort between the two tagging periods, recaptures were more 163

numerous in the 1960s than the 1990s (Table 1, Figure 4a,b). In the 1960s, locations with the 164

highest recapture rates were Huntington Beach Flats (29%) and Newport Bay (16%; Figure 4a). 165

Return rates at these locations were 13% (n = 231) and 22% (n = 224), respectively. In the 1990s, 166

Huntington Beach Flats again had the highest number of recaptures (49%; Figure 4b); however tag 167

returns at this site were relatively low (6%) compared to the 1960s. The 1990s had fewer long 168

term recaptures (> 90 days at liberty) than the 1960s, with the majority of recaptures (75%) 169

10

occurring within just 63 days at liberty compared with 315 days in the 1960s (Figure 5). Although 170

the maximum days at liberty were similar between the two tagging periods (Table 1), there was a 171

significant difference in the average number of days at liberty between the 1960s and the 1990s 172

(student t-test, t = 7.156, p = 0.000; Table 1). Overall, the majority of recaptures occurred within 173

close proximity (< 10 km) to the site of tagging, although one recapture occurred 92 km S (Los 174

Alamitos to Oceanside). It is not clear how many recaptured fish were released versus how many 175

were kept. 176

Spawning season residency 177

We identified 117 Huntington Beach Flats same-year returns in the 1960s and 55 in the 178

1990s. The rate of returns for fish tagged during peak spawning season decreased after 179

approximately 34 days at liberty, regardless of tagging month (Figure 6). This period accounted 180

for 73, 93, and 80% of June-, July-, and August-tagged fish returns, respectively. Subsequently, 181

return rates of July-, and August-tagged fish both decreased by 88%. In contrast, the decrease in 182

the return rate for June-tagged fish was more gradual (40%). This gradual decrease was attributed 183

to a subsequent increase in the frequency of June-tagged fish returns that occurred between 46 and 184

55 days at liberty (Figure 6); these returns occurred during the month of August. The maximum 185

number of days at liberty was greatest for August- (119 days) and June-tagged fish (77 days), 186

compared to only 56 days for July-tagged fish. 187

Fifteen fish tagged at Huntington Beach Flats during peak spawning season were 188

recaptured at a different location during the same peak spawning season; recapture locations for 189

these migratory fish included Horseshoe Kelp (n = 9), Seal Beach (n = 1), Santa Ana River Jetty (n 190

= 3), Corona Del Mar (n = 1), and Dana Point (n = 1). Most of these migratory fish (13 of 15) 191

were tagged in July. 192

11

Movement to non-spawning season locations 193

Non-spawning season recapture distances were variable among individuals, with some 194

suggesting resident behavior and others demonstrating migratory behavior. There were 59 barred 195

sand bass that were tagged during peak spawning season and recaptured during non-spawning 196

season (1960s, n = 50; 1990s, n = 9). The frequency distribution of recapture distances revealed 197

an interesting trend, with 64% of fish being recaptured within 1 km of the tagging site; the 198

remainder showed a normal distribution around 15 km (Figure 7). In the 1960s, the overall 199

average (±SD) non-spawning season recapture distance was 4 ± 7 km, but fish recaptured greater 200

than 1 km away from the tagging location had an average recapture distance of 13 ± 8 km. In the 201

1990s, eight of nine fish were recaptured greater than 1 km away from the tag site; the average 202

non-spawning season recapture distance was 19 ± 14 km. Fish size (TL) showed a weak, positive 203

correlation with non-spawning season “migration” distance (rs(57) = 0.31, p = 0.02). 204

Carlsbad and Huntington Beach Flats tag locations had the highest number of recaptures 205

occurring during non-spawning season, but fish tagged at Huntington Beach Flats showed higher 206

variability in recapture distances (Table 2). The farthest non-spawning season recapture distances 207

occurred between Ventura and Carbon Canyon (40 km S) and Tijuana, Mexico and La Jolla (35 208

km N). The farthest non-spawning season recapture location from Huntington Beach Flats was the 209

Palos Verdes Peninsula (29 km N). Most non-spawning season recapture locations were located 210

north of peak spawning season tagging locations (Table 2). 211

Movement to spawning locations 212

Fish tagged in a presumed non-spawning season residence (Newport Bay) during non-213

spawning season were primarily recaptured outside of Newport Bay during peak spawning season 214

(Table 4). We identified at least 15 different peak spawning season recapture sites that were 215

12

typically located south of Newport Bay by an average (±SD) distance of 17 ± 15 km (Table 3). No 216

fish were tagged in Newport Bay during non-spawning season in the 1990s; however, a fish tagged 217

in San Diego Bay during non-spawning season was recaptured during peak spawning season off 218

the coast of San Onofre (75 km N). In contrast to the correlation results reported above, no 219

correlation was found between fish size (TL) and “migration” distance from Newport Bay to 220

spawning grounds (rs(71) = 0.23, p = 0.05). 221

Spawning and non-spawning season site fidelity 222

Several fish were tagged during peak spawning season and recaptured during subsequent 223

peak spawning seasons (1960s, n = 162; 1990s, n = 7). Eighty-nine percent of these recaptures 224

occurred after 1 yr at liberty, 8% after 2 yr, and 2% after 3 yr. Of the 169 recaptures, 80% 225

occurred back at the same tagging location. The average recapture distance (±SD) for the 20% 226

that were recaptured elsewhere was 18 ± 16 km. Overall, the recapture matrix plot identified a 227

high degree of breeding site fidelity as indicated by the arrangement of recaptures occurring along 228

corresponding tag/recapture locations (Figure 8). Tagging locations with the highest measure of 229

breeding site fidelity appeared to be Huntington Beach Flats, Ocean Park Venice, San Onofre 230

Power Plant, Carlsbad, and Twintrees; however, these return rates may be biased by differences in 231

fishing and tagging effort across sites. We also identified two fish that were twice recaptured in 232

subsequent peak spawning seasons at the same locations (Twintrees and Huntington Beach Flats, 233

Table 4). 234

Annual trends in Newport Bay recaptures revealed fish were still present and/or had left 235

and returned to Newport Bay during subsequent non-spawning and early spawning seasons (Figure 236

9). Two fish were twice recaptured at Newport Bay over subsequent non-spawning seasons (Table 237

13

4). Fish recaptures outside of Newport Bay typically occurred during the middle of peak spawning 238

season through late spawning season (Figure 9). 239

240

Discussion 241

Spawning season residency 242

The spawning residency of migrant barred sand bass at Huntington Beach Flats appears to 243

be approximately one month. This is within the reported spawning residency for other fishes (4 – 244

80 days; Robichaud et al. 2003; Bendal et al. 2005; Douglas et al. 2009). For some species, 245

spawning residency may vary by year and by sex, but we were unable to test this for barred sand 246

bass. Nevertheless, individual barred sand bass with shorter-term residency (e.g., up to one 247

month) likely represent migratory individuals, while fish demonstrating longer-term residency 248

(e.g., up to four months) likely represent fish whose home ranges are located within or near to 249

Huntington Beach Flats. Fish tagged in June or August (the beginning and end of peak spawning 250

season) demonstrated longer-term residency to the spawning grounds than fish tagged in July, 251

suggesting the peak in spawning migrations occurs in July. This timing is in agreement with the 252

month of highest catch numbers and fishing effort for barred sand bass in the southern California 253

CPFV fishery (CDFG unpublished data). Moreover, the majority of fish that were recaptured 254

away from Huntington Beach Flats during peak spawning season were tagged in July. Finally, the 255

expected peak in immigration/emigration rates and associated fishing pressure was apparent in the 256

frequency of June-tagged returns over time. For example, return rates of June-tagged fish 257

decreased after a month and then subsequently increased again before leveling off. We attribute 258

this initial decrease to a peak in immigration to the spawning grounds occurring some time in July, 259

while the subsequent increase and leveling off in return rates is likely associated with a peak in 260

14

emigration from the spawning grounds, followed by an associated decrease in fishing pressure 261

there. 262

Due to the use of standard tag and recapture methods, we were unable to quantify diel site 263

fidelity at spawning locations. Barred sand bass individuals may use neighboring reefs to “rest” or 264

forage between spawning events or may travel to and from other nearby spawning locations during 265

this period. Actively tracked coral trout, Plectropomus leopardus, an aggregate spawning 266

Epinephelid (= Serranidae in part; Smith and Craig 2007), were shown to make multiple trips 267

between their home ranges and nearby spawning sites (< 1 km) during the spawning season (Zeller 268

1998). Zeller (1998) also reported that in some cases, visits to spawning grounds occurred 269

multiple times during the day or lasted from two to three days. Several large artificial reefs and 270

structures located near Huntington Beach Flats include Bolsa Chica Artificial Reef (2.7 km W), oil 271

platforms Eva (2.7 km NW), Emma (2.3 km NE), and Esther (6.7 km NW), and the Los Angeles 272

Federal Breakwater (11.8 km NW). Data collected by the California Seafloor Mapping Program 273

(CSMP) reveal patches of hard substrate (i.e., smaller patch reefs or structures) also exist within 274

the Huntington Beach Flats area (CSUMB 2009). Increases of barred sand bass densities during 275

peak spawning season have been observed on SCUBA at Platform Eva (Martin 2008) and the Los 276

Angeles Federal Breakwater (Froeschke et al. 2005). The degree to which these reefs/structures 277

may act as reproductive staging areas for migratory individuals, resting/foraging areas during peak 278

spawning season, or even distinct spawning areas, remains unknown. 279

Movement to and from spawning locations 280

Movement to and from spawning locations is perhaps one of the more interesting and yet 281

least known behaviors of barred sand bass. Our results further suggest that some barred sand bass 282

individuals migrate to spawn, while others are year-round residents. Mason (2008) demonstrated 283

15

that a portion of acoustically monitored adult barred sand bass at Santa Catalina Island, CA, 284

exhibited year-round site fidelity to their home ranges, while others were not detected in these 285

areas during spawning season. This type of “polymorphic movement behavior” has been 286

described for other aggregate spawners including coral trout, Nassau grouper, Epinephelus 287

striatus, and snapper, Pagrus auratus (Colin 1992; Zeller 1998; Egli and Babcock 2004). 288

Not every fish migrated to the same location after spawning season. Likewise, barred sand 289

bass that shared a non-spawning season residence (Newport Bay) did not necessarily migrate to the 290

same spawning grounds. Zeller (1998) reported that coral trout with overlapping home ranges did 291

not necessarily make excursions to the same spawning grounds, regardless of spawning ground 292

proximity to home ranges. Spawning migration distance has been strongly linked to body 293

condition, size-at-age/maturity, and fishing pressure in Atlantic cod (Gadus morhua; Jørgensen et 294

al. 2008); fish that migrate longer distances are generally older, bigger fish with higher overall 295

fitness. Although a relationship between fish size (TL) and “migration” distance in this study was 296

inconclusive, we can not rule out bioenergetics as a possible explanation for the observed 297

variability in migration distances, as maturity and fitness were not determined for tagged fish. 298

Moreover, it is impossible to know for certain the accuracy of the reported fish tag and recapture 299

locations or if some fish were early or late spawners/migrators. These caveats, in addition to the 300

possibility that a portion of the adult barred sand bass population simply doesn’t spawn (i.e., 301

practices “skipped spawning”), may explain the variability in reported migration distances 302

(Jørgensen et al. 2005). 303

Spawning and non-spawning season site fidelity 304

Most fish tagged during peak spawning season were recaptured at the same location during 305

subsequent peak spawning seasons. Whether these individuals represent year-round residents or 306

16

repeat migrants, barred sand bass appear to show a high degree of breeding site fidelity. 307

Historically, breeding site fidelity was studied extensively in salmonids, but less so in other fishes. 308

With the advent of more sophisticated tracking methods (e.g. electronic tags), breeding site fidelity 309

in fishes has been increasingly documented (Hunter et al. 2003; Block et al. 2005; Douglas et al. 310

2009). Tradition (or learned behavior) has been shown to play a primary role in spawning site 311

selection over annual reassessment of resources, especially if resources are relatively unchanging 312

from one year to the next (Warner 1988, 1990). Due to annual differences in tagging effort across 313

tagging locations, it was not possible to accurately quantify long-term inter-annual variability in 314

spawning site fidelity by tagging location. Recaptures that occurred at different locations in 315

subsequent years may reflect individual variability in the timing of spawning-related movements, 316

movement among aggregation sites, or a certain degree of annual reassessment. 317

We also identified individuals that demonstrated site fidelity to a non-spawning season 318

tagging location (Newport Bay). Bays are noted as habitat for young-of-the year (YOY) and 319

juvenile barred sand bass (Feder et al. 1974; Valle et al. 1999; Allen et al. 2002), while adults 320

prefer sand/rock ecotone habitat to 30 m depth (Feder et al. 1974; Johnson et al. 1990; Mason 321

2008). In San Diego Bay, YOY barred sand bass were shown to develop there throughout their 322

life history and adults utilized the bay habitat throughout the year (Pondella et al. 2006). In this 323

study, a fish that was tagged and recaptured during non-spawning season in Newport Bay may 324

represent a fish that remained there year-round or one that migrated to spawn and returned in the 325

winter. Our data indicate that a portion of adult barred sand bass in Newport Bay migrate to 326

locations outside of the bay during spawning season. This identifies Newport Bay as an important 327

non-spawning season residence for barred sand bass, as well as an important source of migrants to 328

various spawning locations outside of the bay. Although it is unknown for certain whether the 329

17

migrant recaptures in this study would have returned to Newport Bay following peak spawning 330

season, the overall pattern in recaptures is highly suggestive. 331

Recapture rates 332

The most striking difference between the two tagging periods is the recapture rate: 17% in 333

the 1960s compared to 4% in the 1990s. A typical range of recapture rates using standard tag and 334

recapture methods is 3 to 10% (Lowe and Bray 2006). Given that tagging effort and numbers of 335

tagged fish did not dramatically differ between these two time periods, recapture rates may have 336

been influenced by changes in barred sand bass availability, in addition to advances in fishing 337

technology (e.g., fish finders, GPS), the willingness of fishers to report tag returns, and an 338

increased knowledge of spawning aggregation sites. In general, high recapture rates in open 339

systems reflect relatively lower population sizes due to the higher probability of encountering the 340

same fish at a later date. This may explain the higher number of long-term recaptures in the 1960s 341

dataset. CPFV barred sand bass catch values were nearly four times greater in the 1990s than in 342

the 1960s despite only a doubling of fishing effort (CDFG unpublished data). Historical Fish and 343

Game publications report that barred sand bass were scarce during the 1950s (a cold water period) 344

and occurred more frequently along the coast “in and subsequent to periods of warmer waters” 345

(Young 1969; Feder et al. 1974). Kelp bass and barred sand bass larvae densities were also lower 346

during the cool regime (1950s – 1970s) and higher in the warm regime (1980s – 1990s), especially 347

during the 1980s (Moser et al. 2001). While it appears that barred sand bass populations may have 348

increased in the 1990s relative to the 1960s, barred sand bass stock-recruitment relationships and 349

the effects on these relationships by natural and anthropogenic influences remain unknown. 350

18

Management implications 351

Currently, a sustainable level of harvest for barred sand bass is unknown because no 352

biomass estimates exist. Due to concerns over barred sand bass catch declines in recent years and 353

the uncertainty in the status of the population, barred sand bass may require further fishery 354

protection beyond current minimum size limits and bag limits. Information gathered on the 355

spawning-related movements of barred sand bass may be useful for identifying further protection 356

measures. For instance, an aggregate spawner that demonstrates multiple spawning events over a 357

relatively long spawning residency period would likely benefit from a series of temporary fishing 358

closures during the peak in fishing effort. Non-spawning residences that are shown to supply 359

migrant spawners to spawning aggregation sites may be important sites to consider for seasonal 360

protection of barred sand bass. Further consideration of barred sand bass movement patterns and 361

life history traits, in addition to important feasibility concerns, may help to define additional 362

alternatives to protect barred sand bass until a harvest guideline can be developed. 363

364

Acknowledgements 365

CDFG lead investigators for the barred sand bass tagging studies in the 1960s and 1990s 366

were P. “Bud” Young and J.R. Raymond Ally, respectively. Over the two tagging periods, CDFG 367

tagging efforts were augmented by [and listed in no particular order] R. Izor (Izorline 368

International), Orange County Marine Institute, County Sanitation Districts of Orange County, Los 369

Angeles Rod and Reel Foundation, and many other individual volunteers. We thank L.G. Allen 370

and T. Mason for their critical review of the drafts of this manuscript. Funding was supported in 371

part by the Los Angeles County Fish and Game Commission and the Federal Aid in Sportfish 372

Restoration Act (also known as the Dingell-Johnson Act; Grant #F-50-R-20). 373

19

Literature Cited 374

Aburto-Oropeza, O., B. Erisman, V. Valdez-Ornelas, and G. Danemann. 2008. Commercially 375

Important Serranid Fishes from the Gulf of California: Ecology, Fisheries, and Conservation. 376

Ciencia y Conservación 2008:1-23. 377

Allen, L.G., and T.E. Hovey. 2001. Barred sand bass. Pp. 224-225 in California's Living Marine 378

Resources: A Status Report (W.S. Leet, C.M. Dewees, R. Klingbeil & E.J. Larson, eds.), 379

Calif. Fish Game Resources Agency, 592 pp. 380

Allen, L.G., A.M. Findlay, and C.M. Phalen. 2002. Structure and standing stock of the fish 381

assemblages of San Diego Bay, California from 1994 to 1999. Bull. So. Cal. Acad. Sci., 382

1012:49-85. 383

Ally, J.R.R, D.S. Ono, R.B. Read, M.D. Harris, and M. Wallace. 1990. Southern California 384

Partyboat Sport Fish Survey: Procedures Manual. Calif. Fish Game Resources Agency, 130 385

pp. 386

Bendall, B., A. Moore, and V. Quayle. 2005. The post-spawning movements of migratory brown 387

trout, Salmo trutta L. J. Fish Biol., 67:809-822. 388

Block, B.A., S.L.H. Teo, A. Walli, A. Boustany, M.J.W. Stokesbury, C.J. Farwell, K.C. Weng, H. 389

Dewar, And T.D. Williams. 2005. Electronic tagging and population structure of Atlantic 390

bluefin tuna. Nature, 434:1121-1127. 391

Collyer, R.D. 1949. Rockbass. Pp 113-115 in The commercial fish catch of California for the 392

year 1947 with an historical review 1916-1947. Calif. Fish Game, Fish Bull., 74. 393

[CSUMB] California State University, Monterey Bay. 2009. 2008 CSMP Surveys: San Diego to 394

Point Conception. (1 December, 2009; http://seafloor.csumb.edu). 395

20

Domeier, M.L., and P.L. Colin. 1997. Tropical reef fish spawning aggregations: defined and 396

reviewed. Bull. Mar. Sci., 60:698-726. 397

Douglas, S.G., G. Chaput, J. Hayward, and J. Sheasgreen. 2009. Prespawning, spawning, and 398

postspawning behavior of striped bass in the Miramichi River. Trans. Am. Fish. Soc., 399

138:121-134. 400

Egli, D.P. and R.C. Babcock. 2004. Ultrasonic tracking reveals multiple behavioural modes of 401

snapper (Pagrus auratus) in a temperate no-take marine reserve. ICES J. Mar. Sci., 402

61:1137-1143. 403

Erisman, B.E. and L.G. Allen. 2006. Reproductive behavior of a temperate serranid fish, 404

Paralabrax clathratus, from Santa Catalina Island, California, U.S.A. J. Fish Biol., 68:157-405

184. 406

Feder, H.M., C.H. Turner, and C. Limbaugh. 1974. Observations on fishes associated with kelp 407

beds in southern California. Calif. Fish Game Fish Bull., 160:31-32. 408

Froeschke, J.T., L.G. Allen, and D.J. Pondella. 2005. The reef fish assemblage of the outer Los 409

Angeles Federal Breakwater, 2002-2003. Bull. So. Cal. Acad. Sci., 104:63-74. 410

Hovey, T.E. and L.G. Allen. 2000. Reproductive patterns of six populations of the spotted sand 411

bass, Paralabrax maculatofasciatus, from Southern and Baja California. Copeia, 2000:459-412

468. 413

Hunter, E., J.D. Metcalfe, and J.D. Reynolds. 2003. Migration route and spawning area fidelity 414

by North Sea plaice. Proc. R. Soc. Lond. B, 270:2097-2103. 415

Johnson, T.D., A.M. Barnett, E.E. DeMartini, L.L. Craft, R.F. Ambrose, and L.J. Purcell. 1994. 416

Fish production and habitat utilization on a southern California artificial reef. Bull. Mar. Sci., 417

55:709-723. 418

21

Jørgensen, C., B. Ernande, Ø. Fiksen, and U. Dieckmann. 2006. The logic of skipped spawning in 419

fish. Can. J. Fish. Aquat. Sci., 63:200-211. 420

Jørgensen, C., E.S. Dunlop, A.F. Opdal, and Ø. Fiksen. 2008. The evolution of spawning 421

migrations: state dependence and fishing-induced changes. Ecology, 89:3436-3448. 422

Love, M.S., A. Brooks, and J.R.R. Ally. 1996a. An analysis of commercial passenger fishing 423

vessel fisheries for kelp bass and barred sand bass in the southern California Bight. Calif. Fish 424

Game, 82:105-121. 425

Love, M.S., A. Brooks, D. Busatto, J. Stephens, and P. Gregory. 1996b. Aspects of the life 426

histories of the kelp bass, Paralabrax clathratus, and barred sand bass, P. nebulifer, from the 427

southern California Bight. U.S. Fish. Bull., 94:472-481. 428

Lowe, C.G., and R.N. Bray. 2006. Movement and activity patterns. Pp. 524-553 in The Ecology 429

of California Marine Fishes (L.G. Allen, M.H. Horn, and D.J. Pondella, eds.), University of 430

California Press, 670 pp. 431

Martin, C.J.B. 2009. San Pedro Shelf platform fish assemblages and relations to habitat quality. 432

Master’s Thesis. California State University Long Beach. 433

Mason, T. 2008. Home range size, habitat use, and the effects of habitat breaks on the movements 434

of temperate reef gamefishes in a southern California marine protected area. Master’s Thesis. 435

California State University Long Beach. 52 pp. 436

Moser, H.G., R.L. Charter, P.E. Smith, D.A. Ambrose, W. Watson, S.R. Charter, and E.M. 437

Sandknop. 2001. Distributional atlas of fish larvae and eggs in the Southern California Bight 438

region: 1951-1998. Calif. Coop. Oceanic Fish. Invest. Atlas, 34:46. 439

[PSMFC] Pacific States Marine Fisheries Commission. 2010a. Recreational Fisheries 440

Information Network (RecFIN) Estimated Total Catch with Releases (A+B1+B2) in 441

22

Thousands of Fish Caught By Marine Recreational Anglers By Species and Year For All 442

Modes of Fishing in All Marine Areas in Southern California where Common Name 443

Contains Barred Sand Bass. (20 January, 2010; www.recfin.org) 444

PSMFC. 2010b. RecFIN Estimated Total Harvested Dead Catch (A+B1) in Thousands of Fish 445

Caught by Marine Recreational Anglers Fishing for All Possible Species by Month and State 446

Fishing Mode for All Modes Of Fishing in All Marine Areas in Southern California from 447

January 2004 - December 2008 where Common Name Contains Barred Sand Bass. (1 448

February, 2010; www.recfin.org) 449

Pondella, D.J., L.G. Allen, M.T. Craig, and B. Gintert, B. 2006. Evaluation of eelgrass mitigation 450

and fishery enhancement structures in San Diego Bay, California. Bull. Mar. Sci., 78:115-451

131. 452

Robichaud, D. and G.A. Rose. 2003. Sex differences in cod residency on a spawning ground. 453

Fish. Res., 60:33-43. 454

Sadovy, Y. and M.L. Domeier. 2005. Are aggregation-fisheries sustainable? Reef fish fisheries as 455

a case study. Coral Reefs, 24:254-262. 456

Sadovy, Y. and M.L. Domeier. 2005. Perplexing problems of sexual patterns in the fish genus 457

Paralabrax. J. Zool. Lond., 267:121-133. 458

Sala, Enric, O. Aburto-Oropeza, G. Paredes, and G. Thompson. 2003. Spawning aggregations 459

and reproductive behavior of reef fishes in the Gulf of California. Bull. Mar. Sci.,72:103-460

121. 461

Smith, W. and M.T. Craig. 2007. Casting the Percomorph net widely: the importance of broad 462

taxonomic sampling in the search for the placement of Serranid and Percid fishes. Copeia, 463

1:35-55. 464

23

Turner, C.H., E.E. Ebert, and R.R. Given. 1969. Man-made reef ecology. Calif. Fish Game Fish. 465

Bull., 146:176-177. 466

Valle, C.F., J.W. O’Brien, and K.B. Wiese. 1999. Differential habitat use by California halibut, 467

Paralichthys californicus, barred sand bass, Paralabrax nebulifer, and other juvenile fishes in 468

Alamitos Bay. U.S. Fish. Bull., 97:646-660. 469

Warner, R.R. 1988. Traditionality of mating-site preferences in a coral reef fish. Nature, 470

335:719-721. 471

Warner, R.R. 1990. Resource assessment versus tradition in mating-site determination. Am. Nat., 472

135:205-217. 473

Young, P.H. 1969. The California partyboat fishery 1947-1967. Calif. Fish Game Fish. Bull., 474

145:1-91. 475

Zeller, D.C. 1998. Spawning aggregations: patterns of movement of the coral trout Plectropomus 476

leopardus as determined by ultrasonic telemetry. Mar. Ecol. Prog. Ser., 162:253-263. 477

24

Table 1. Tag and recapture summary statistics for barred sand bass tagged in southern California, 478

historical California Department of Fish and Game tagging project (1960s and 1990s). 479

Tag and recapture results 1960s 1990sTagged fish 4,687 3,947Tagging effort days 174 153 fish/day 27 ± 32 26 ± 58 Avg (±SD) (mo/yr) 4 ± 3 6 ± 3Capture method hook-and-line 100% 74% bottom trawl -- 26%Avg (±SD) capture depth (m) overall -- 15 ± 17 hook-and-line -- 22 ± 7 bottom trawl -- 25 ± 12Avg (±SD) TL (mm) 306 ± 38 337 ± 72 % mature 89% 93% % legal size 41% 70% Recaptures 801 171Return rate overall 17% 4% hook-and-line 17% 5% bottom trawl -- 3%Avg (±SD) recapture depth (m) -- 23 ± 9Avg (±SD) TL (mm) 326 ± 43 343 ± 46 % mature 96% 98% % legal size 68% 86%Days at liberty Avg (±SD) 200 ± 197 90 ± 187 Max 1,211 1,258Recapture distance (km) Avg (±SD) - All fish 6 ± 12 7 ± 9 Avg (±SD) - Only movers 18 ± 15 10 ± 9 Max 92 76

480

25

Table 2. Recapture distances (km) of barred sand bass tagged during peak spawning season 481

(Jun-Aug) and recaptured during non-spawning season (Nov-Mar), historical California 482

Department of Fish and Game tagging project (1960s and 1990s). Dir. = direction of recapture 483

location from tagging location. 484

485

26

Table. 3. Recapture distances (km) of barred sand bass tagged in Newport Bay during the non-486

spawning season (Nov-Mar) and recaptured during peak spawning season (Jun-Aug), historical 487

California Department of Fish and Game tagging project (1960s). Dir. = direction of recapture 488

location from tagging location. 489

490

Recapture distance (km)

Peak Spawning Season Recapture Location N Avg Stdev Dir.

Horseshoe Kelp 1 24.1 -- W

Huntington Beach Flats 13 17.6 2.2 W

Santa Ana River Jetty 3 8.0 0.0 W

Newport Harbor 19 0.5 1.5 --

Corona Del Mar 2 2.4 1.1 S

Crystal Cove 1 1.6 -- S

North Laguna Beach 3 10.2 1.9 S

South Laguna Beach 1 12.9 -- S

Salt Creek 2 18.5 1.1 S

Dana Point 6 20.4 1.3 S

Capistrano Beach 1 24.1 -- S

San Mateo Point 4 31.4 1.6 S

San Clemente 1 29.0 -- S

Middle Kelp 1 24.1 -- S

San Onofre Power Plant 5 32.8 0.9 S

Barn Kelp 8 42.8 1.5 S

Oceanside 1 51.5 -- S

27

Table 4. Tag and recapture dates and locations of barred sand bass recaptured on two separate occasions, historical California 491

Department of Fish and Game tagging project (1960s). 492

493

494

* Asterisks denote fish that were released in Newport Harbor following weigh-in at a fishing tournament.495

28

List of Figures 496

497

Figure 1. Map of barred sand bass tagging locations in southern California, historical California 498

Department of Fish and Game tagging project (1960s and 1990s). 499

500

Figure 2. Numbers of barred sand bass tagged by the California Department of Fish and Game 501

in southern California during the a) 1960s and b) 1990s. 502

503

Figure 3. Length-frequency distributions of barred sand bass by tagging location, historical 504

California Department of Fish and Game tagging project (1960s and 1990s). 1960s and 1990s 505

sites are represented by grey and white bars, respectively. Only locations where at least 100 506

tagged individuals are listed. The vertical lines represent minimum size of legal take (305 mm). 507

508

Figure 4. Numbers of tagged barred sand bass recaptured during the a) 1960s and b) 1990s, 509

historical California Department of Fish and Game tagging project (1960s and 1990s). 510

511

Figure 5. Numbers of barred sand bass recaptured by days since tagged for fish tagged in the a) 512

1960s and b) 1990s, historical California Department of Fish and Game tagging project (1960s 513

and 1990s). 514

515

Figure 6. Cumulative number of barred sand bass recaptures (dots) and frequency of recaptures 516

(bars) by days since tagged for fish tagged during each month of peak spawning season (June, 517

29

July, August) at Huntington Beach Flats, CA and recaptured within the same year at the same 518

location, historical California Department of Fish and Game tagging project (1960s and 1990s). 519

520

Figure 7. Recapture distances for barred sand bass tagged during peak spawning season (Jun-521

Aug) and recaptured during non-spawning season (Nov-Mar), historical California Department 522

of Fish and Game tagging project (1960s and 1990s). 1960s = dark bars, 1990s = grey bars. 523

524

Figure 8. Recapture matrix plot of barred sand bass tagged during peak spawning season and 525

recaptured in subsequent peak spawning seasons, historical California Department of Fish and 526

Game tagging project (1960s and 1990s). Shaded, darker boxes occurring along the diagonal 527

line indicate a higher degree of breeding site fidelity. 528

529

Figure 9. Recapture plot of barred sand bass tagged in Newport Bay, California during non-530

spawning season (Nov-Mar, shaded areas), historical California Department of Fish and Game 531

tagging project (1960s). Recaptures occurring within 90 d of tagging were excluded. Recapture 532

locations are arranged north to south (top to bottom).533

30

534

Figure 1. 535

31

536

537 Figure 2. 538

32

539 Figure 3. 540

33

541

542 Figure 4 543

34

544 Figure 5. 545

35

546 547 Figure 6. 548

36

549 Figure 7. 550

37

551 Figure 8. 552

38

553 Figure 9. 554