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Jarvisetal bsb movements_01_mar10
1. 1 SPAWNING-RELATED MOVEMENTS OF BARRED SAND BASS, PARALABRAX
2 NEBULIFER, IN SOUTHERN CALIFORNIA: INTERPRETATIONS FROM TWO DECADES
3 OF HISTORICAL TAG AND RECAPTURE DATA.
4 E.T. Jarvis, C. Linardich, and C.F. Valle. California Department of Fish and Game, Marine
5 Region, Los Alamitos, CA, 90720.
2. 6 Abstract
7 During the 1960s and 1990s, the California Department of Fish and Game tagged 8,634
8 barred sand bass in southern California. Tag and recapture data suggest migrant individuals reside
9 at spawning grounds for approximately one month. Although 64% of non-spawning season
10 recaptures occurred within 1 km of the spawning location, the average (± SD) recapture distance of
11 migrant fish was 13 ± 8 km. Fish size (TL) was weakly correlated with “migration” distance from
12 spawning locations (rs(57) = 0.31, p = 0.02). Spatial and temporal trends in recaptures suggest a
13 high degree of spawning and non-spawning season site fidelity.
2
3. 14 Introduction
15 Barred sand bass, Paralabrax nebulifer, continues to be one of the most sought-after sport fish
16 in southern California. In the early 1900s, barred sand bass were landed in both the commercial
17 and recreational fisheries; however, due to limited demand in the commercial fishery and scarcity
18 of the resource during the 1950s, commercial take was banned in 1953 and a 12 in (30.5 cm)
19 minimum size limit was implemented in 1959 (Collyer 1949; Young 1969). Since the 1960s,
20 barred sand bass have ranked among the top 10 sport fish caught by commercial passenger fishing
21 vessels (CPFVs = party boats) in southern California, with total annual catches averaging nearly
22 two million fish per year between 1980 and 2003* (Allen and Hovey 2001; PSMFC 2010a).
23 Although no commercial fishery currently exists for barred sand bass in California, “heavy annual
24 landings” of barred sand bass have been reported in the commercial fisheries of Baja California,
25 Mexico, and the Gulf of Mexico in recent years (Aburto-Oropeza et al. 2008).
26 In southern California, fishing effort for barred sand bass by both party and private boaters is
27 highest during peak spawning season (June to August). In recent years, approximately 77% of the
28 June, July, and August barred sand bass harvest was taken by party/charter boats (PSMFC 2010b).
29 Well-known spawning aggregation sites include the Ventura Flats, inner Santa Monica Bay,
30 Huntington Beach Flats, San Onofre, and Silver Strand in San Diego. According to California
31 Department of Fish and Game (CDFG) CPFV logbook data, Huntington Beach Flats ranks the
32 highest in numbers of barred sand bass caught in the past 40 years (CDFG unpublished data).
33 From 1961 to 2008, barred sand bass CPFV catch-per-unit-effort (CPUE = barred sand bass
34 kept/angler*hr) increased to a high in 2000, and has recently decreased to the lowest CPUE since
*
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.
3
4. 35 1985 (CDFG unpublished data), causing concern regarding the vulnerability of the population to
36 future harvest impacts.
37 Fish species that are targeted during their spawning aggregations are especially susceptible to
38 overexploitation because harvest effects may not be immediately evident (Sadovy and Domeier
39 2005). Furthermore, overexploitation of an aggregate spawner can result in the complete absence
40 of spawning aggregation formations at historic sites (Domeier and Colin 1997; Sadovy and
41 Domeier 2005). Consequences of this occurring in the barred sand bass fishery might include
42 negative ecological impacts, as well as severe economic impacts to the sport fishing industry in
43 southern California.
44 Barred sand bass is one of three temperate serranine fishes (sea basses), including kelp
45 bass, Paralabrax clathratus, and spotted sand bass, Paralabrax maculatofasciatus, that participate
46 to varying degrees in aggregate spawning behavior (Hovey and Allen 2000; Erisman and Allen
47 2006; Miller and Allen 2006). Barred sand bass breeding aggregations are reported to occur over
48 sand flats in depths of 20 to 40 m (Turner 1969; Feder et al. 1974; Love et al. 1996), and based on
49 the exceptionally high landings of barred sand bass during the summer, it is possible that these
50 aggregations consist of thousands of fish. However, underwater video documentation of these
51 aggregations has never been reported. Following peak spawning, considerably fewer barred sand
52 bass are caught over the sand flats and catches typically resume inshore in bays or near low relief
53 natural or artificial reefs, but not in such high numbers (Love et al. 1996). These catch trends may
54 suggest transient spawning aggregation behavior, in which large aggregations form at specific,
55 predictable locations at higher than average densities for a period of several weeks to months
56 (Domeier and Colin 1997). Transient spawning aggregations are characterized by having
57 individuals that migrate relatively longer distances to breed than “resident” aggregate spawners,
4
5. 58 but it is still unclear to what degree this may occur with barred sand bass. Knowledge of the
59 origins and destinations of barred sand bass spawning migrations, and understanding whether all or
60 some of the adult population participates in spawning aggregations will be important for effective
61 management of this species.
62 In the 1960s and 1990s, biologists with the California Department of Fish and Game (CDFG)
63 tagged nearly 9,000 barred sand bass in southern California. The recapture information from these
64 two time periods enables us to document the historical spawning-related movements of barred
65 sand bass for the first time. Specifically, our objectives of this study are to examine these
66 historical data for trends in 1) residency at spawning locations, 2) movement to and from spawning
67 locations, and 3) breeding site fidelity.
68
69 Methods
70 Tagging Effort
71 During the 1960s and 1990s, barred sand bass were tagged along the coast of southern
72 California and at one location in Baja California, Mexico (Figure 1). Tagging locations included
73 sand flats, reefs, and bay habitat. During both tagging periods, fish were captured by hook-and-
74 line, measured to the nearest mm total length (TL), externally tagged with spaghetti or T-bar tags,
75 and released. In the 1990s, fish were also captured by bottom trawl, and upon release, tagged fish
76 suffering from barotrauma were recompressed to depth using milk crates. Loran or GPS
77 coordinates of the tagging sites were recorded; otherwise, a site name or geographic landmark was
78 provided. In addition, depth (m) and release condition were recorded for some but not all fish.
79 Rewards for recaptures of tagged fish were offered during both tagging periods. Recapture
5
6. 80 information included date, location, TL (mm), and tag ID number. In the 1990s, recapture depth
81 (m) and Loran or GPS coordinates were also provided when available.
82 Analysis
83 All historical barred sand bass tag and recapture data were archived into a relational
84 database for analysis. To standardize tagging effort across the two tagging periods, locations for
85 all records were assigned a fishing site code based on historical southern California CPFV sport
86 fish surveys (Ally et al. 1990). Spawning season codes were also assigned to each tag and
87 recapture record based on capture month (Nov.–Mar. = non-spawning season, Apr.–May = early
88 spawning season, Jun.–Aug. = peak spawning season, Sept.–Oct. = late spawning season). Days at
89 liberty, recapture distance (estimated or actual km), and general direction of movement were
90 calculated and incorporated into the database. Recapture distances were measured as linear
91 distances between approximate or actual tagging and recapture locations. In this paper we report
92 recapture rates and return rates. Recapture rates refer to the number of fish recaptured at a given
93 site divided by the total number fish recaptures. Return rates refer to the number of fish recaptured
94 at a given site divided by the total number of fish tagged at that site.
95 Spawning season residency
96 To investigate the residence time of individuals at spawning locations, we selected fish
97 tagged at Huntington Beach Flats during peak spawning season and recaptured at Huntington
98 Beach Flats within the same year (peak spawning season through December). This location was
99 chosen because of the high return rate and due to the relatively high historical catch numbers of
100 barred sand bass. For each group of fish tagged in June, July, and August, we plotted the
101 cumulative number of tag returns over time (days at liberty), as well as the frequency of recaptures
102 over time. Each trend line was examined for the point in time after which the rate of tag returns
6
7. 103 decreased; this point was assumed to represent the minimum residence time. We also report the
104 locations and recapture distances of fish that were recaptured away from Huntington Beach Flats
105 during the same peak spawning season.
106 Movement to non-spawning season locations
107 Movement from peak spawning season tagging locations to non-spawning season recapture
108 locations was assumed to be movement from spawning grounds to non-spawning season
109 residences. To estimate the proximity of non-spawning season residences to spawning grounds,
110 we grouped non-spawning season recapture distances for fish tagged during peak spawning season
111 into 5 km bins. We then calculated the average non-spawning season recapture distances from
112 each tagging location to determine whether non-spawning season “migration” distances varied by
113 spawning location. Finally, we tested for a relationship between fish length (TL) and “migration”
114 distance to non-spawning season residences using a Spearman Rho rank test.
115 Movement to spawning locations
116 We examined peak spawning season recaptures of fish tagged in Newport Bay during the
117 non-spawning season to identify if and where Newport Bay residents migrate to spawn. Spawning
118 “migration” distances from Newport Bay to spawning grounds were reported and also tested for a
119 relationship with fish length (TL) using a Spearman Rho rank test. Lastly, we looked for seasonal
120 patterns in site fidelity to Newport Bay by creating a recapture plot of fish tagged in Newport Bay
121 (Nov.-May) for the years 1964 to 1973.
122 Spawning and non-spawning season site fidelity
123 To investigate annual site fidelity of barred sand bass to specific peak spawning season
124 tagging locations (i.e., presumed spawning grounds) we considered fish that were only tagged
125 during peak spawning season and recaptured during subsequent peak spawning seasons. We
7
8. 126 constructed a matrix of the number of fish recaptured by tagging location and recapture location,
127 with tag and recapture locations arranged from N to S. A higher number of recaptures that occur
128 along a series of corresponding tag/recapture locations within the matrix (i.e., where recapture
129 location = tag location) indicates a higher degree of spawning site fidelity than an arrangement of
130 non-corresponding tag/recapture locations or few corresponding tag/recapture locations within the
131 matrix.
132
133 Results
134 Tagging Effort
135 From 1962 to 1976 barred sand bass were tagged from Santa Barbara to San Diego Bay (Table
136 1, Figure 2a). Most of this tagging effort occurred in the 1960s. Thirty-eight percent were tagged
137 at Huntington Beach Flats during peak spawning season (Jun.–Aug.), while 21% were tagged in
138 Newport Bay during the non-spawning season and early spawning season (Nov.–May). Tagging
139 at other locations primarily occurred during peak spawning season. Between 1989 and 1999,
140 barred sand bass were tagged from Santa Barbara to Baja California, including Santa Catalina
141 Island (Table 1, Figure 2b); 20 of these locations had also been visited in the 1960s. In the 1990s,
142 50% of fish were tagged at Huntington Beach Flats (32%) and Horseshoe Kelp, mostly during
143 peak spawning season. Most fish were captured by hook-and-line, while others were captured by
144 trawl (Table 1). Tagging effort (= average number of fish tagged per day and average number of
145 tagging months per year) was similar between the two tagging periods (Table 1), although in the
146 1990s, most of the tags were out within the first four years, versus six years in the 1960s.
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9. 147 Demographics of tagged barred sand bass
148 Length frequency distributions of tagged fish were similar at locations with the highest
149 numbers of tagged fish (Manhattan Reef, Horseshoe Kelp, Huntington Beach Flats, and Newport
150 Bay, Figure 3). On average, fish tagged in the 1990s were larger than fish tagged in the 1960s
151 (Table 1), although this trend was likely influenced by the larger fish tagged in Ventura and
152 Mexico in the 1990s (Figure 3). The majority of fish tagged in Ventura were captured by trawl,
153 with no difference in capture depth between line-caught and trawl-caught barred sand bass (Table
154 1). There was a weak but significant positive linear relationship between capture depth and TL (r2
155 = 0.14, p = 0.001).
156 Most tagged fish were greater than the size at 100% maturity (~ 270 mm); however, the
157 proportion of legal sized (= 305 mm) barred sand bass was higher in the 1990s than the 1960s
158 (Table 1). In the 1990s, the proportion of tagged adults decreased from 88% in the non-spawning
159 season to 59% in the late spawning season; likewise, the proportion of tagged legal fish decreased
160 from 65% to 38%. This trend was also apparent for fish tagged in the 1960s, but was less
161 pronounced.
162 Recaptures
163 Despite similar tagging effort between the two tagging periods, recaptures were more
164 numerous in the 1960s than the 1990s (Table 1, Figure 4a,b). In the 1960s, locations with the
165 highest recapture rates were Huntington Beach Flats (29%) and Newport Bay (16%; Figure 4a).
166 Return rates at these locations were 13% (n = 231) and 22% (n = 224), respectively. In the 1990s,
167 Huntington Beach Flats again had the highest number of recaptures (49%; Figure 4b); however tag
168 returns at this site were relatively low (6%) compared to the 1960s. The 1990s had fewer long
169 term recaptures (> 90 days at liberty) than the 1960s, with the majority of recaptures (75%)
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10. 170 occurring within just 63 days at liberty compared with 315 days in the 1960s (Figure 5). Although
171 the maximum days at liberty were similar between the two tagging periods (Table 1), there was a
172 significant difference in the average number of days at liberty between the 1960s and the 1990s
173 (student t-test, t = 7.156, p = 0.000; Table 1). Overall, the majority of recaptures occurred within
174 close proximity (< 10 km) to the site of tagging, although one recapture occurred 92 km S (Los
175 Alamitos to Oceanside). It is not clear how many recaptured fish were released versus how many
176 were kept.
177 Spawning season residency
178 We identified 117 Huntington Beach Flats same-year returns in the 1960s and 55 in the
179 1990s. The rate of returns for fish tagged during peak spawning season decreased after
180 approximately 34 days at liberty, regardless of tagging month (Figure 6). This period accounted
181 for 73, 93, and 80% of June-, July-, and August-tagged fish returns, respectively. Subsequently,
182 return rates of July-, and August-tagged fish both decreased by 88%. In contrast, the decrease in
183 the return rate for June-tagged fish was more gradual (40%). This gradual decrease was attributed
184 to a subsequent increase in the frequency of June-tagged fish returns that occurred between 46 and
185 55 days at liberty (Figure 6); these returns occurred during the month of August. The maximum
186 number of days at liberty was greatest for August- (119 days) and June-tagged fish (77 days),
187 compared to only 56 days for July-tagged fish.
188 Fifteen fish tagged at Huntington Beach Flats during peak spawning season were
189 recaptured at a different location during the same peak spawning season; recapture locations for
190 these migratory fish included Horseshoe Kelp (n = 9), Seal Beach (n = 1), Santa Ana River Jetty (n
191 = 3), Corona Del Mar (n = 1), and Dana Point (n = 1). Most of these migratory fish (13 of 15)
192 were tagged in July.
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11. 193 Movement to non-spawning season locations
194 Non-spawning season recapture distances were variable among individuals, with some
195 suggesting resident behavior and others demonstrating migratory behavior. There were 59 barred
196 sand bass that were tagged during peak spawning season and recaptured during non-spawning
197 season (1960s, n = 50; 1990s, n = 9). The frequency distribution of recapture distances revealed
198 an interesting trend, with 64% of fish being recaptured within 1 km of the tagging site; the
199 remainder showed a normal distribution around 15 km (Figure 7). In the 1960s, the overall
200 average (±SD) non-spawning season recapture distance was 4 ± 7 km, but fish recaptured greater
201 than 1 km away from the tagging location had an average recapture distance of 13 ± 8 km. In the
202 1990s, eight of nine fish were recaptured greater than 1 km away from the tag site; the average
203 non-spawning season recapture distance was 19 ± 14 km. Fish size (TL) showed a weak, positive
204 correlation with non-spawning season “migration” distance (rs(57) = 0.31, p = 0.02).
205 Carlsbad and Huntington Beach Flats tag locations had the highest number of recaptures
206 occurring during non-spawning season, but fish tagged at Huntington Beach Flats showed higher
207 variability in recapture distances (Table 2). The farthest non-spawning season recapture distances
208 occurred between Ventura and Carbon Canyon (40 km S) and Tijuana, Mexico and La Jolla (35
209 km N). The farthest non-spawning season recapture location from Huntington Beach Flats was the
210 Palos Verdes Peninsula (29 km N). Most non-spawning season recapture locations were located
211 north of peak spawning season tagging locations (Table 2).
212 Movement to spawning locations
213 Fish tagged in a presumed non-spawning season residence (Newport Bay) during non-
214 spawning season were primarily recaptured outside of Newport Bay during peak spawning season
215 (Table 4). We identified at least 15 different peak spawning season recapture sites that were
11
12. 216 typically located south of Newport Bay by an average (±SD) distance of 17 ± 15 km (Table 3). No
217 fish were tagged in Newport Bay during non-spawning season in the 1990s; however, a fish tagged
218 in San Diego Bay during non-spawning season was recaptured during peak spawning season off
219 the coast of San Onofre (75 km N). In contrast to the correlation results reported above, no
220 correlation was found between fish size (TL) and “migration” distance from Newport Bay to
221 spawning grounds (rs(71) = 0.23, p = 0.05).
222 Spawning and non-spawning season site fidelity
223 Several fish were tagged during peak spawning season and recaptured during subsequent
224 peak spawning seasons (1960s, n = 162; 1990s, n = 7). Eighty-nine percent of these recaptures
225 occurred after 1 yr at liberty, 8% after 2 yr, and 2% after 3 yr. Of the 169 recaptures, 80%
226 occurred back at the same tagging location. The average recapture distance (±SD) for the 20%
227 that were recaptured elsewhere was 18 ± 16 km. Overall, the recapture matrix plot identified a
228 high degree of breeding site fidelity as indicated by the arrangement of recaptures occurring along
229 corresponding tag/recapture locations (Figure 8). Tagging locations with the highest measure of
230 breeding site fidelity appeared to be Huntington Beach Flats, Ocean Park Venice, San Onofre
231 Power Plant, Carlsbad, and Twintrees; however, these return rates may be biased by differences in
232 fishing and tagging effort across sites. We also identified two fish that were twice recaptured in
233 subsequent peak spawning seasons at the same locations (Twintrees and Huntington Beach Flats,
234 Table 4).
235 Annual trends in Newport Bay recaptures revealed fish were still present and/or had left
236 and returned to Newport Bay during subsequent non-spawning and early spawning seasons (Figure
237 9). Two fish were twice recaptured at Newport Bay over subsequent non-spawning seasons (Table
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13. 238 4). Fish recaptures outside of Newport Bay typically occurred during the middle of peak spawning
239 season through late spawning season (Figure 9).
240
241 Discussion
242 Spawning season residency
243 The spawning residency of migrant barred sand bass at Huntington Beach Flats appears to
244 be approximately one month. This is within the reported spawning residency for other fishes (4 –
245 80 days; Robichaud et al. 2003; Bendal et al. 2005; Douglas et al. 2009). For some species,
246 spawning residency may vary by year and by sex, but we were unable to test this for barred sand
247 bass. Nevertheless, individual barred sand bass with shorter-term residency (e.g., up to one
248 month) likely represent migratory individuals, while fish demonstrating longer-term residency
249 (e.g., up to four months) likely represent fish whose home ranges are located within or near to
250 Huntington Beach Flats. Fish tagged in June or August (the beginning and end of peak spawning
251 season) demonstrated longer-term residency to the spawning grounds than fish tagged in July,
252 suggesting the peak in spawning migrations occurs in July. This timing is in agreement with the
253 month of highest catch numbers and fishing effort for barred sand bass in the southern California
254 CPFV fishery (CDFG unpublished data). Moreover, the majority of fish that were recaptured
255 away from Huntington Beach Flats during peak spawning season were tagged in July. Finally, the
256 expected peak in immigration/emigration rates and associated fishing pressure was apparent in the
257 frequency of June-tagged returns over time. For example, return rates of June-tagged fish
258 decreased after a month and then subsequently increased again before leveling off. We attribute
259 this initial decrease to a peak in immigration to the spawning grounds occurring some time in July,
260 while the subsequent increase and leveling off in return rates is likely associated with a peak in
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14. 261 emigration from the spawning grounds, followed by an associated decrease in fishing pressure
262 there.
263 Due to the use of standard tag and recapture methods, we were unable to quantify diel site
264 fidelity at spawning locations. Barred sand bass individuals may use neighboring reefs to “rest” or
265 forage between spawning events or may travel to and from other nearby spawning locations during
266 this period. Actively tracked coral trout, Plectropomus leopardus, an aggregate spawning
267 Epinephelid (= Serranidae in part; Smith and Craig 2007), were shown to make multiple trips
268 between their home ranges and nearby spawning sites (< 1 km) during the spawning season (Zeller
269 1998). Zeller (1998) also reported that in some cases, visits to spawning grounds occurred
270 multiple times during the day or lasted from two to three days. Several large artificial reefs and
271 structures located near Huntington Beach Flats include Bolsa Chica Artificial Reef (2.7 km W), oil
272 platforms Eva (2.7 km NW), Emma (2.3 km NE), and Esther (6.7 km NW), and the Los Angeles
273 Federal Breakwater (11.8 km NW). Data collected by the California Seafloor Mapping Program
274 (CSMP) reveal patches of hard substrate (i.e., smaller patch reefs or structures) also exist within
275 the Huntington Beach Flats area (CSUMB 2009). Increases of barred sand bass densities during
276 peak spawning season have been observed on SCUBA at Platform Eva (Martin 2008) and the Los
277 Angeles Federal Breakwater (Froeschke et al. 2005). The degree to which these reefs/structures
278 may act as reproductive staging areas for migratory individuals, resting/foraging areas during peak
279 spawning season, or even distinct spawning areas, remains unknown.
280 Movement to and from spawning locations
281 Movement to and from spawning locations is perhaps one of the more interesting and yet
282 least known behaviors of barred sand bass. Our results further suggest that some barred sand bass
283 individuals migrate to spawn, while others are year-round residents. Mason (2008) demonstrated
14
15. 284 that a portion of acoustically monitored adult barred sand bass at Santa Catalina Island, CA,
285 exhibited year-round site fidelity to their home ranges, while others were not detected in these
286 areas during spawning season. This type of “polymorphic movement behavior” has been
287 described for other aggregate spawners including coral trout, Nassau grouper, Epinephelus
288 striatus, and snapper, Pagrus auratus (Colin 1992; Zeller 1998; Egli and Babcock 2004).
289 Not every fish migrated to the same location after spawning season. Likewise, barred sand
290 bass that shared a non-spawning season residence (Newport Bay) did not necessarily migrate to the
291 same spawning grounds. Zeller (1998) reported that coral trout with overlapping home ranges did
292 not necessarily make excursions to the same spawning grounds, regardless of spawning ground
293 proximity to home ranges. Spawning migration distance has been strongly linked to body
294 condition, size-at-age/maturity, and fishing pressure in Atlantic cod (Gadus morhua; Jørgensen et
295 al. 2008); fish that migrate longer distances are generally older, bigger fish with higher overall
296 fitness. Although a relationship between fish size (TL) and “migration” distance in this study was
297 inconclusive, we can not rule out bioenergetics as a possible explanation for the observed
298 variability in migration distances, as maturity and fitness were not determined for tagged fish.
299 Moreover, it is impossible to know for certain the accuracy of the reported fish tag and recapture
300 locations or if some fish were early or late spawners/migrators. These caveats, in addition to the
301 possibility that a portion of the adult barred sand bass population simply doesn’t spawn (i.e.,
302 practices “skipped spawning”), may explain the variability in reported migration distances
303 (Jørgensen et al. 2005).
304 Spawning and non-spawning season site fidelity
305 Most fish tagged during peak spawning season were recaptured at the same location during
306 subsequent peak spawning seasons. Whether these individuals represent year-round residents or
15
16. 307 repeat migrants, barred sand bass appear to show a high degree of breeding site fidelity.
308 Historically, breeding site fidelity was studied extensively in salmonids, but less so in other fishes.
309 With the advent of more sophisticated tracking methods (e.g. electronic tags), breeding site fidelity
310 in fishes has been increasingly documented (Hunter et al. 2003; Block et al. 2005; Douglas et al.
311 2009). Tradition (or learned behavior) has been shown to play a primary role in spawning site
312 selection over annual reassessment of resources, especially if resources are relatively unchanging
313 from one year to the next (Warner 1988, 1990). Due to annual differences in tagging effort across
314 tagging locations, it was not possible to accurately quantify long-term inter-annual variability in
315 spawning site fidelity by tagging location. Recaptures that occurred at different locations in
316 subsequent years may reflect individual variability in the timing of spawning-related movements,
317 movement among aggregation sites, or a certain degree of annual reassessment.
318 We also identified individuals that demonstrated site fidelity to a non-spawning season
319 tagging location (Newport Bay). Bays are noted as habitat for young-of-the year (YOY) and
320 juvenile barred sand bass (Feder et al. 1974; Valle et al. 1999; Allen et al. 2002), while adults
321 prefer sand/rock ecotone habitat to 30 m depth (Feder et al. 1974; Johnson et al. 1990; Mason
322 2008). In San Diego Bay, YOY barred sand bass were shown to develop there throughout their
323 life history and adults utilized the bay habitat throughout the year (Pondella et al. 2006). In this
324 study, a fish that was tagged and recaptured during non-spawning season in Newport Bay may
325 represent a fish that remained there year-round or one that migrated to spawn and returned in the
326 winter. Our data indicate that a portion of adult barred sand bass in Newport Bay migrate to
327 locations outside of the bay during spawning season. This identifies Newport Bay as an important
328 non-spawning season residence for barred sand bass, as well as an important source of migrants to
329 various spawning locations outside of the bay. Although it is unknown for certain whether the
16
17. 330 migrant recaptures in this study would have returned to Newport Bay following peak spawning
331 season, the overall pattern in recaptures is highly suggestive.
332 Recapture rates
333 The most striking difference between the two tagging periods is the recapture rate: 17% in
334 the 1960s compared to 4% in the 1990s. A typical range of recapture rates using standard tag and
335 recapture methods is 3 to 10% (Lowe and Bray 2006). Given that tagging effort and numbers of
336 tagged fish did not dramatically differ between these two time periods, recapture rates may have
337 been influenced by changes in barred sand bass availability, in addition to advances in fishing
338 technology (e.g., fish finders, GPS), the willingness of fishers to report tag returns, and an
339 increased knowledge of spawning aggregation sites. In general, high recapture rates in open
340 systems reflect relatively lower population sizes due to the higher probability of encountering the
341 same fish at a later date. This may explain the higher number of long-term recaptures in the 1960s
342 dataset. CPFV barred sand bass catch values were nearly four times greater in the 1990s than in
343 the 1960s despite only a doubling of fishing effort (CDFG unpublished data). Historical Fish and
344 Game publications report that barred sand bass were scarce during the 1950s (a cold water period)
345 and occurred more frequently along the coast “in and subsequent to periods of warmer waters”
346 (Young 1969; Feder et al. 1974). Kelp bass and barred sand bass larvae densities were also lower
347 during the cool regime (1950s – 1970s) and higher in the warm regime (1980s – 1990s), especially
348 during the 1980s (Moser et al. 2001). While it appears that barred sand bass populations may have
349 increased in the 1990s relative to the 1960s, barred sand bass stock-recruitment relationships and
350 the effects on these relationships by natural and anthropogenic influences remain unknown.
17
18. 351 Management implications
352 Currently, a sustainable level of harvest for barred sand bass is unknown because no
353 biomass estimates exist. Due to concerns over barred sand bass catch declines in recent years and
354 the uncertainty in the status of the population, barred sand bass may require further fishery
355 protection beyond current minimum size limits and bag limits. Information gathered on the
356 spawning-related movements of barred sand bass may be useful for identifying further protection
357 measures. For instance, an aggregate spawner that demonstrates multiple spawning events over a
358 relatively long spawning residency period would likely benefit from a series of temporary fishing
359 closures during the peak in fishing effort. Non-spawning residences that are shown to supply
360 migrant spawners to spawning aggregation sites may be important sites to consider for seasonal
361 protection of barred sand bass. Further consideration of barred sand bass movement patterns and
362 life history traits, in addition to important feasibility concerns, may help to define additional
363 alternatives to protect barred sand bass until a harvest guideline can be developed.
364
365 Acknowledgements
366 CDFG lead investigators for the barred sand bass tagging studies in the 1960s and 1990s
367 were P. “Bud” Young and J.R. Raymond Ally, respectively. Over the two tagging periods, CDFG
368 tagging efforts were augmented by [and listed in no particular order] R. Izor (Izorline
369 International), Orange County Marine Institute, County Sanitation Districts of Orange County, Los
370 Angeles Rod and Reel Foundation, and many other individual volunteers. We thank L.G. Allen
371 and T. Mason for their critical review of the drafts of this manuscript. Funding was supported in
372 part by the Los Angeles County Fish and Game Commission and the Federal Aid in Sportfish
373 Restoration Act (also known as the Dingell-Johnson Act; Grant #F-50-R-20).
18
19. 374 Literature Cited
375 Aburto-Oropeza, O., B. Erisman, V. Valdez-Ornelas, and G. Danemann. 2008. Commercially
376 Important Serranid Fishes from the Gulf of California: Ecology, Fisheries, and Conservation.
377 Ciencia y Conservación 2008:1-23.
378 Allen, L.G., and T.E. Hovey. 2001. Barred sand bass. Pp. 224-225 in California's Living Marine
379 Resources: A Status Report (W.S. Leet, C.M. Dewees, R. Klingbeil & E.J. Larson, eds.),
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23
24. 478 Table 1. Tag and recapture summary statistics for barred sand bass tagged in southern California,
479 historical California Department of Fish and Game tagging project (1960s and 1990s).
Tag and recapture results 1960s 1990s
Tagged fish 4,687 3,947
Tagging effort
days 174 153
fish/day 27 ± 32 26 ± 58
Avg (±SD) (mo/yr) 4±3 6±3
Capture 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 ± 12
Avg (±SD) TL (mm) 306 ± 38 337 ± 72
% mature 89% 93%
% legal size 41% 70%
Recaptures 801 171
Return rate
overall 17% 4%
hook-and-line 17% 5%
bottom trawl -- 3%
Avg (±SD) recapture depth (m) -- 23 ± 9
Avg (±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 1,258
Recapture distance (km)
Avg (±SD) - All fish 6 ± 12 7±9
Avg (±SD) - Only movers 18 ± 15 10 ± 9
Max 92 76
480
24
25. 481 Table 2. Recapture distances (km) of barred sand bass tagged during peak spawning season
482 (Jun-Aug) and recaptured during non-spawning season (Nov-Mar), historical California
483 Department of Fish and Game tagging project (1960s and 1990s). Dir. = direction of recapture
484 location from tagging location.
485
25
26. 486 Table. 3. Recapture distances (km) of barred sand bass tagged in Newport Bay during the non-
487 spawning season (Nov-Mar) and recaptured during peak spawning season (Jun-Aug), historical
488 California Department of Fish and Game tagging project (1960s). Dir. = direction of recapture
489 location from tagging location.
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
26
27. 491 Table 4. Tag and recapture dates and locations of barred sand bass recaptured on two separate occasions, historical California
492 Department of Fish and Game tagging project (1960s).
493
494
495 * Asterisks denote fish that were released in Newport Harbor following weigh-in at a fishing tournament.
27
28. 496 List of Figures
497
498 Figure 1. Map of barred sand bass tagging locations in southern California, historical California
499 Department of Fish and Game tagging project (1960s and 1990s).
500
501 Figure 2. Numbers of barred sand bass tagged by the California Department of Fish and Game
502 in southern California during the a) 1960s and b) 1990s.
503
504 Figure 3. Length-frequency distributions of barred sand bass by tagging location, historical
505 California Department of Fish and Game tagging project (1960s and 1990s). 1960s and 1990s
506 sites are represented by grey and white bars, respectively. Only locations where at least 100
507 tagged individuals are listed. The vertical lines represent minimum size of legal take (305 mm).
508
509 Figure 4. Numbers of tagged barred sand bass recaptured during the a) 1960s and b) 1990s,
510 historical California Department of Fish and Game tagging project (1960s and 1990s).
511
512 Figure 5. Numbers of barred sand bass recaptured by days since tagged for fish tagged in the a)
513 1960s and b) 1990s, historical California Department of Fish and Game tagging project (1960s
514 and 1990s).
515
516 Figure 6. Cumulative number of barred sand bass recaptures (dots) and frequency of recaptures
517 (bars) by days since tagged for fish tagged during each month of peak spawning season (June,
28
29. 518 July, August) at Huntington Beach Flats, CA and recaptured within the same year at the same
519 location, historical California Department of Fish and Game tagging project (1960s and 1990s).
520
521 Figure 7. Recapture distances for barred sand bass tagged during peak spawning season (Jun-
522 Aug) and recaptured during non-spawning season (Nov-Mar), historical California Department
523 of Fish and Game tagging project (1960s and 1990s). 1960s = dark bars, 1990s = grey bars.
524
525 Figure 8. Recapture matrix plot of barred sand bass tagged during peak spawning season and
526 recaptured in subsequent peak spawning seasons, historical California Department of Fish and
527 Game tagging project (1960s and 1990s). Shaded, darker boxes occurring along the diagonal
528 line indicate a higher degree of breeding site fidelity.
529
530 Figure 9. Recapture plot of barred sand bass tagged in Newport Bay, California during non-
531 spawning season (Nov-Mar, shaded areas), historical California Department of Fish and Game
532 tagging project (1960s). Recaptures occurring within 90 d of tagging were excluded. Recapture
533 locations are arranged north to south (top to bottom).
29
