Sepioteuthis Sepioidea (Cephalopoda: Loliginidae) ∗ 5 S.E

+Model FISH 2256 1–6 ARTICLE IN PRESS

Fisheries Research xxx (2006) xxx–xxx

3 A preliminary investigation of the use of subcutaneous tagging in Caribbean

4 reef squid Sepioteuthis sepioidea (Cephalopoda: Loliginidae) ∗ 5 S.E. Replinger, J.B. Wood

6 Bermuda Institute of Ocean Science, St. George’s GE 01, Bermuda Received 18 July 2006; received in revised form 2 November 2006; accepted 9 November 2006

8 Abstract

9 This paper describes a new method developed to directly measure size and temperature specific growth rates of individual wild squid. Our 10 tag and recapture method is complimentary to previously employed statolith aging methods but provides finer scale detail. Caribbean reef squid, 11 Sepioteuthis sepioidea, is an ideal model organism for field work on squid life-history as they live in shallow coastal areas and are accessible. 12 These squid were tagged and monitored for tag retention and growth rates over a period of 56 days in the laboratory and 27 days in the field. A total 13 of 103 squid were tagged, 10 in the laboratory and 93 in the field. Visible Implant Elastomer (VIE) tags and Visible Implant Alphanumeric (VI 14 Alpha) tags were used during this experiment. None of the VI Alpha tags were retained in laboratory animals, but VIE tags remained visible for 15 the duration of the laboratory study and were used in all field studies. The 10 recaptured squid weighed from 19.1 to 122.9 g with an average and 16 standard deviation recapture weight of 56.9 ±35.5 g, compared to their original weights of 48.0 ±30.5 g. In the field, these squid had instantaneous ◦ 17 growth rates of 1.19–3.10 with an average growth rate of 1.93 ± 0.71 at temperatures ranging from 19.5 to 23.7 C. 18 © 2006 Published by Elsevier B.V.

19 Keywords: Squid; Tag; Growth; Sepioteuthis; VIE; Cephalopoda

1 1. Introduction ious life cycles make rational fisheries management a challenge 19 (Rodhouse, 2001). 20 2 Fisheries around the world are shifting focus to species of Statolith aging methods have provided invaluable informa- 21 3 lower trophic levels, such as squid, as fish from the higher tion regarding the life-history strategies of squid. However, there 22 4 trophic levels become less abundant (Caddy and Rodhouse, are limitations with this technique: (1) statoliths give only an esti- 23 5 1998; Pauly et al., 1998; Piatkowski et al., 2001; Jereb and Roper, mate of the age of a squid, although these estimates have been 24 6 2005). Squid are popular food items in many areas of the world validated for many species such as Loligo plei (Jackson and 25 7 (Sugiuama et al., 1989; Okuzumi and Fujii, 2000; Rocha and Forsythe, 2002), Loligo vulgaris (Arkhipkin, 1995) and Sepi- 26 8 Vega, 2003), and humans currently harvest over 2.5 million tons oteuthis lessoniana (Jackson, 1990); (2) the initial hatchling 27 9 of cephalopods (including squid, octopus, and cuttlefish) each weight of squid is often estimated when using statoliths to deter- 28 10 year (World Fisheries, 2003; Jereb and Roper, 2005). As a result mine growth rates and this estimate may have a large impact on 29 11 of the increasing pressure on squid fisheries worldwide, a better squid growth models (Pecl et al., 2004) and (3) statolith work is 30 12 understanding of their life histories and growth rates is important usually done after several seasons of growth. The range of tem- 31 13 to properly manage the fishery (Jackson et al., 2000). To date, a peratures experienced is likely to vary considerably between 32 14 fine scale method that allows individual squid to be tracked so seasons, which can be problematic since squid are extremely 33 15 that their size specific growth rates in the wild can be obtained responsive to changes in temperature. While it is possible to 34 16 over short time periods has not been developed. Squid popu- consider the impact of temperature on the age and/or size spe- 35 17 lations are highly variable with recruitment being responsive to cific growth rates of squid using statolith studies (Arkhipkin and 36 18 environmental change. This variability and their short semelpar- Laptikhovsky, 1994; Ikeda et al., 2003), it can be difficult with 37 wild squid (e.g., Forsythe, 1993; Jackson et al., 1997; Forsythe 38 et al., 2001; Jackson and Moltschaniwskyj, 2002). 39 ∗ Corresponding author. Tel.: +441 297 1880. While statolith analysis provides an average growth rate for 40 E-mail address: [email protected](J.B. Wood). a populationPROOF of squid over a lifetime, tagging studies allow 41

Please cite this article in press as: Replinger, S.E., Wood, J.B., A preliminary investigation of the use of subcutaneous tagging in Caribbean reef squid Sepioteuthis sepioidea (Cephalopoda: Loliginidae), Fish. Res. (2006), doi:10.1016/j.ﬁshres.2006.11.028 +Model FISH 2256 1–6 ARTICLE IN PRESS

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42 growth rates of individual squid to be studied over shorter 43 time periods. This will allow for a better understanding of 44 how other factors such as temperature (Forsythe, 1993; Jackson 45 et al., 1997; Forsythe et al., 2001; Wood and O’Dor, 2000; 46 Jackson and Moltschaniwskyj, 2002), food rations (Jackson and 47 Moltschaniwskyj, 2001), and age (Forsythe, 1993; Forsythe et 48 al., 2001; Pecl et al., 2004) can affect the growth rates of squid. 49 Although there has been a considerable amount of cephalo- 50 pod growth data from captive rearing studies, fisheries scientists 51 remain reluctant to use this laboratory generated data. Captive Fig. 1. Double-tagging of squid using VIE tags. The ventral side of the squid 52 squid do not grow in a manner identical to wild squid, so culture is shown (a) in the diagram with lines indicating the locations of the VIE tags. 53 studies may not accurately reflect wild populations (Pecl and Tagging of the squid (b) required the squid to be out of the water for a short time 54 Moltschaniwskyj, 1999). An alternative method to determine while the tag was inserted. 55 growth rates of wild squid populations is to periodically capture 56 schools of squid in the same area. However, this has proven to of tag tested in this study were the VI Alpha tags, which are 99 57 be an unsuccessful technique due to the migration of microco- small pieces of plastic with a unique three-digit alphanumeric 100 58 horts of squid through time and space (O’Dor, 1998). The squid code that are also inserted under the skin of the animal. While 101 59 captured in the same area were not representative of the squid a pattern of lines must be used to identify individuals with the 102 60 that had been previously caught at a particular location. VIE tags, VI Alpha tags can provide a more definitive identifi- 103 61 We developed a method to tag individual Caribbean reef cation of individuals. Six families of amphibians, 3 families of 104 62 squid, Sepioteuthis sepioidea, in order to better understand the crustaceans, 24 families of fish and 1 reptilian family have been 105 63 size specific growth of wild squid. Tagging and recapturing tagged with VI Alpha tags (NMT, 2006). 106 64 individual squid in the wild allows growth rates to be directly S. sepioidea were chosen for this experiment because they 107 65 measured over shorter time periods, which is important because live in shallow coastal waters (Boycott, 1965) and therefore are 108 ◦ 66 even a small increase in temperature, such as a change of 1 C, accessible. Individual adult squid often remain in the same loca- 109 67 will greatly impact the growth rates of fast growing ectotherms tion on successive days, which enhances the ability to recapture 110 68 such as hatchling and juvenile squid (Fulton, 1904; Rodhouse individuals, and thus monitor growth rates and tag retention 111 69 and Hatfield, 1990; Forsythe, 1993, 2004). The ability to follow (Moynihan and Rodaniche, 1982; Mather and Wood personal 112 70 individuals in the wild and study how growth rates change during observations). The ability to successfully raise and keep S. sepi- 113 71 seasonal water temperature variations will enhance our under- oidea in captivity (La Roe, 1971) is also important because it 114 72 standing of this principle. Additionally, variability in hatchling suggests that this species may be relatively hardy and better able 115 73 size (Steer et al., 2003; Pecl et al., 2004) and size specific sur- to survive the stresses associated with tagging. Keeping squid 116 74 vivorship (Steer et al., 2003) could overestimate squid growth in captivity was important for this study because it allowed us 117 75 rates as current models often assume an average hatchling size to develop a tagging method in the laboratory as well as obtain 118 76 and ignore the effect of hatchling size on survivorship. Directly an estimate of wet weight error data. 119 77 measuring growth in individual squid provides data that is not

78 subject to assumptions of hatchling size and survivorship. 2. Materials and methods 120 79 In this study, we tested both VIE tags and VI Alpha tags, both

80 of which are small and inserted subcutaneously which leaves We investigated the tag retention rate for Visible Implant 121 81 nothing hanging outside of the cephalopods body, this reduces Elastomer (VIE) tags and Visible Implant Alphanumeric (VI 122 82 the chance of infection. Both the VIE and VI Alpha tags ﬂuoresce Alpha) tags to measure the growth of individual Caribbean reef 123 83 under a blue LED light, improving tag visibility even through the squid S. sepioidea in the laboratory and in the ﬁeld. The tags 124 84 pigmented skin of the squid. These tags have not previously been were provided by Northwest Marine Technology, Shaw Island, 125 85 used with cephalopods, but past studies with external tags have Washington, USA. 126 86 shown that the behavior of squid post-tagging appears normal

87 (Sauer et al., 2000). However, external tags commonly damage 2.1. Growth rates 127 88 the delicate skin of squid, resulting in wounds that do not heal

89 (Sauer et al., 2000). Therefore, in this study we chose internal Growth rates were calculated for the periods between each 128 90 tags in an attempt to reduce the physical effects of the tags on weighing using the instantaneous growth rate equation (Forsythe 129 130 91 the squid. and Van Heukelem, 1987): 92 The VIE tags consist of a liquid elastomer that is inserted ln W2 − ln W1 93 subcutaneously and hardens to form a colored marking. Mul- G = × 131 T − T 100 94 tiple lines of different colors placed in different locations can 2 1 95 be used to identify individualsUNCORRECTED (Fig. 1). These tags have been where PROOFW1 and W2 are the wet weights at time 1 and 2 and 132 96 successfully used on over 40 families of ﬁsh (e.g., FitzGerald et (T2 − T1) is the time interval in days. 133 97 al., 2004), 5 families of crustaceans (e.g., Sellars et al., 2004), as Before weighing, the squid were held head down to drain 134 98 well as amphibians and reptiles (NMT, 2006). The second type excess water out of the mantle cavity before placing it on a 135

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136 balance with an error of 0.1 g or less. This source of error inherent repeated for each individual. During this process, the time that 189 137 in the weighing process was quantified by weighing four squid the squid spend out of the water was minimized. In all subse- 190 138 four to six times each in the laboratory to calculate the amount quent rounds, each squid was examined through the bottom of 191 139 of variation caused by water. It was determined that the wet a clear glass fish bowl to determine its identity. 192 140 weight error for squid was 1.4% of their body weight. In order Visible Implant Alphanumeric (VI Alpha) tags were also 193 141 to minimize the influence of this wet weight weighing error on tested. The ventral side of the fins, close to the area where the 194 142 calculated growth rates, we only used recapture data with an body and fin of the squid meet was determined to be a more 195 143 interval of 7 days or more between each weighing. acceptable location for the tags than the ventral side of the 196 mantle. As these tags were not being retained, we tried NWT 197 144 2.2. Maintaining squid in captivity suggestion of adding a small amount of veterinary glue (Vet 198 Bond) in the insertion hole to help prevent the tag from coming 199 145 S. sepioidea were captured using a seine net from Whale- out. 200 ◦ ◦ 146 bone Bay (32 21.872 64 42.781 ), St. George Island, Bermuda Approximately every 10 days, the tags (VIE and VI Alpha) 201 147 and were acclimated to captivity over a period of approximately were examined and the squid’s weight and mantle lengths were 202 148 6 weeks in the lab at the Bermuda Institute of Ocean Science recorded. A blue LED light and amber-filter glasses were used 203 149 (BIOS). Squid were kept in a long, shallow, rectangular cement when necessary to illuminate the tags. This continued for the 204 150 tank approximately 525 cm long and 100 cm wide with a water duration of the 56 days that the squid were monitored during this 205 151 depth of about 40 cm. The tank was enriched with natural rocks study. Growth rates and average temperatures were calculated 206 152 and cement bricks. A fresh supply of seawater was continu- for each period. 207 153 ally fed into the approximately 2100 l tank at a rate of 1.5 l per 154 minute, and the water was circulated with airstones. Screening 2.4. Squid tagging in the field 208 155 was placed over the tank to prevent mortality caused by squid 156 jumping out of the tank. The tank was setup in a quiet lab to Following the success of laboratory tagging of squid using 209 157 avoid disturbances. The lights in the room were slowly turned VIE tags, a field tagging trial commenced at various loca- 210 158 off in the evening and were turned on again in the mornings tions on St. George’s Island in Bermuda. These locations were 211 ◦ ◦ 159 over a period of several minutes to simulate a natural sunrise. Whalebone Bay, 32 21.872 N, 64 42.781 W(n = 3 groups of 212 ◦ ◦ 160 Temperature was recorded daily. squid), Ferry Reach Bridge, 32 21.727 N, 64 42.923 W(n = 1), 213 ◦ ◦ 161 Approximately four times a day, squid were fed a diet of Achilles Bay, 32 23.430 N64 40.528 W(n = 4), and Concrete 214 ◦ ◦ 162 frozen silversides (Family Clupeidae and Engraulidae, mostly Beach, 32 20.178 N, 64 41.820 W(n = 5). Three squid were 215 163 Anchoa choerostoma and Jenkinsia lamprotaenia) which were also tagged near the Bermuda airport on St. David’s Island just 216 ◦ ◦ 164 thawed before feeding. These fish were caught locally using a off the causeway, 32 21.872 N, 64 42.781 W. Overall, a total 217 165 beach seine net or were bought from a local fisherman. Also, of 93 squid were tagged in the field between 21 October 2004 218 166 live silversides, small reef squirrelfish (Holocentrus coruscus) and 17 November 2004, 10 of which were recaptured with a 219 167 and small french grunts (Haemulon falvolineatum) were occa- long enough time interval that we could use the data to calculate 220 168 sionally captured and fed live to the squid. growth rates. We required a 7-day interval between captures in 221 order to minimize the effect of wet weight error in our growth 222 169 2.3. Squid tagging in captivity rates. During this period, we made 19 attempts to tag squid in 223 the field. Of those 19 attempts we successfully tagged one or 224 170 The initial tagging treatment was done with Visible Implant more squid on 14 occasions. 225 171 Elastomer (VIE) Tags following NMT’s instructions. Dive Squid were caught using a beach seine net. Two people held 226 172 gloves were worn when handling the squid since they can bite the the seine net at each end so that the leading edge rested on the 227 173 handler. In addition, a mesh holding tank was set up at one end bottom while the other floated at the surface. Small rocks were 228 174 of the squid tank to separate the tagged from the untagged indi- placed inside the net to weigh down the mesh. Additional helpers 229 175 viduals. Ten captive squid were tagged and their mantle length gently guided the squid over the net, at which point the edges 230 176 and weight were recorded. The squid were double-tagged based were quickly lifted out of the water to trap the squid. After being 231 177 on a pre-designated pattern of horizontal, diagonal, and verti- caught, squid were kept on location in a floating mesh holding 232 178 cal lines (Fig. 1). Double tagging was done to aid in studying tank that was approximately 130 cm long by 80 cm wide with a 233 179 the tag retention rate and so that the squid could be more accu- water depth of 30 cm. The sides of the tank floated about 50 cm 234 180 rately identified. All tagging was done on the ventral side of above the water to prevent the squid from jumping out. The 235 181 the mantle since this is generally the least pigmented part of the holding tank was divided into two parts, which enabled us to 236 182 squid. As a backup for the tags, the dorsal side of the squid were separate the squid that we had already tagged. 237 183 photographed with a digital camera so that individuals could be A portable battery-powered scale allowed processing of the 238 184 identified using their unique dot patterns (Byrne et al., 2002). squid to be done on location in order to minimize transportation 239 185 Finally, squid were placed in aUNCORRECTED fish tank with water that had been and PROOF thus stress on the squid. Care was taken to reduce the time 240 186 previously tarred and were weighed using an electronic balance. that squid spent out of water, as this was the greatest cause of 241 187 After being weighed, squid were gently released into the hold- stress during the tagging process and the most likely cause of 242 188 ing tank. This procedure, which took about 3 min per squid, was mortalities in laboratory work. When the squid were tagged, 243

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244 one individual held the squid ventral side up while a second individually identified. The squid ranged in size from 1.8 g with 281 245 individual tagged the squid. a mantle length of 19 mm to 149.4 g with a mantle length of 282 246 Each squid was individually photographed, weighed, had its 123 mm. The average weight of the squid was 40.2 g and the 283 247 mantle length measured, and was tagged using VIE tags. Pre- average mantle length was 67 mm. No mortalities were observed 284 248 determined tagging patterns similar to those used for the captive prior to re-release in the field. 285 249 squid were used in the field (Fig. 1). All squid were then returned We recaptured 10 of the 93 squid tagged in the field between 286 250 to the holding tank until they could all be released together 21 October and 20 November 2004 (Table 1). While one squid 287 251 approximately 15 min after tagging ended. Since three of the was found over a nautical mile from where it was originally 288 252 four mortalities of captive squid occurred within 15 min after tagged, the other nine squid that were recaptured were caught 289 253 they were tagged, this waiting period allowed us to monitor at the location they were tagged. On three separate occasions 290 254 tagging mortality in the field. groups of similarly sized squid were found in the same part of 291 Achilles Bay, but none of the squid that were captured had been 292

255 3. Results previously tagged. 293

256 Visible Implant Elastomer (VIE) tags were successfully used 3.3. Growth rates 294 257 to directly measure size speciﬁc growth rates in individual squid

258 both in the laboratory and in the ﬁeld. In captivity, the average growth rate for the squid was 1.66% 295 body weight per day, with a minimum of −1.4 and a maxi- 296

259 3.1. Squid tagging in captivity mum of 5.3. The average temperatures for the growth periods 297 ◦ ◦ ranged from 27.1 C down to 22.9 C over the duration of the 298

260 In captivity, both VIE tags and Visible Implant Alphanumeric experiment. In the ﬁeld, the growth rates for the 10 recaptured 299 261 (VI Alpha) tags were tested for ease of use and retention. The squid ranged from 1.2 to 3.1, with an average growth rate of 1.9 300 262 VIE tags on ventral side of the mantle and on the ﬁns of the (Table 1). The water temperatures for the squid tagged in the 301 ◦ 263 captive squid were retained for the duration of the 56 days of the wild ranged from 25.3 to 19.5 C. 302 264 study and continued to be usable for identifying the squid. With

265 increasing user experience, the tags became easier to insert and 4. Discussion 303 266 the mortally rate decreased.

267 The VI Alpha tags were not retained in any of the first two Growth rate data generated by the tag and recapture method 304 268 trials with captive squid (n =7,n = 5). As these initial attempts developed in this study compliments that of statolith and captive 305 269 to insert these tags on the ventral side of the mantle resulted in culture work. While our current knowledge of squid growth rates 306 270 the deaths of two squid, subsequent attempts to use these tags in the field has been greatly enhanced by the use of statolith aging 307 271 were done on the fins. In the final set of laboratory trials, six methods, tagging studies help to address the unknowns that are 308 272 squid were tagged, and veterinary glue (Vet Bond) was used to inherent in the statolith method of studying squid growth: mainly 309 273 prevent the tags from coming out of the insertion holes. Two the instantaneous growth rates of squid at specific sizes, and the 310 274 additional mortalities can be attributed to this attempt to use the impact of small temperature changes on these growth rates. 311 275 VI Alpha tags, one immediately following tagging and the other Small changes in temperatures have been shown to have 312 276 after several days has passed. When squid were examined 5 days a large effect on the exponential growth phase of hatchling 313 277 later, only one of the VI Alpha tags was retained. squid (Forsythe, 1993; Jackson et al., 1997; Forsythe et al., 314 2001; Wood and O’Dor, 2000; Jackson and Moltschaniwskyj, 315 278 3.2. Squid tagging in the field 2002). Most statolith field work is over relatively long time 316 intervals where temperature is likely to be extremely variable. 317 279 Overall, a total of 93 squid were tagged in the field using Researchers are starting to attack this problem of unknown tem- 318 280 VIE tags in unique patterns (Fig. 1) so that all squid could be peratures over long time periods by investigating the correlation 319

Table 1 Summary of data for the 10 squid tagged and recaptured in the ﬁeld Squid number Location tagged Location recaptured Start weight (g) End weight (g) Average temperature (◦C) Instantaneous growth rate

24 Whalebone Whalebone 22.2 29.1 22.8 2.71 26 Whalebone BIOS 19.7 22.2 – 1.99 51 BIOS BIOS 102.4 122.9 21.4 1.66 52 BIOS BIOS 50.7 69.2 21.4 2.83 53 BIOS BIOS 14.9 19.1 21.8 3.10 54 BIOS BIOS 50.8 59.3 21.4 1.41 56 BIOSUNCORRECTED BIOS 42.1 48.7 PROOF 23.0 1.82 57 BIOS BIOS 37.2 40.9 23.0 1.19 57 BIOS BIOS 40.9 46.6 21.4 1.19 60 BIOS BIOS 99.5 111.4 23.0 1.41

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320 between temperature and strontium to calcium ratios is statoliths tags. Scientists interested in real time tracking cephalopods 377 321 (Ikeda et al., 2003). When using tag and recapture methods, the and direct continuous environmental sampling should consider 378 322 temperature can be directly measured at the location where the larger acoustic/archival tags (Jackson et al., 2005) or pop-up 379 323 squid are captured. The shorter time intervals in tagging studies, tags (Block et al., 2002). Currently, the smallest coded trans- 380 324 on the order of weeks, provides less temperature variation than mitters available are the Vemco V7-1L which measures 7 mm 381 325 the longer time intervals of months or seasons that are typically by 17.5 mm and weighs only 1.4 g (Vemco, 2006). However, 382 326 used in statolith work. these tags are not yet small enough for growth work on juvenile 383 327 A combination of both statolith and tag and recapture meth- cephalopods because they are similar in size to the smaller squid 384 328 ods would provide both age and size specific growth rates. we tagged in this study (1.8 g with 19 mm mantle length). 385 329 Statolith work gives an overall estimate of the instantaneous Of all physical tags available, we believe that Visible Implant 386 330 growth rate of squid for a long time period, while the tag and Elastomer (VIE) tags and other similar tags are the least likely 387 331 recapture method provides the instantaneous growth rate for a to affect the behavior and growth in all species of cephalopods. 388 332 much shorter time period. Knowing both of these growth rates These tags, made of medical grade elastomer, are small, internal 389 333 would allow scientists to: (1) calculate both size and age spe- and flexible, and therefore, they do not cause drag, infection, or 390 334 cific growth rates during the tagging interval and (2) use these irritation of contracting muscles (Roberts et al., 1973a,b,c). After 391 335 known growth rates to calculate and better understand the rapid the success of squid tagging using VIE tags, the corresponding 392 336 growth rate for paralarval and juvenile squid data that has been author has also started preliminary use of these tags on other 393 337 notoriously difficult to obtain in the field. species of cephalopods such as Octopus vulgaris and Octopus 394 338 There are two main limitations that can hinder the use of macropus. 395 339 tagging studies. The first is the need to recapture the same squid The growth rates of individual reef squid can be directly mea- 396 340 in order to obtain data. Oceanic and continental shelf squid can sured in the field using VIE tags. Reef squid are ideal model 397 341 migrate great distances, moving rapidly through time and space, squid as they are relatively hardy (for squid), can be kept in cap- 398 342 thus squid caught in the same area are not representative of the tivity to validate the method, and are found near shore. Future 399 343 same population from month to month (O’Dor, 1998). Ikeda et studies using this method will be able to investigate the effects 400 344 al. (2003) tagged 10,354 squid (Todarodes pacificus), and even of biotic and abiotic factors such as temperature and size on 401 345 with help of Japanese fishermen, was only able to recaptured squid growth rates. In combination with statolith methods, this 402 346 0.88%. When it is necessary to initially tag such large numbers method can be used to examine the temporal variation in age 403 347 of squid to ensure some recaptures, it would be very difficult to specific growth rates as well as provide finer scale resolution for 404 348 process and weigh each squid so that growth could be directly better understanding the multi-phase growth of squid species. 405 349 measured. As we continue to fish down the food chain and rely more 406 350 While we have shown that coastal squid such as S. sepi- on short live opportunist species such as squid (Pauly, 1998; 407 351 oidea can be successfully studied with our tag and recapture Rodhouse, 2001), understanding the life-history of squid will 408 352 method, we believe that many of them, especially the paralar- become increasingly important. 409 353 val and smaller juveniles also move through time and space 354 on a smaller scale. At Achilles Bay, schools of similarly sized Acknowledgements 410 355 squid were caught in the same location of the bay on three sep- 356 arate occasions without obtaining a single recapture. Although We thank Northwest Marine Technology (Shaw Island, 411 357 we looked extensively, we did not find additional schools within Washington, USA) for providing us with the tagging materi- 412 358 1 km of the area or locate any of the squid that we had previously als for this study. Additionally, we thank the National Science 413 359 tagged. Foundation (NSF) for supporting the Research Experience for 414 360 The second limitation of our tagging process is squid moral- Undergraduates (REU) program at the Bermuda Biological Sta- 415 361 ities that occur due to excessive time spent out of the water. tion for Research which supported the primary author during 416 362 Although S. sepioidea is a relatively hardy species for a squid, this study. 417 363 increased handler experience with the tags and tagging process We also appreciate all the help we received in our efforts to 418 364 was necessary to obtain low mortality rates. While tagging Sep- catch and tag squid from many volunteers at the BBSR. Helpers 419 365 tioteuthis sepioidea, the most common cause of death appeared included NSF-REU students Walt Carlson, Victoria Conti, Erin 420 366 not to be from the tags themselves, but rather from stresses asso- Culpepper, Jason Helyer, Kristen Henderson, Ashley Maloney, 421 367 ciated with the out of water handling time needed to insert the and Paul Pawelzik; as well as Amanda Pilch, Ariane Graf, 422 368 tags and especially the time needed to weigh the squid. Jeff Zola, Justin Anderson, Nancee Kumpfmiller, and Kevin 423 424 369 In the past, tag and recapture research has been aimed at inves- Kumpfmiller. We also thank John Forsythe, Alison King, Kim 370 tigating the migration of squid, and recaptures were obtained Zeeh, and our two anonymous reviewers for comments on drafts 425 426 371 through the reporting of tags by fishermen (Ikeda et al., 2003). of this manuscript. 372 However, this past research has evaluated the effects of tags on References 427 373 squid behavior. One study foundUNCORRECTED that after being tagged, Chokka PROOF 374 squid (Loligo vulgaris reynaudii) were able to swim normally, Arkhipkin, A., 1995. Age, growth, and maturation of the European squid Loligo 428 375 and males were observed engaging in normal courtship behav- vulgaris (Myopsida, Loliginidae) on the West Saharan Shelf. J. Mar. Biol. 429 376 iors (Sauer et al., 2000), despite the use of external, highly visible Ass. UK 75, 593–604. 430

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