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2006 Elsevier B.V. All rights reserved.
Keywords: Squid; Tag; Growth; Sepioteuthis; VIE; Cephalopoda
1. Introduction
Fisheries around the world are shifting focus to species of
lower trophic levels, such as squid, as sh from the higher
trophic levels become less abundant (Caddy and Rodhouse,
1998; Paulyet al., 1998; Piatkowski et al., 2001; JerebandRoper,
2005). Squid are popular food items in many areas of the world
(Sugiuama et al., 1989; Okuzumi and Fujii, 2000; Rocha and
Vega, 2003), and humans currently harvest over 2.5 million tons
of cephalopods (including squid, octopus, and cuttlesh) each
year (World Fisheries, 2003; Jereb and Roper, 2005). As a result
of the increasing pressure on squid sheries worldwide, a better
understanding of their life histories and growth rates is important
to properly manage the shery (Jackson et al., 2000). To date, a
ne scale method that allows individual squid to be tracked so
that their size specic growth rates in the wild can be obtained
over short time periods has not been developed. Squid popu-
lations are highly variable with recruitment being responsive to
environmental change. This variability and their short semelpar-
C,
will greatly impact the growth rates of fast growing ectotherms
such as hatchling and juvenile squid (Fulton, 1904; Rodhouse
and Hateld, 1990; Forsythe, 1993, 2004). The ability to follow
individuals in the wild and study howgrowth rates change during
seasonal water temperature variations will enhance our under-
standing of this principle. Additionally, variability in hatchling
size (Steer et al., 2003; Pecl et al., 2004) and size specic sur-
vivorship (Steer et al., 2003) could overestimate squid growth
rates as current models often assume an average hatchling size
and ignore the effect of hatchling size on survivorship. Directly
measuring growth in individual squid provides data that is not
subject to assumptions of hatchling size and survivorship.
In this study, we tested both VIE tags and VI Alpha tags, both
of which are small and inserted subcutaneously which leaves
nothing hanging outside of the cephalopods body, and reduces
the chance of infection. Boththe VIEandVI Alpha tags uoresce
under a blue LEDlight, improving tag visibility even through the
pigmented skin of the squid. These tags have not previously been
used with cephalopods, but past studies with external tags have
shown that the behavior of squid post-tagging appears normal
(Sauer et al., 2000). However, external tags commonly damage
the delicate skin of squid, resulting in wounds that do not heal
(Sauer et al., 2000). Therefore, in this study we chose internal
tags in an attempt to reduce the physical effects of the tags on
the squid.
The VIE tags consist of a liquid elastomer that is inserted
subcutaneously and hardens to form a colored marking. Mul-
tiple lines of different colors placed in different locations can
be used to identify individuals (Fig. 1). These tags have been
successfully used on over 40 families of sh (e.g., FitzGerald et
al., 2004), 5 families of crustaceans (e.g., Sellars et al., 2004), as
well as amphibians and reptiles (NMT, 2006). The second type
Fig. 1. Double-tagging of squid using VIE tags. The ventral side of the squid
is shown (a) in the diagram with lines indicating the locations of the VIE tags.
Tagging of the squid (b) required the squid to be out of the water for a short time
while the tag was inserted.
of tag tested in this study were the VI Alpha tags, which are
small pieces of plastic with a unique three-digit alphanumeric
code that are also inserted under the skin of the animal. While
a pattern of lines must be used to identify individuals with the
VIE tags, VI Alpha tags can provide a more denitive identi-
cation of individuals. Six families of amphibians, 3 families of
crustaceans, 24 families of sh and 1 reptilian family have been
tagged with VI Alpha tags (NMT, 2006).
S. sepioidea were chosen for this experiment because they
live in shallow coastal waters (Boycott, 1965) and therefore are
accessible. Individual adult squid often remain in the same loca-
tion on successive days, which enhances the ability to recapture
individuals, and thus monitor growth rates and tag retention
(Moynihan and Rodaniche, 1982; Mather and Wood personal
observations). The ability to successfully raise and keep S. sepi-
oidea in captivity (La Roe, 1971) is also important because it
suggests that this species may be relatively hardy and better able
to survive the stresses associated with tagging. Keeping squid
in captivity was important for this study because it allowed us
to develop a tagging method in the laboratory as well as obtain
an estimate of wet weight error data.
2. Materials and methods
We investigated the tag retention rate for Visible Implant
Elastomer (VIE) tags and Visible Implant Alphanumeric (VI
Alpha) tags to measure the growth of individual Caribbean reef
squid S. sepioidea in the laboratory and in the eld. The tags
were provided by Northwest Marine Technology, Shaw Island,
Washington, USA.
2.1. Growth rates
Growth rates were calculated for the periods between each
weighing using the instantaneous growth rate equation (Forsythe
and Van Heukelem, 1987):
G =
ln W
2
ln W
1
T
2
T
1
100
where W
1
and W
2
are the wet weights at time 1 and 2 and
(T
2
T
1
) is the time interval in days.
Before weighing, the squid were held head down to drain
excess water out of the mantle cavity before placing it on a
310 S.E. Replinger, J.B. Wood / Fisheries Research 84 (2007) 308313
balance withanerror of 0.1 gor less. This source of error inherent
in the weighing process was quantied by weighing four squid
four to six times each in the laboratory to calculate the amount
of variation caused by water. It was determined that the wet
weight error for squid was 1.4% of their body weight. In order
to minimize the inuence of this wet weight weighing error on
calculated growth rates, we only used recapture data with an
interval of 7 days or more between each weighing.
2.2. Maintaining squid in captivity
S. sepioidea were captured fromWhalebone Bay (32
21.872
64
42.781
21.872
N, 64
42.781
W (n =3 groups of
squid), Ferry Reach Bridge, 32
21.727
N, 64
42.923
W (n =1),
Achilles Bay, 32
23.430
N 64
40.528
20.178
N, 64
41.820
21.872
N, 64
42.781
W. Overall, a total
of 93 squid were tagged in the eld between 21 October 2004
and 17 November 2004, 10 of which were recaptured with a
long enough time interval that we could use the data to calculate
growth rates. We required a 7-day interval between captures in
order to minimize the effect of wet weight error in our growth
rates. During this period, we made 19 attempts to tag squid in
the eld. Of those 19 attempts we successfully tagged one or
more squid on 14 occasions.
Squid were caught using a beach seine net. Two people held
the seine net at each end so that the leading edge rested on the
bottom while the other oated at the surface. Small rocks were
placed inside the net to weigh down the mesh. Additional helpers
gently guided the squid over the net, at which point the edges
were quickly lifted out of the water to trap the squid. After being
caught, squid were kept on location in a oating mesh holding
tank that was approximately 130 cm long by 80 cm wide with a
water depth of 30 cm. The sides of the tank oated about 50 cm
above the water to prevent the squid from jumping out. The
holding tank was divided into two parts, which enabled us to
separate the squid that we had already tagged.
A portable battery-powered scale allowed processing of the
squid to be done on location in order to minimize transportation
and thus stress on the squid. Care was taken to reduce the time
that squid spent out of water, as this was the greatest cause of
stress during the tagging process and the most likely cause of
mortalities in laboratory work. When the squid were tagged,
S.E. Replinger, J.B. Wood / Fisheries Research 84 (2007) 308313 311
one individual held the squid ventral side up while a second
individual tagged the squid.
Each squid was individually photographed, weighed, had its
mantle length measured, and was tagged using VIE tags. Pre-
determined tagging patterns similar to those used for the captive
squid were used in the eld (Fig. 1). All squid were then returned
to the holding tank until they could all be released together
approximately 15 min after tagging ended. Since three of the
four mortalities of captive squid occurred within 15 min after
they were tagged, this waiting period allowed us to monitor
tagging mortality in the eld.
3. Results
Visible Implant Elastomer (VIE) tags were successfully used
to directly measure size specic growth rates in individual squid
both in the laboratory and in the eld.
3.1. Squid tagging in captivity
In captivity, both VIEtags and Visible Implant Alphanumeric
(VI Alpha) tags were tested for ease of use and retention. The
VIE tags on ventral side of the mantle and on the ns of the
captive squid were retained for the duration of the 56 days of the
study and continued to be usable for identifying the squid. With
increasing user experience, the tags became easier to insert and
the mortally rate decreased.
The VI Alpha tags were not retained in any of the rst two
trials with captive squid (n =7, n =5). As these initial attempts
to insert these tags on the ventral side of the mantle resulted in
the deaths of two squid, subsequent attempts to use these tags
were done on the ns. In the nal set of laboratory trials, six
squid were tagged, and veterinary glue (Vet Bond) was used to
prevent the tags from coming out of the insertion holes. Two
additional mortalities can be attributed to this attempt to use the
VI Alpha tags, one immediately following tagging and the other
after several days has passed. When squid were examined 5 days
later, only one of the VI Alpha tags was retained.
3.2. Squid tagging in the eld
Overall, a total of 93 squid were tagged in the eld using
VIE tags in unique patterns (Fig. 1) so that all squid could be
individually identied. The squid ranged in size from 1.8 g with
a mantle length of 19 mm to 149.4 g with a mantle length of
123 mm. The average weight of the squid was 40.2 g and the
average mantle length was 67 mm. No mortalities were observed
prior to re-release in the eld.
We recaptured 10 of the 93 squid tagged in the eld between
21 October and 20 November 2004 (Table 1). While one squid
was found over a nautical mile from where it was originally
tagged, the other nine squid that were recaptured were caught
at the location they were tagged. On three separate occasions
groups of similarly sized squid were found in the same part of
Achilles Bay, but none of the squid that were captured had been
previously tagged.
3.3. Growth rates
In captivity, the average growth rate for the squid was 1.66%
body weight per day, with a minimum of 1.4 and a maxi-
mum of 5.3. The average temperatures for the growth periods
ranged from 27.1
C down to 22.9
C.
4. Discussion
Growth rate data generated by the tag and recapture method
developed in this study compliments that of statolith and captive
culture work. While our current knowledge of squid growth rates
in the eld has been greatly enhanced by the use of statolith aging
methods, tagging studies help to address the unknowns that are
inherent in the statolith method of studying squid growth: mainly
the instantaneous growth rates of squid at specic sizes, and the
impact of small temperature changes on these growth rates.
Small changes in temperatures have been shown to have
a large effect on the exponential growth phase of hatchling
squid (Forsythe, 1993; Jackson et al., 1997; Forsythe et al.,
2001; Wood and ODor, 2000; Jackson and Moltschaniwskyj,
2002). Most statolith eld work is over relatively long time
intervals where temperature is likely to be extremely variable.
Researchers are starting to attack this problem of unknown tem-
peratures over long time periods by investigating the correlation
Table 1
Summary of data for the 10 squid tagged and recaptured in the eld
Squid number Location tagged Location recaptured Start weight (g) End weight (g) Average temperature (