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Fisheries Research 129–130 (2012) 1–7
Contents lists available at SciVerse ScienceDirect
Fisheries Research
journal homepage: www.elsevier .com/ locate / f ishres
Tag loss in the lobster Palinurus elephas (Fabricius, 1787) and implications forpopulation assessment with capture-mark-recapture methods
Lucía González-Vicente∗, David Díaz, Sandra Mallol, Raquel Goni
CentroOceanográficode Baleares, IEO, Muelle de Poniente s/n07015 Palma deMallorca, Spain
a r t i c l e i n f o
Article history:
Received 3 February 2012
Received in revised form 25 May 2012Accepted 26 May 2012
Keywords:
Capture-mark-recapture
Double tagging
Tag loss
Return rate bias
Palinurus elephas
a b s t r a c t
Assessments of animal populations by mark-recapture methods rest on the assumption that all marked
animals are recognizable, yet a variety of processes, such as molting in crustaceans, can lead to tag
shedding. In this paper we estimate the shedding rate of T-bar anchor tags in the European spiny lobster
Palinuruselephas from double tagging experiments conducted in the wild. During four annual consecutive
surveys (1999–2002) lobsters were caught, double-tagged and released. The evolution of the subsequent
double- and single-tag recaptures was traced during the following ten years (2000–2009). The estimated
instantaneous rate of tag loss was 6.8% year−1 in males and 5.0% year−1 in females tagged during inter-
molt. These tag-shedding rates are low compared to most published values for similar species. Data also
suggested that there is poorer tag retention when tagging immature individuals or during premolt. We
simulate and discuss the impact of undetected recaptures on the return rate and the implications for
population assessment in the particular case of P. elephas.
© 2012 Elsevier B.V. All rights reserved.
1. Introduction
Capture-mark-recapture (CMR) methods are widely used tostudy wild animal populations (Williams et al., 2002) since they
allow the estimation of population parameters such as abundance,
survival and recruitment. For marine exploited populations, CMR
techniques can be applied to determine the response of a popula-
tion to the introduction of marine reserves (or ‘marine protected
areas’, MPA), allowing the assessment the closure effectiveness on
population recovery and the estimation of movement and biomass
exportation outside the reserve (e.g. Afonso et al., 2008; Goni
et al., 2006, 2010). The accuracy of parameter estimates such as
abundance and survival obtained by CMR methods relies on the
correctness of assumptions, including the accurate identification
of tagged specimens, their traceability over time and complete tag
reporting (Lebreton et al., 1992; Seber, 1986).
However some recaptures could be missed as result of tagdetachment. Tagloss canoccur shortly after tagging as resultof tis-
sue rejection or insufficient anchoring of the tag (“type I tag loss”),
or tags maybe lost overtimeas resultof the normalaging, wear and
tear or molting (“typeII tagloss”) (Beverton and Holt, 1957; Parker,
1962). Hence, the use of CMR techniques for estimating population
parameters is hampered since this removal from the tagged pool is
indistinguishable from mortality or emigration.
∗ Corresponding author.
E-mail address: [email protected] (L. González-Vicente).
In this study we assess the rate of tag loss of T-bar anchor tags
in the European spiny lobster Palinurus elephas (Fabricius, 1787)
basedon double-taggingexperiments in the wild. The study has thedouble aimof quantifying therates of short-and long-term tagloss
andevaluatinghow tagloss affects returnratesand itsimplications
for population assessments, using P. elephas for illustration.
2. Materials and methods
2.1. Study area and species
The study area is situated in the Western Mediterranean and
includes the Columbretes islands marine reserve (hereafter “the
MPA”) and its adjacent fishing grounds. The MPA was established
in 1990 to protect species such as P. elephas, the most commer-
cially important spiny lobster species in the Northeast Atlantic and
Mediterranean.P. elephas is a long-lived, slow growing species that
in the study region matures at about 80mm carapace length (CL)
(Goni et al., 2003a) or 3–4years (Díaz, 2010). Due to its long life-
span and limited adult mobility, along with its high unit value and
excessive fishing effort, P. elephas populations are heavily over-
fished (Goni and Latrouite, 2005). For this reason, management
effectiveness of this valuable resource is crucial to its recovery and
sustainability. At present in the Spanish Mediterranean this fishery
is managed through a 6-month closed season, the prohibition of
retainingberried females, a minimum landing size (90mm CL) and
limited soak duration, in addition to area closures.
0165-7836/$ – seefrontmatter © 2012 Elsevier B.V. All rights reserved.
http://dx.doi.org/10.1016/j.fishres.2012.05.014
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2 L. González-Vicente et al./ Fisheries Research 129–130 (2012) 1–7
Table 1
Number of P. elephas double tagged and subsequently recaptured from 2000 to
2009 with twotags (double) andone tag (single) by sex. Only first time encounters
occurring earlier than thesixth year at liberty used in thestudy areshown.
Released
Female Male
1597 889
Recaptured
Female Male
648 468Double Single Double Single
45.0% 13.1% 31.7% 10.2%
Several previous studies comparing P. elephas inthe MPA and in
fished areas have shown that protection objectives are being met,
benefiting lobster in terms of abundance, mean size and reproduc-
tive potential (Díaz et al., 2011; Goni et al., 2001, 2003a,), as well
as increasing the local fishery yield as result of the outward flux of
above-average-sized lobsters (Goni et al., 2006, 2010).
2.2. Data collection
A total of 2486 lobsters (64.2% females) were captured using
trammel nets inside the MPA during annual experimental fishingsurveys conducted in September 1999 and June 2000, 2001 and
2002 (Table 1) (see Goni et al., 2003bf or details of survey methods).
All lobster caught were double tagged withcoded T-baranchortags
(Hallprint®, Australia) using an ordinary tagging gun. Tags were
inserted dorso-laterally on both sides of the midline in the first
abdominal inter-segmental membrane.
During the following ten years (2000–2009), tagged lobsters
were recaptured either inside the MPAin subsequentexperimental
fishing surveys conducted annuallyin June,or in the fishery outside
the MPA during the annual lobster fishing season (March–August).
Thanks to a close relationship with the fishermen operating in
the region and a reward offered for every recapture notified, the
detection and reporting rate from the fishermen involved in the
tag-recovery program was very high (Goni et al., 2010) and similarto that by scientific staff in the surveys inside the MPA.
2.3. Data analysis
2.3.1. Tag retentionmodel
Recapture data were aggregated by months at liberty, a time
interval (t ) at which fishing and natural mortality can be assumed
constant (Gulland, 1963). The rate of tag loss (L) and the prob-
ability of tag loss (P t ) were estimated applying a least squares
regression on the log-transformed proportion of tags retained
[2·N double/(2·N double +N single)] as a function of time (Eq. (1)).
P t = 1 − e(−Lt ) (1)
N double and N single represent the number of double and singletagged lobsters recaptured; 1- refers to the immediate tag loss
(“Type I loss”) (Bayliff and Mobrand, 1972) and L represents the
instantaneous rate of tag loss (“Type II loss”) (Bayliff and Mobrand,
1972;Chapmanet al., 1965). The number of reencountered animals
(N double +N single) at every time interval t was used as weights in the
regression.
Since the reduction of the number of reencounters over time
led to unrepresentative extreme values (0% or 100%) at the end of
the series, only the first-time encounters occurring earlier than the
sixth year at liberty were used in the study.
The probability of each of three theoretical tag-combinations
was calculated as follows: (1) P D = (1−P t )2, probability of a lob-
ster retaining two-tags; (2)P S = 2 ( 1−P t )·P t , probability of a lobster
retaining one-tag, and, (3)P 0 = (P t )2
, probability of no tagretention.
Subsequently and conditional on retention of at least one tag, the
expected probabilities of retaining either two tags (P RD) or one tag
(P RS) at recapture were computed (Eq. (2)):
P RD =
P D
1− P 0
and P RS =
P S
1 − P 0
(2)
Since this was a double tagging study, to assess the fit of the
tag loss model, the computed probabilities of being single-tagged
(P RS) at recapture (Eq. (2)) were compared with the observed per-
centage of single recaptures over time. Model selection was based
on the parameters of goodness of fit of the weighted least squares
regression and the examination of the residual plots. The level of
significance of all tests was˛= 0.05.
The following modeling assumptions were made (Hearn et al.,
1991): (1)thelossofanytagisindependentofthepositionofthetag
(left orright)andof the presenceof any other tag onthesamespec-
imen; (2)all the tags belongto therandom sample of tags available
for tagging and all of them have similar characteristics, and there-
fore similar chances to be retained; (3) the natural mortality and
catchability of an individual is independent on its tag-status; (4)
the tags are conspicuous and the same reporting reward is offered
for a single or double tagged animal, so they have equal reporting
rate; and (5) dispersal of individuals within the population is ran-
dom and independent of their tag-status. Therefore the proportionof double vs. single tagged animals reencountered is representa-
tive and independent of recapture location. Assumptions 1 and 2
relate to the independence and homogeneity of tags applied, while
assumptions 3–5 pertain to the representativeness of the tagged
population. Assumptions based on tag-status independence could
be violated when features associated with greater likelihood of tag
loss (e.g. molt state, size at tagging) are present, since lobsters lose
tags mainly through molting.
2.3.2. Factors affecting tag loss
The effect of factors time-at-liberty, size-at-tagging (log-
transformed), tagging season and sex on the probability of tag loss
was explored by multipleregression analysis.Tag losswas recorded
as a binomial response variable (retained or lost) with a logit linkfunction. The model which showed the lowest value of the Akaike
information criterion (AIC) was selected using a stepwise selection
procedure (Zuur et al., 2007).
Tag loss as a function of time-at-liberty was evaluated using
data for all mature lobsters (CL at tagging≥80mm CL) tagged and
released in June (surveys 2000–2002). Data for assessing the effect
of size at tagging on the probability of tag loss included all lobsters
tagged and released in June (surveys 2000–2002) grouped in two
size classes: small (immaturewith CL at tagging < 80mm) and large
(adults with CL at tagging≥80 mm). For assessing the effect of tag-
ging season on tag loss the dataset included recaptures grouped
according to month of tagging, June and September, the latter a
proxy for proximity to the males autumn molt.
2.3.3. Effects of ignoring tag loss on the return rate
To assess the effectof the tag loss on the return rate (the ratio of
individuals expected to be reencountered over time, relative to the
initial number of tagged animals released), we created a stochastic
model which allowed forecasting the reencounter reduction with
time. The following parameter values were used as input data: (i)
number of releasesR♀ =R♂ = 200 individuals per cohort; (ii) appar-
ent survivalPhi♂ = 0.6, Phi♀ = 0.8 (assuming lower values for males
based on their higher probability of mortality and emigration, Goni
et al., 2010), and (iii)recapture probability identical for bothgroups
p♀ = p♂ =0.5. We simulated 100 realizations from the log-normal
distribution of tag loss probabilities previously estimated (P t ±SD)
andestimated the mean proportion of tags lost over time. Then, the
expected return rate with time in the no-tag loss scenario (Phin · p)
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L. González-Vicente et al. / Fisheries Research 129–130 (2012) 1–7 3
Fig. 1. Proportion of single-tag recaptures for lobster tagged in September (autumn, gray bubbles) and June (summer, white bubbles). Recaptures accumulated in 30days-
at-liberty bins. Size of the bubble is proportional to thenumber of recaptures within each bin. Forsimplicity only single-tagged data (%) are represented. Females (top) and
males (bottom).
and the expected return rate affected by tag loss (Phin · p− P t ) were
computed, where Phin is the cumulative survival up to time (n).
Thus, theexpectedfrequencies of returnedanimalsover timeunder
both scenarios were reported.
3. Results
A total of 1116 first-time recaptures were obtained from 2000
to 2009, which represented 40.6% of the tagged females and 52.6%of the tagged males (Table 1). The majority of the recaptures took
place during the first three years-at-liberty, followed by a sharp
reductionof the numberof recoveries with time (Fig.1). From these
recaptures, 502(45.0%)femalesand 354(31.7%)males retainedtwo
tags.
Data exploration suggested model assumptions were generally
met, not finding statistical differences in the number of ani-
mals reencountered depending on tag-status, neither for males or
females (2-test testing the position of the tag p-value> 0.05, n.s.;
t -test testing for dispersal of single and double tagged animals in
the MPA and in the fished area p-value> 0.05, n.s.; QAICc values
indicated models accounting for recapture probability dependent
on tag-status performed worse and showed a lower statistical sup-
port).
3.1. Factors affecting tag loss
Multiple regression analyses indicated that the model includ-
ing factors time-at-liberty, size-at-tagging, tagging season, sex and
the interaction time:size-at-tagging explained the most deviance
and that these five attributes were significant ( p< 0.05) (Table 2).
Therefore, data were split and analyzed separately.
Modeling results for June tag releases indicated that the model
accounting for immediatetag loss 1- was not well supported, sug-
gesting that tags were adequately applied and retained in the shortterm for individuals tagged during the intermolt. However there
was a statistically significant long-term tag shedding L, reaching
5.0%year−1 in females and 6.8%year−1 in males (Table 3a). When
this constant rate was back-transformed into a probability P t (Eq.
(1)), the cumulative probability of losing a tag increased monoton-
ically. As a result, a population of P. elephas tagged with a single
tag would be expected to lose 50% of the tags after 10years in
males and after 14years in females (Fig. 2). No overall pattern was
found in the residuals when comparing the expected probability
of single-tagged recaptures (P RS) and the observed percentage of
single recaptures (% S Obs) (Fig. 3).
Size-at-tagging and its interaction with time were also signifi-
cant factors affecting tag loss (Table 2) and a higher proportion of
tags were shed in the group which was tagged when undersized.
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4 L. González-Vicente et al./ Fisheries Research 129–130 (2012) 1–7
Table 2
Deviance tableof multiple regressionof predictor variables for tag loss recorded as binomial response (logit link): time-at-liberty, size-at-tagging (log-transformed), tagging
season and sex. Statistically significant p-values in bold.
Estimate SE Deviance Residual deviance F p-Value
Intercept 21.251 19.719 1211.10
Time-at-liberty −11.865 5.463 177.86 1033.24 177.862 0.000
Size-at-tagging −5.708 4.262 21.2 1012.04 21.198 0.000
Factor (tagging season) −0.446 0.190 5.77 1006.27 5.767 0.016
Factor (sex) −11.611 27.534 5.82 1000.46 5.816 0.016
Time:size 2.832 1.181 9.38 991.08 9.383 0.002Time:factor (sex) 6.568 7.856 0.99 990.08 0.992 0.319
Size:factor (sex) 2.427 5.922 3.23 986.85 3.233 0.072
Time:size:factor (Sex) −1.366 1.690 0.64 986.21 0.639 0.424
Tagging season had a significant effect on the probability of tag
shedding (Table 2) in line with the observed pattern (Fig. 1). Long
term tag loss L was sensibly higher in male lobsters tagged in
September than in June (Table 3), although immediate tag loss 1-
again was not statistically different from zero.
3.2. Effects of ignoring tag loss on the return rate
Simulations mimicking the returns under the no-tag loss and
tag-loss scenarios showed a noticeable reduction of the expectedreencounters over time in the latter, more marked in males than in
females (Fig. 4). That is, after the first year at liberty, the number of
reencounters would be reduced by 22.1% and 12.2% for males and
females, respectively.
4. Discussion
The use of CMR methods to study animal populations relies
on the assumption that all tagged animals will be accurately rec-
ognized (Lebreton et al., 1992; Seber, 1986). Since tag shedding
constitutes a removal from the tagged pool that is indistinguish-
able from mortality or emigration, its knowledge is critical. In this
double-tagging experiment of the European spiny lobster P. ele-
phas in the wild, T-bar anchor tag shedding rates were amongthe lowest estimated for lobster species marked with external
tags. In spite of this relatively low rate, in an ordinary CMR study
where a single tag is used, half of the tagged population would
not be recognized after 10–14years (male and females, respec-
tively).
Published lobster tag-loss estimates vary depending not only
on the species, but also on tag type, size-at-tagging, tag-
ging season, duration of experiment, insertion location and
experimental conditions (wild, captivity) (Melville-Smith and
Chubb, 1997; Montgomery and Brett, 1996; Taylor and Hoenig,
1991; Trendall, 1989; Scarrat, 1970), hampering comparisons and
probably explaining the high variability of results (Table 4). Tag
loss rates estimated in this study (6.8–5.0% year−1 for males and
females, respectively) were lower than expected on the basis of
studies using comparable tag methods on other lobster species,
which can be as high as 87% year−1 in Homarus, 56% year−1 in Jasus
or 54%year−1 in Panulirus (Table 4). Nevertheless, the limited infor-
mation available for comparison suggests that T-bar anchor tagsinserted dorsally provide the highest retention in spiny lobsters, as
is the case of this study and that of Jasus verreauxi in tank experi-
ments of 8%year−1 (Table 4). Field experiments on Jasus edwardsii
with T-bar anchor tags inserted ventrally, which allegedly offers a
better protection against friction and entanglement, yielded a sim-
ilar shedding rate for females (6.1% year−1) but a higher rate for
males (13.4% year−1) (Frusher et al., 2008).
Due to theirhighergrowthrates andconsequent moltfrequency
(Díaz, 2010), undersized specimens of P. elephas were more likely
to lose tags than adults, being this probability of tag loss reduced
as growth rate slows down with time, as inferred from the mul-
tiple regression analyses. Similarly, males which grow faster than
females showed poorer tag retentionthan females. This is expected
because the main cause of crustacean tag shedding is molting(Davis, 1978; Restrepo and Hoening, 1988), although other causes
have been identified, such as removal by congeners (Krouse and
Nutting, 1990) or predators(Rowe and Haedrich, 2001), and entan-
glement with surfaces (Ennis, 1986). For the same reason, tags
inserted duringpre-molt, such as those of males tagged in autumn,
showed poorer tag retention than tags inserted in summer during
post-molt (adult females) or inter-molt (males). Similarly, in other
lobster species, tagging duringthe late pre-molt hasbeen observed
Table 3
Estimatesof short-term (1-) andlong-term (L) tag loss(± SE)by sexin lobsters≥80mm CL attagging;(a) tagged inJuneand(b) inSeptember.The modelswithlowest AIC
score are highlighted in bold.
Sex Model Tag loss (year−1 ± SE) p-Value AIC
(a)
Female p= 1-e−Lt 1- 0.021 ± 0.04 0.966
L 0.040 ± 0.01 0.000 −65.48
p= 1-e−Lt L 0.050 ± 0.01 0.000 −67.48
Male p= 1-e−Lt 1- 0.021 ± 0.04 0.637
L 0.072 ± 0.01 0.000 −55.94
p= 1-e−Lt L 0.068 ± 0.01 0.000 −57.71
Female p= 1-e−Lt 1- 0.021 ± 0.036 0.567
L 0.040 ± 0.011 0.001 −19.67
p= 1-e−Lt L 0.045 ± 0.007 0.000 −21.32
Male p= 1-e−Lt 1- 0.021 ± 0.036 0.540
L 0.081 ± 0.019 0.000 5.11
p= 1-e−Lt L 0.090 ± 0.013 0.000 3.52
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L. González-Vicente et al. / Fisheries Research 129–130 (2012) 1–7 5
Table 4
Publishedestimates of lobster tag loss rates. Tag loss estimatesfrom short term studies (<1year) were scaled up to oneyear at liberty. (*)Juveniles.
Species Reference Tag type Tag loss (year−1) Experiment
Palinurus elephas This study, 2012 T-bar anchor tag 6.8%♂ Field
5.0% ♀
Panulirus cygnus Chittleborough (1974) Sphyrion tag 27% Field
Rock lobster tag 30%
Panulirus argus (*) Davis (1978) Floy FD-68B tag 45% Field
Panulirus marginatus O‘Malley (2008) Streamer tag 54% Field
Jasus novaehollandiae Winstanley (1976) Dart tag 56%♂ Field41% ♀
Jasus verreauxi Montgomery and Brett (1996) Toggle tag 6% Laboratory
T-bar anchor tag 8%
Dart tag 8%
Jasus edwardsii Frusher et al. (2008) T-bar anchor tag 13.4%♂ Field
6.1% ♀
Homarus americanus Cooper (1970) Sphyrion tag 22% Laboratory
Scarrat (1970) 32% Laboratory
Fogarty et al. (1980) 87% Field
Ennis (1986) 36% Molted Field
24% Non molted
Homarus gammarus Bennett and Lovewell (1983) Sphyrion tag 36% Laboratory
to increase the risk of tag loss (Comeau and Mallet, 2003; Moriyasu
et al., 1995).
Despite the comparably low rate of tag shedding obtained forP. elephas in this study, this loss has a significant impact on the
return rate. To that proportion of the population which is missed
with time as result of shedding itself, we have to add the effect of
the decreasing return rate with time, given that the detection of
those tagged animals is conditional on their survival up to time t
and recaptureat that particular event.This wouldbe more acute for
those strata with higher growth rates i.e. differential growth rate
Fig.2. Expected cumulativeprobabilityand 95% confidenceintervalsof tag loss(P t )
for lobsters taggedin Junemeasuring≥80mm CLat tagging.Females() and males
().
related to sex, size or molt stage.Our simulations forthe returnrate
based on constant parameters and number of single-tag releases,
forecast the failure to detect recaptured animals beyond the fourthyear at liberty as result of the impact of tag loss. Because our field
data come from a double tagging experiment, we obtained reen-
countersfar beyond this time limit (>10 years). Certainlythe steady
state simulated here is used just for illustrating the importance of
accounting for tag loss, since in fact we suspect that several fac-
tors, such as sex-related differences in movement probability or
catchability, are jointly influencing the reencounter pattern of P.
elephas in the study area in the long term (González-Vicente et al.,
2011). Therefore, P. elephas true survival may be generally higher
0%
20%
40%
60%
80%
100%
876543210Years at liberty
% S Obs PRS
0%
20%
40%
60%
80%
100%
876543210
Years at liberty
% S Obs PRS
Fig. 3. Expected frequency of single-tag lobsters at recapture (P SR ) versus observed
proportion (June releases≥80mm CL). Females () and males ().
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6 L. González-Vicente et al./ Fisheries Research 129–130 (2012) 1–7
0
20
40
60
80
100
876543210
A b
s o l u t e f r e q u e n c y
Years at liberty
Expected returns
with tag lossExpected returns
0
20
40
60
80
100
876543210
A b
s o l u t e f r e q u e n c y
Years at liberty
Expected returns
with tag lossExpected returns
Fig. 4. Expected number of returned individualsin scenariosunaffected (solid line)
and affected (dashed line) by tag loss. Females (top) and males (bottom).
than the apparent survival simulated here, and so are the return
rates, leading to this long term reencounter time series.
Although simulation studies have previously shown that the
‘type I tagloss’ does not lead to importantbiason survival estimates
(McGarvey et al., 2009), when studying the parameters involved
in the population dynamics, unaccounted ‘type II tag loss’ wouldlead to erroneous estimates of recapture and survival. Moreover,
recaptured but unidentified animals would be misinterpreted as
immigrants, biasing upwards recruitment estimates. As a result of
this, inadequate management measures to protect the population
could be enforced, while the originating cause of animal loss (tag
detachment) would remain unidentified.
Tag loss has been shown to bias abundance and survival esti-
mates in field studies (McDonald et al., 2003; Seber and Felton,
1981), a problem which is commonly remedied by adjusting sur-
vival estimates a posteriori (Pollock, 1981). Also, when population
parameters are affected by complex time-, location- or state-
dependency factors, the correction for tag loss can be incorporated
into the likelihood to obtain specific estimates for each category
and time event. For instance, Oosthuizen et al. (2010) determinedcohort- and site-specific tag loss estimates for southern elephant
seal (Mirounga leonina), in order to avoid the heterogeneity among
the different age and location strata. Similarly, it is possible to
include sampling methodology aspects into the likelihood estima-
tion. For instance, Juillet et al. (2011) combined information from
liverecaptures, deadrecoveries, and single anddouble tagged spec-
imens of greater snow goose (Chen caerulescens) to estimate tag
loss.
In conclusion, the service life of T-bar tags for this species
spans beyond 15years, validating this tagging method and provid-
ing a reasonably high retention in the long term. In view of their
long-termretention, endurance against ecdysis and easy detection,
tagging dorsally with T-bar tags anchor is a suitable method for
field studies of lobsters such as P. elephas andpresumably of others
with similar molt rates. For maximizing survival and tag retention,
tagging is advised during the inter-molt period.
Acknowledgements
The authors are grateful to the skippers and crews of the fishing
vessels operatingin thestudy areafor their invaluable collaboration
in recapture notification over the years. We also thank A. Quet-
glas and O. Renones and all the collaborators who contributed tothe data collection in the early years of the project. We would like
to thank to Dr. McGarvey and one anonymous referee for his/her
constructive comments. This work was funded by the Spanish Sec-
retaría General del Mar by means of a grant for the LANGOSTA &
ERICOL projects (1997–2012). L.G.V. was supported by a FPI-IEO
scholarship (2008–2012).
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