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Integrating Anthropogenic and Climatic Factors in the Assessment of the

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 International Journal of Marine Science 2013, Vol.3, No.6, 36-45
http://ijms.sophiapublisher.com

Research Report Open Access

Integrating Anthropogenic and Climatic Factors in the Assessment of the
Caribbean Spiny Lobster (Panulirus argus) in Cuba: Implications for
Fishery Management
Rafael Puga , Roberto Piñeiro , Romina Alzugaray , Lisset Susana Cobas , María Estela de León , Ofelia
Morales
Fisheries Research Centre, 5th. Ave. and 246, Barlovento, Playa, Havana, Cuba
Corresponding author email: rpuga@cip.telemar.cu; Authors
International Journal of Marine Science, 2013, Vol.3, No.6 doi: 10.5376/ijms.2013.03.0006
Received: 18 Dec., 2012
Accepted: 09 Jan., 2013
Published: 24 Jan., 2013
Copyright: © 2013 Puga et al., This is an open access article published under the terms of the Creative Commons Attribution License, which permits unrestricted
use, distribution, and reproduction in any medium, provided the original work is properly cited.
Preferred citation for this article:
Puga et al., 2013, Integrating Anthropogenic and Climatic Factors in the Assessment of the Caribbean Spiny Lobster (Panulirus argus) in Cuba: Implications
for Fishery Management, International Journal of Marine Science, Vol.3, No.6 36-45 (doi: 10.5376/ijms. 2013.03.0006)

Abstract The Caribbean spiny lobster Panulirus argus, the most valuable Cuban fishery resource, is managed with a set of input
and biological controls. The aim of this study was to integrate two indices that relates water and nutrients supply and tropical
cyclones activity in the stock assessment, through simultaneous estimation of the parameters of a spawning stock–recruitment
function in a statistical catch-at-age analysis. The population dynamics model allowed estimating key Reference Points for
management at fixed levels of fishing mortality rate and environmental conditions. The results indicated that the reduction of
recruitment and catches in the Cuban spiny lobster fishery could be a result of synergistic cumulative effects because of the
anthropogenic reduction of nutrients supplies and the increase of the potential destructiveness of tropical cyclones since 1994, mainly
from 2001 on. Although the degradation of the coastal habitat quality in Cuba was apparent and perhaps unavoidable, the assessment
of the impact allowed implementing management actions for the spiny lobster fishery sustainability. The implementation of annual
TAC depending on the stock status, have maintained the fishery around the more conservative Reference Points F40% and F0.1 during
the current unfavorable environment period.
Keywords Caribbean spiny lobster (Panulirus argus); Stock assessment; Fishery management; Catch-at-age analysis; Environmental
effects; Cuba

Background length (CL), a maximum landing size of 140 mm CL

Since the Caribbean spiny lobster Panulirus argus is
the most valuable fishery resource in Cuba, many
studies have been made on the assessment of the
population dynamics of the species, to enable the
fishery management to achieve sustainable
exploitation (Puga et al., 1991; Cruz et al., 1995; Puga
et al., 1996; de León et al., 2001; Puga et al., 2005;
González-Yáñez et al., 2006; Morales and Puga, 2008;
Alzugaray and Puga, 2010).
for females, a prohibition on taking berried females
and a Total Allowable Catch (TAC) since 2008 which
is set annually depending on the previous stock
assessment. The status of the stock is evaluated by the
stock assessment group and the management system is
reviewed annually in an adaptive framework by an
advisory board with the stakeholders. Any changes in
the management regulations are declared as state laws
which are enforced by a fisheries inspection unit.

The fishery is managed with input (effort restrictions),
output (catch quotas) and biological controls
(protection of reproduction activity and recruits),
which include: a state property limited entry regime,
territorial rights to fishery enterprises, gear restrictions,
a 145-days closed season from February to June,
permanently closed areas to protect juveniles and
spawners, a minimum landing size of 76 mm carapace
It was recognized that, despite good management and
control, the spiny lobster populations in Cuba, Mexico
and the United States of America were showing signs
of declining, because a number of factors outside the
fishing sector, such as the increased frequency and
intensity of hurricanes and the human development in
coastal areas, were negatively impacting the lobster
habitat (FAO, 2007; Ehrhardt et al., 2011).

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The analysis of causes and consequences of change in
diverse coastal areas worldwide show similar patterns:
multiple human impacts have destroyed seagrass and
wetland habitats and depleted many important species
(Lotze et al., 2006). The increasing anthropogenic
pressures and the susceptibility to climate change have
led to a gradual global loss of seagrass beds,
mangrove forests, salt marshes, and coral reefs
(Duarte et al., 2008). The mechanisms of these losses
are multiple, including coastal land loss,
eutrophication and hypoxia, hydrologic changes in
coastal wetlands made for flood control, and climate
change (Cowan, 2009).
There is compelling evidence of the effects of dams
on limiting the flow of nutrients and the quantity of
water needed for maintaining downstream ecology
(Berkamp et al., 2000; Humborg et al., 2000; Baisre
and Arboleya, 2006). On the other hand, Oczkowski et
al (2009) found that the fishery production can be
stimulated by fertilizer and sewage runoff that
increases nutrient loading in an oligotrophic sea.
Inorganic fertilizer consumption by agricultural
activities is the largest single source of nitrogen and
probably of other nutrients to Cuban landscape (Baisre,
2006).
The increase in the frequency of hurricane activity
may cause longer-term biogeochemical and trophic
changes in estuarine and coastal habitats (Paerl et al.,
2001). Numerous studies have addressed the impact of
hurricanes on marine communities such as mangroves
forests (Ward and Smith, 2007), coral reefs (Gardner
et al., 2005), and seagrass beds (Fourqurean and
Rutten, 2004), all of them important habitats of P.
argus (Butler et al., 2006).
An adequate shallow-water nursery habitat is crucial
for successful postlarval settlement and the survival
and growth of juvenile lobsters that recruit to fisheries
(Butler et al., 2006). Therefore the aim of this article
is to evaluate anthropogenic and climate factors that
could impact the Cuban spiny lobster stock, through
their possible direct or indirect effects on the
recruitment success. Some previous results examined
the anthropogenic effect on the supply of water and
nutrients from the terrestrial system to the Cuban
marine coastal area (Baisre, 2006; Baisre and
Arboleya, 2006) and the increase of the potential
destructiveness of tropical cyclones in the North
Atlantic (Emanuel, 2005). Therefore, we integrated
two indices related to water and nutrients supply and
tropical cyclones activity in the assessment of Cuban
spiny lobster dynamics.
1 Results
The log-likelihood resulting from the fitting of the
simplified model and the proposed environmental
model (Table 1) show a best fitting for the proposed
model, because it provided lower values for the
components (mainly for recruitment) and for the total
log-likelihood used as objective function.
Table 1 The log-likelihood components resulting from the fitting of
the simplified model and the proposed environmental model
Variables Simplified model Environmental model
Rt 0.037 0.019
Ca,t 12.992 12.185
Yt 0.050 0.032
Ft 0.017 0.016
Total 13.096 12.252
The model fit is adequate in that the model is
generally able to follow the overall trends of the
observed catch, although the model overestimates the
catch from 1968~1973 and underestimates the catch
during the period 1978~1991 (Figure 1). The fishery
was undeveloped before 1955, in developing phase till
1965 and stable around 9 100 tones between 1965 and
1976. After the maximum catches around 11 200 tones
between 1982 and 1989, the fishery decreased to another
relatively stable catches period around 9 410 tones in
1991~1999. As consequence, there is no a defined
general trend between 1965 and 1999. Following a
declining trend with high inter-annual variability
since 1999, the observed catches are quite stable
around 4 870 tones during 2005~2011.
Figure 1 Observed and predicted catch in weight in the Cuban
spiny lobster fishery

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The fit of the model to the fully recruited fishing
mortality rate is very good (Figure 2), which is shown
by the significant relationship in the linear regression
between the model predicted fishing mortality rate and
the fishing effort in boat-days Ft = qft (R2 = 0.95,
n = 40). The predicted F increasing trend till the
1970’s reflects the fishery developing phase. After a
gradual increase from 1993 to 1999, the fishing effort
and F sharply declined and stabilized at low levels
during the current period 2008~2011.
Figure 2 Observed and predicted fully recruited fishing
mortality rate and fishing effort in the Cuban spiny lobster fishery
The recruitment stayed at average level between 1955
and 1965, and increased above average during
1966-1994 (Figure 3), coincident with favorable
environmental conditions given by high level of AEI
between 1965 and 1991 and low level of PDI between
1976 and 1993. Despite the unfavorable high PDI
values between 1966 and 1973, it seems the
recruitment was enhanced by the highest AEI level
during this period.
Figure 3 Time trajectories of anomalies of lobster recruitment
(R) in year t, Anthropogenic Effect Index (AEI) and Power
Dissipation Index (PDI) in year t-1
Note: Horizontal broken line represents the average level for
the time series
38
AEI remained below average level since 1993, and
PDI show a sustained increasing trend from 1994 with
values above average level since 2001, which
determine unfavorable cumulative environmental
conditions. As consequence, the recruitment fell below
average level since 1996 to the lowest values during
2006~2011.
The Spawning Potential Ratio as function of F in
Figure 4 shows F0.1 as the most conservative Target
Reference Point (TRP), and FMSY as a Limit Reference
Point (LRP) that should not be exceeded to avoid
overfishing.
Figure 4 Equilibrium curve of Spawning Potential Ratio (SPR)
as function of fishing mortality rate and Reference Points F0.1,
F40% and FMSY
There were considerable differences in stock
production for key Reference Points associated with
variations in environmental conditions over their
ranges (Figure 5). Observed and estimated equilibrium
catches were considerably higher during periods of
favorable conditions. A summary of results of interest
for fishery management is presented in Table 2.
Figure 5 Equilibrium curves of catch in weight (Y) as function
of fishing mortality rate (F) for three levels of environmental
conditions, 5-year moving average of observed Y versus F from
1954 to 2011 and Reference Points F0.1, F40% and FMSY
Note: The arrows indicate the trajectories of observed Y versus
F and the numbers represent the last two numbers of some years
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Table 2 Estimates for quantities of interest to the management of Cuban spiny lobster resource with three levels of environmental
conditions
f SPR
Reference point F
(boat-days)
F0.1 0.162 20 193
F40% 0.184 22 878
FMSY 0.383 47 763
The observed catch trajectory followed the average
equilibrium curve during the fishery developing phase
(Figure 5). Between 1971 and 1976 the FMSY LRP was
lightly exceeded, but the catches increased later even
for lesser F values, coincident with the effect of
several years with favorable environment.
After catches peaked in 1984~1987, the catches
declined because an F decreasing trend, and a shift
toward average environmental conditions, determined
by a gradual drop in AEI level. Despite the F
increasing in 1993-1997, the catches remained
relatively stable during 1991-1999 and definitively
descended with F, as a result of low recruitment due
to an unfavorable environment shift determined by
low AEI since 1993 and strengthened by PDI
increasing since 2001. The fishery was close to the
F40% TRP during 2005~2007 and virtually on the more
conservative F0.1 TRP since the implementation of
TAC in 2008.
2 Discussion
In the Cuban P. argus fishery, the general decreasing
trends in fishing effort and F since 1972 and mainly
since 1999, made it difficult to associate a senescent
fishery phase with the increase of the exploitation
intensity. The fishing effort and F remained high in
1972~1984 and declined to low values in 1990~1993,
because of the partial collapse of the Cuban economy
due to the loss of imports from the Soviet Union,
which led to a reduction of imported fuel. Therefore,
the current decreasing catches are more likely due to a
reduction in the recruitment and its consequence on
the lobster population dynamics, because of the effect
of fishery independent factors. The Cuban spiny
lobster fishery, like other Caribbean ones, is
recruitment driven and catches mostly correspond to
fluctuations in recruitment. The effects on recruitment
of environmental conditions independent of fishery
exploitation, are impacting the recruitment to the
Catch in weight (tones)
(%) Favorable Average Unfavorable
44 9 428 7 409 4 694
40 9 892 7 774 4 925
17 11 266 8 853 5 609
fishery two to three years later in Cuba and Florida,
where regulations maintain controlled fisheries
(Ehrhardt et al., 2011). Anthropogenic and environmental
effects on the spiny lobster nursery habitat may be
playing a role in the decline of the recruitment and the
subsequent catches.
A significant feature with the Florida and
Nicaragua-Honduras lobster stocks is the shift in
recruitment success, associated with Caribbean mean
sea level shifts (Ehrhardt et al., 2011). Like the high
PDI in Cuba, a high sea level produces low
recruitment in Florida and Nicaragua-Honduras and
that such environmental effects have been stronger
during the 2000’s than before. A study on surface
wind variability showed that tropical storms and
hurricanes increase the Ekman pumping and transport
in southern Cuban waters, which could negatively
impact the success of postlarval settlement
(Perez-Santos et al., 2010). The frequency of storms
and westerly winds also affects the oceanographic
conditions in the lower coast of Western Australia and
hence the oceanic larval phase of the western rock
lobster P. cygnus (Caputi and Brown, 1993).
It is widely recognized the combined effect of
anthropogenic and climatic factors on the coastal
habitat degradation can have on the productivity of
marine ecosystems and fisheries (Gardner et al., 2005;
Lotze et al., 2006; Duarte et al., 2008; Cowan, 2009;
Sumaila et al., 2011). Coral reefs, mangrove forests
and seagrass beds are important components of the
spiny lobster habitat. The decline of corals on tropical
reefs is usually attributed to a combination of
anthropogenic and natural factors. The contribution of
hurricanes to Caribbean coral cover decline over the
past two decades was quantified to be 17%, on
average, in the year following the hurricane impact
(Gardner et al., 2005). Seagrass beds with little to
moderate sediment deposition may recover within 1 year,
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while areas with substantial erosion may have
recovered very little during the 3 years after the storm
(Fourqurean and Rutten, 2004). Because of the
complex interactions among a variety of physical
factors, the damages caused by hurricanes may have
unexpected consequences that spill throughout the
species dependent on mangrove forests ecosystem
(Ward and Smith, 2007).
On the other hand, human activities affect nutrient
fluxes to the aquatic communities, trough their effects
on the biogeochemical cycling. The nutrient supply
from land to the coastal area is strongly influenced by
the nature of catchment areas that supply the water,
via rivers, to the sea. Concentrations of phosphorus or
nitrogen may limit productivity in these areas, but
additional anthropogenic effects on the chemistry of
river water have special significance for planktonic
communities. The export of silicate, an essential
component of the cells of planktonic diatoms, are
dramatically lower in the dammed rivers (Begon et al.,
2006).
The social-economics benefits of dams in Cuba are
obvious, and include the modulation of river flow
between rainy and dry periods, the production of
hydroelectric power, and water supply for human
consumption and agriculture irrigation, however, the
ecological consequence for the coastal area should not
be ignored. Since Cuba is located in the oligotrophic
Caribbean Sea (Richards and Bohnsack 1990),
without significant coastal upwelling and tidal mixing,
the river discharges are the main source of nutrients
supporting the productivity of marine coastal fisheries
(Baisre and Arboleya, 2006).
The dam capacity in Cuba increased from 1.1 million m3
in 1968 to 9.7 million m3 in 1991, which represents
approximately 30% of the total volume of Cuban
hydrologic resources (http://www.hidro.cu/). The
fertilizer consumption in Cuban agriculture activity
increased from 229 thousands tones in 1972 to around
602 thousands tones in 1981~1990, and decreased
later due to the drastic shortage of funds for the
purchase of inorganic fertilizer since 1990, therefore,
nitrogen and other nutrients inputs were reduced
(Baisre, 2006). The consequence of both factors, the
anthropogenic effect on nutrient and water supplies to
the coastal area is reflected in the AEI time series,
40
with high levels in 1965~1991 and low values since
1993, coincident with the periods of higher and lower
recruitment and catches. Inorganic fertilizer was the
largest single source of reactive nitrogen to the Cuban
marine environment (Baisre, 2006). In a similar way,
river damming and fertilizer consumption in
agriculture activity played a significant role in the
Mediterranean fishery. Oczkowski et al (2009) relate
the collapse of the coastal Mediterranean fishery off
the Nile River delta, with the completion of the Aswan
High Dam in 1965. The fishery recovered
dramatically since the mid-1980s, coincident with
large increases in fertilizer application in the Egyptian
agriculture. They use stable isotopes of nitrogen to
confirm that more than 60% of the fishery production
was from primary production stimulated by nutrients
from fertilizer and sewage runoff.
Humborg et al (2000) found far-reaching effects in the
Baltic Sea and the Black Sea, of disturbance on the
coastal biogeochemical cycles and food web structure,
because dams in river basins had substantially reduced
silicon loads to the coastal area.
Arias-Schreiber et al (2008) found that adverse
changes have occurred in the benthic assemblages in
the areas of south west Cuba with a great reduction in
species diversity and seagrass coverage. Also, the prey
abundance of juvenile lobsters and shrimps has been
significantly reduced in the nursery areas to less than
20% compared to the 1980’s (Lopeztegui-Castillo and
Capetillo-Piñar, 2008; Cantón-Machín et al., 2010), a
fact that could be constraining the success of the
recruitment and therefore the subsequent catches of
both species.
Baisre and Arboleya (2006) concluded that two factors
have acted synergistically, leading to the decline of
marine coastal fisheries since 1990 in Cuba: a drastic
reduction in nutrient inputs from land sources due to
reduced fertilizer use, and the trapping of nutrients by
river damming. Our results, based on the effects of
AEI and PDI on the lobster recruitment success,
indicate that the reduction of recruitment and catches
in the Cuban spiny lobster fishery could be a result of
synergistic cumulative effects because of the
anthropogenic reduction of fresh water and nutrients
supplies to the coastal zone (AEI) and the increase of
the potential destructiveness of tropical cyclones (PDI)
 International Journal of Marine Science 2013, Vol.3, No.6, 36-45
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since 1994, mainly from 2001 on. The increase in
tropical cyclone power in the North Atlantic was
previously presented by Emanuel (2005) with this
index of the potential destructiveness of hurricanes
(PDI) based on the total dissipation of power,
integrated over the lifetime of the cyclone.
Lotze et al (2006) indicated that synergistic effects
related with habitat loss and exploitation have been
significant for species depletions and extinctions.
Cumulative impacts were even more important for
recovery, with significant lag times. Cowan (2009)
pointed out that detecting important impacts of habitat
loss on fishery resources should be relatively easy,
nevertheless, a direct cause-effect relationship
between habitat loss and fisheries productivity has
remained elusive, because our inability to detect
proper relationships between the degradation of
coastal habitats, and the animals that are dependent
upon these habitats. That is why cumulative impacts
could likely drive a fishery ecosystem to a precipitous
and perhaps irreversible decline. Barange et al (2010)
highlighted the importance of develop coupled
modeling frameworks to explore the synergistic
effects between climate change and human activity on
marine ecosystems.
Using inappropriate models or data could lead to
misleading or contradictory results. Based on a
generalized fisheries simulation model for
data-limited situations, Chavez (2001) concluded that
the P. argus Cuban fishery was underexploited and
recommended increasing twice the current F to obtain
14 453 tones at MSY. Later on, Chavez (2009)
concluded that the fishery was overexploited between
1991 and 2004, and recommended a 30% F reduction
to achieve 11 100 tones at MSY.
Our results do not support overfishing as responsible
for the recruitment decline, as Cruz et al (2001)
suggested based on 10 years analysis of relative
abundance in early life-cycle stages.
Environmental effects on the recruitment of Cuban P.
argus stock are evident, in spite of strict fishery
regulations, low catches has not been avoided. The
improvement of the management system, particularly
the reduction of fishing effort and F, has failed to
reverse the decline in catches, which have remained
below 8 000 tones since year 2000 and below 6 000 tones
since 2005. The implementation of annual TAC
41
depending on the stock status, have maintained the
fishery around the more conservative Reference Points
F40% and F0.1 during the current unfavorable
environment period. The reduced effort in response to
the lower recruitment has provided greater protection
to the breeding stock as recruitment overfishing is
often caused by a combination of heavy fishing
pressure being maintained on a series of low
recruitment year-classes.
In similar way, after the collapse of the spiny lobster P.
marginatus because of the regime shift in the
Hawaiian Archipelago, the fishing effort was greatly
reduced in subsequent years without recovery. Even
areas which never open to lobster fishing, experienced
the same collapse in recruitment and subsequent
decline in the spiny lobster population (Polovina,
2005). Therefore, it was decided that overfishing was
not the cause for the decline, so the managers chose to
continue the fishery at exploitation rates with low risk
of overfishing, based on population dynamics model
results which include the environmental effects.
There is evidence that the targets and limits set for the
Cuban spiny lobster management are sensitive to
environmental changes. As the resilience may be
reduced by unfavorable shifts in the environment, the
management rules should take account of effects of
past changes and allow for the consequences of future
changes. Caputi et al (2010) also highlighted the
importance of assessing the impact of environmental
climate changes on stock assessment and management
of the fishery of the western rock lobster in Western
Australia.
Although the degradation of the coastal habitat quality
in Cuba is apparent and perhaps unavoidable, the
assessment of the impact of the water and nutrients
supplies and the tropical cyclones, allows
implementing management actions for the spiny
lobster fishery sustainability.
3 Data and Methods
3.1 Data source and processing
Annual catches in tones from 1954 to 2011, fishing
effort in boat-days from 1972 to 2011 and commercial
weight categories from 1979 to 2011 were obtained
from official compulsory records of the fishing
enterprises of the Cuban Ministry of Food.
The commercial weight categories were converted
into length and age structures of total catch in
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numbers, using the length-weight relationship from
Cruz et al (1981) W = 0.00243CL2.764 , where W is the
lobster weight (g) and CL is the carapace length (mm),
and the growth parameters for the von Bertalanffy
relationship obtained by León et al (2005) L∞ (Males)
= 185 mm CL, L∞ (Females) = 155 mm CL, K (Males)
= 0.23, K (Females) = 0.19, t0 (Males) = 0.44, t0
(Females) = 0.37
The natural mortality rate M = 0.34 derived from Cruz
et al (1981), has been used in previous stock
assessment analysis (Puga et al., 1991; Puga et al.,
1996; Puga et al., 2005; Morales and Puga, 2008;
Alzugaray and Puga, 2010).
The two indices included in the stock assessment are:
The Anthropogenic Effect Index (AEI) as an index of
the effect of human activities on the supply of water
and nutrients from the terrestrial system to the aquatic
one. AEI = Fert/DamCap, where Fert is the amount of
inorganic fertilizer used in Cuban agriculture
(http://faostat.fao.org/site/575/default.aspx#ancor) and
DamCap is the water capacity in Cuban dams
(http://www.hidro.cu/).
The annual activity of tropical cyclones on Cuban
marine area was quantified by the Power Dissipation
Index, PDI = ∑V3 from Emanuel (2005), where V is
the maximum sustained wind speed measured each 6
hours while the cyclones remain in the area
(http://weather.unisys.com/hurricane/atlantic/index.php).
The time series data of Fert, DamCap and PDI were
ranked from the lowest (number one) to the largest
value, with the method of Chen et al (2007) to obtain
non-parametric environmental indices. AEI was then
calculated for each year and a simple exponential
smoother in a similar way that Emanuel (2005)
procedure, was applied to both non-parametric indices
(AEI and PDI), to take into account the cumulative
effect of both factors on the habitat, and to reduce the
effect of interannual variability.
High AEI and low PDI indicate favorable
environmental conditions for lobster recruitment,
while low AEI and high PDI indicate unfavorable
environments compared with their average levels.
Thus, AEI and PDI are related to both the relative
recruitment success and environmental conditions, and
we used it as relative indices together with the
spawning stock to estimate annual recruitment
42
numbers in the stock-recruitment relationship, in
similar way to the assessment of the Pacific cod
(Sinclair and Crawford, 2005).
3.2 Population dynamics model
The statistical catch-at-age analysis (SCAA) as
discussed by Haddon (2001), Maunder and Watters
(2003), and Butterworth and Rademeyer (2008), was
applied to fit the available data to a population
dynamics model. This method allows the
simultaneous estimation of the parameters of a
spawning stock–recruitment function and allows the
inclusion of relationships between environmental
variables and processes in the population dynamics
(like recruitment) to improve estimation of model
parameters and prediction of population variables.
SCAA, unlike Virtual Population Analysis (VPA), does
not require that catch-at-age data be available for each
year covered by the assessment.
The master equations of the SCAA used to estimate
the parameters in the population dynamics model
implemented in Microsoft Excel are:
For recruitment:
Rt + 1 =
St
(ln(δAEIt ) + ln(ϕPDIt ))
(α + βSt )
where St is the population egg production in year t,
AEIt and PDIt are the Anthropogenic Effect Index and
the Power Dissipation Index in year t, Rt+1 is the
number of one year old lobsters in year t+1, and α, β,
δ, φ, are parameters in the modified Beverton-Holt
recruitment model. St was estimated in the model with
the surviving numbers at age by year and the egg
production by female by age (Table 3) derived from
Cruz and Bertelsen (2008) maturity and fecundity
relationships.
Table 3 Egg production by female by age derived from Cruz
and Bertelsen (2008)
Age Number of eggs
1 0
2 68
3 3 043
4 72 779
5 236 000
6 391 314
7 523 042
8 640 416
9+ 942 680
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− (M + Fa ,t )
For survived number: N a +1,t +1 = N a ,t e
For fishery catch in number:
Ca,t =
Fa,t
M + Fa,t
(
N a,t 1 − e
−(M + Fa ,t )
) The log-likelihood components were estimated for the
where Na,t and Na+1,t+1 are the surviving numbers at
ages a and a+1 in years t and t+1, M is the natural
mortality rate and Fa,t is the fishing mortality rate at
age a in year t.
For fishery catch in weight: Yt = ∑C
where Wa is the lobster weight at age a.
For fishing mortality: Fa ,t = sa Ft and Ft = qft
where Ft is the fully recruited fishing mortality, sa is
the age-specific selectivity, q is the catchability, and ft
is the annual fishing effort in boat-days. In the
relationships between the catch in weight and F and
between the Spawning Potential Ratio and F, and for
Reference Points estimate, we use the average value
of F over all the ages groups a.
1 1 − λ λ eφλ (γ −a )
For selectivity: sa = ( ) (
1− λ λ 1 + eφ ( γ − a )
where a is the age and λ, ϕ, γ, are parameters in the
Thompson (1994) exponential-logistic model, which
can represent flat-topped or dome-shaped selectivity
patterns and has the useful property that the maximum
selectivity value is unity.
We fitted the population model using a
likelihood-based approach where we used the
Microsoft Excel’s solver to find parameter values that
minimized the objective function. The objective
function was the sum of the log-likelihood
components associated with observed and predicted
variables.
ln L =
∑ (ln I t − ln Iˆt ) 2
2nCV
where It is the observed variable, Iˆt is the predicted
variable and n is the number of observation years. The
coefficient of variation CV was arbitrarily set to an
standard value of 0.2 because several authors
considered values around 0.2 to be reasonable for the
process errors in the same variables like recruitment,
catch, catchability and fishing mortality rate in other
studies (Punt et al., 2001; Jurado-Molina et al., 2005;
Sinclair and Crawford, 2005; Wilberg and Bence,
2006).
variables Rt (n = 57); Ca,t (n = 33); Yt (n = 58) and Ft
(n = 40).
During the model fitting, we carefully chose the
starting values and we repeated the fitting several
times until we obtained the best fit that minimized the
a ,t Wa log-likelihood objective function. The estimated
parameters included the initial age structure (from
ages 2 to 9) in 1954 (8 parameters), age-1 recruitment
from 1954 to 2011 (58 parameters), fully recruited
fishing mortality from 1954 to 2011 (58 parameters),
catchability, three parameters for selectivity and four
parameters for the modified Beverton-Holt model,
giving a total of 132 parameters. A comparison
between the environmental proposed model and a
simplified model (without the inclusion of PDI and
AEI as explanatory variables) was made using the
log-likelihood as a measure of the goodness of fit.
The model parameters were used to estimate
)
equilibrium conditions at fixed levels of fishing
mortality rate (F) and environmental conditions, and
some Reference Points useful for fishery management
were estimated with the Microsoft Excel’s solver.
F0.1: The fishing mortality rate at which the slope of
the yield per recruit curve as a function of fishing
mortality is 10% of its value at the origin.
F40%: The fishing mortality rate that will result in a
Spawning Potential Ratio (SPR) of 40%. (SPR: The
number of eggs that could be produced by an average
recruit in a fished stock divided by the number of eggs
that could be produced by an average recruit in an
unfished stock).
FMSY: The fishing mortality rate for the Maximum
Sustainable Yield.
Author Contributions
RP (corresponding author) designed and guided the research, analyzed data
and drafted the manuscript. RP collected, processed and analyzed
anthropogenic data. RA collected, processed and analyzed fishery data and
implemented the model in Excel. LSC collected, processed and analyzed
data on tropical cyclones. MEdeL and OM collected, processed and
analyzed biologic and fishery data. All authors read and approved the final
manuscript.
43
 International Journal of Marine Science 2013, Vol.3, No.6, 36-45
http://ijms.sophiapublisher.com
Acknowledgements
We are grateful to the fishermen, specialists and managers from the fishing
sector of the Cuban Ministry of Food, for providing support and reliable
information on the spiny lobster fishery.
We acknowledge the useful comments of Dr. Nic Caputi and Dr. Simon de
Lestang from the Western Australian Fisheries and Marine Research
Laboratories. We also acknowledge the manuscript improvement by the
comments from the anonymous reviewers.
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