Species: Isurus oxyrinchus (Shortfin Mako) Justification Shortfin

advertisement
Species: Isurus oxyrinchus (Shortfin Mako)
Justification
Shortfin Mako (Isurus oxyrinchus) is an important target species, a bycatch in tuna and billfish
longline and driftnet fisheries, particularly in high-seas fisheries, and is an important coastal
recreational species. Most catches are inadequately recorded and underestimated and landings
data do not reflect numbers finned and discarded at sea. Various analyses suggest that this
species may have undergone significant declines in abundance over various parts of its range. A
global assessment of Vulnerable is considered appropriate for this species on the basis of
estimated and inferred declines, inadequate management resulting in continuing (if not
increasing) fishing pressure, the high value of its meat and fins, and vulnerable life history
characteristics. Although it is difficult to accurately assess the conservation status of this shark
because it is migratory and caught in numerous poorly monitored fisheries worldwide, it is
reasonable to assume that decreases may be occurring in those areas for which there is limited
or no data.
Mediterranean Region
Several subpopulations of Shortfin Mako h
ave been assessed separately for the IUCN Red List, however further data are required to
determine whether individuals occurring in the Mediterranean constitute a subpopulation
(based on the definition of "subpopualtion" as given in the IUCN Red List Categories and
Criteria). Recent investigations in the Mediterranean suggest that the western basin is a nursery
area where bycatch of Shortfin Mako (Isurus oxyrinchus) from the tuna and swordfish fishery
consists almost exclusively of juveniles. It is possible that this nursery area corresponds to the
Eastern Central Atlantic population, which is affected by the swordfish longline fishery off the
western coast of Africa and the Iberian peninsula. In other areas of the Mediterranean, the
Shortfin Mako is caught sporadically. Reports from the Ligurian Sea show a significant decline
since the 1970s. In the Adriatic Sea, Shortfin Makos were considered common at the end of
19th/beginning of the 20th centuries, but since 1972 there have been no records of this species
reported despite a large increase in fishing pressure and introduction of new fishing gear to the
area. On the basis of the absence of records of this species from some localised areas, evidence
of large declines in others and captures of juveniles in a probable nursery area, this species is
considered Critically Endangered in the Mediterranean, warranting focused attention and
immediate action in order to preserve this species in the region.
Geographic Range
Shortfin Mako is a coastal, oceanic species occurring from the surface to at least 500 m depth
and is widespread in temperate and tropical waters of all oceans from about 50°N (up to 60°N in
the northeast Atlantic) to 50°S. It is occasionally found close inshore where the continental shelf
is narrow. It is not normally found in waters below 16°C (Compagno 2001).
Atlantic
Casey and Kohler (1992) suggest that the core distribution in the western north Atlantic is
between 20?40°N, bordered by the Gulf Stream in the west and the mid-Atlantic ridge in the
east (see Habitat and Ecology section for more details). Shortfin Mako in Atlantic Canadian
waters represent the margins of the distribution of the population (Campana et al. 2005). In the
eastern North Atlantic, it is presumed that the Strait of Gibraltar is nursery (Buencuerpo et al.
1998 and Tudela et al. 2005).
The area between 17° to 35°S off the coast of Brazil is an area of birth, growth and mating
(Amorim et al. 1998). Pregnant females with near term embryos have been found there, but not
females in early pregnancy stages (Costa et al. 1995, Costa 1994). The presence of this species in
Uruguayan waters year round has been confirmed by the observers on board the Uruguayan
tuna fleet. Although a few new borns were captured, no pregnant females have been found
(Domingo pers. comm. 2008).
Mediterranean
Highest abundance is reported in the western Mediterranean and mako are rarely reported in
eastern waters (Aegean Sea and Sea of Marmara). Recent investigations suggest that the
western basin is a nursery area (Buencuerpo et al. 1998). It is possible that this nursery area is
from the eastern central Atlantic population, which is affected by the swordfish longline fishery
off the western coast of Africa and Iberian peninsula. Two Shortfin Makos a few months old
were reported in the western Ligurian Sea as bycatch of the swordfish longline fishery (Orsi
Relini and Garibaldi 2002). They are not reported from the Black Sea. In the eastern Adriatic Sea,
Shortfin Makos were reported as common a century ago (Katuri 1893 and Kosi 1903), recent
publications consider it to be rare (Mili?i? 1994, Jardas 1996). Soldo and Jardas (2002) report
that there have been no records of Shortfin Mako in the eastern Adriatic since 1972.
The International Council for the Exploration of the Sea (ICES) considers there to be a single
stock of shortfin mako throughout the north Atlantic, therefore the ICES area (i.e. northeast
Atlantic) consists of only part of this stock’s distribution range (ICES 2012). The north Atlantic
population does however appear to be isolated genetically from other shortfin mako
populations worldwide. The southern limit of the population is assumed to be 5o North, as for
the swordfish, based on the oceanography of equatorial waters. The relationship between
shortfin mako in the Atlantic and Mediterranean Sea is still unclear, therefore the two areas
should not necessarily be assessed as one.
Population
Shortfin Mako contribute some 9.5?10% of the pelagic sharks caught by Spanish longline fleets
(targeting sharks and swordfish) in the Atlantic and Pacific Oceans (Mejuto et al. 2002, 2005,
2006, 2007).
Shortfin Mako in the north and the south Atlantic constitute genetically distinct groups (Heist et
al. 1996). Casey and Kohler (1992) hypothesized from tag-recapture data that western north
Atlantic makos form a separate population from those in the eastern Atlantic although limited
intermixing is possible as shown by crossings to the Azores and Europe. Mitochondrial DNA data
indicate separation of female makos between the western and eastern north Atlantic, but a lack
of differentiation in nuclear DNA suggests male mixing across the north Atlantic (Heist et al.
1996, Schrey and Heist 2003). For fishery assessment purposes, mixing between the western
and eastern North Atlantic is considered minimal.
Mature males were occasionally caught in the western English Channel in the 1960s and 1970s
but are now rarely encountered (J. D. Stevens, pers. comm.), suggesting possible range
contraction in the north-east Atlantic.
Analyses of catch per unit effort (CPUE) from US pelagic longline fishery logbooks reported that
Isurus spp. may have declined by about 40% in the northwest Atlantic between 1986 and 2000
(Baum et al. 2003). A more recent assessment of observer data for the same fishery found a
similar instantaneous rate of decline of 38% between 1992 and 2005 (Baum et al. in prep). A
similar analysis of the same dataset and species grouping that restricted the areas of analysis to
account for unbalanced observations, resulted in an overall decline of 48% from beginning to
end of the time series (1992-2005; Cortes et al. in press). A 2004 ICCAT stock assessment
workshop reported that stock depletions for north Atlantic Shortfin Mako are likely to have
occurred based on CPUE declines of 50% or more. Demographic model results varied widely,
with one approach suggesting present stock size is about 80% of virgin level, and another
approach suggesting reductions to about 30% of virgin biomass (1950s) (Cortes et al. in press).
In the South Atlantic, the magnitude of decline appears to be smaller than in the north Atlantic
and the stock size appears to lie above MSY, although only one modeling approach could be
applied to the available data and assessments results were more uncertain than for the North
Atlantic.
For both north and south Atlantic populations, uncertainties about demographic parameters
and catches, and the uninformative nature of available catch data indicate that further analysis
is necessary to properly delineate stock status. If historical Shortfin Mako catch is higher than
the estimates in this report, the likelihood of the stock being below the biomass at MSY will
surely increase (ICCAT 2005). A standardized catch rate index from the commercial large pelagic
fishery off Canada suggested a decline in the 1970s and stable abundance since 1988 (Campana
et al. 2005). However, the analysis did not have the statistical power to detect anything less
than a severe decline and these sharks represent the margins of the population. The most
heavily fished areas lie outside of Canadian waters. The median size of mako sharks in the
commercial catch has declined since 1988, possibly indicating a loss of larger sharks (Campana
et al. 2005).
In the Mediterranean, ?Tonnarella? (tuna-trap) catches in the Ligurian Sea from 1950 to the
1970s show a rapid decline and eventual disappearance of the Shortfin Mako (Boero and Carli
1979). Landings data from Maltese waters for 1979?2001 (data from the Maltese fishery
department) shows a decline although the fishing pressure was not changed. Historically
described as common (end of 19th/beginning of 20th century), Soldo and Jardas (2002) report
that there have been no records of Shortfin Mako in the eastern Adriatic since 1972. Since 1998,
there have been few records of mako sharks from the central and eastern Mediterranean (A.
Soldo pers. comm.). Previously, the species was considered common throughout the
Mediterranean.
[The following 10 paragraphs of information are from WGEF 2012, including: table 9.1, page
221; figure 9.2, page 225; figure 9.3 and 9.4, page 226; and figure 9.5, page 227]
In 2010, 4,005 tonnes of landed shortfin mako catch were reported to ICCAT in the north
Atlantic, 3,546 tonnes of which were from longline fleets and 459 tonnes from other fleets (ICES
2012). This is the largest catch recorded since 1970 (the extent of the database), apart from
2004, which was more than 1000 tonnes larger. However, this figure is in the same region as
seen in 2007 and 2009. The main countries reporting catches in the North Atlantic are Spain,
Portugal, USA and Japan, accounting for 52%, 36%, 5% and 3% of total reported landings in 2010
respectively. National landings reported to ICES for 2011 were 20.6 tonnes for the northeast
Atlantic, with the majority of this from the Azores waters by Portuguese fleets (15.6 tonnes),
and smaller amounts reported by Spain (including Basque country) and France.
In the Mediterranean the total reported landings of shortfin mako were just two tonnes, with
Spain, Cyprus and Portugal each reporting catches of less than one tonne (ICES 2012). Since
1997, the Mediterranean catches have always been low (less than nine tonnes), with peak
reported catches of 17 and 10 tonnes in 2005 and 2006.
Previous ICCAT shortfin mako assessments used two other estimates of landings for this stock,
the tuna ratio (logged observations of shark catches relative to tuna catches) and the fin trade
index (shark fin trade observations from the Asian market used to calculate caught shark
weights based on catch effort data; Clarke et al. 2006; ICCAT 2005 and 2008). These figures were
much higher than actual reported landings. Similarly, discard data - where available - are
considered a large underestimation (ICES 2012).
The actual level of shortfin mako bycatch is difficult to estimate because available data are
limited and incomplete (ICES 2012). The species is of high value to both the finning and meat
trade, and many European fisheries land specimens gutted, with the head attached. The
practice of finning is likely to result in undocumented catches and mortality of the species in
some fleets.
The historical use of generic shark categories is problematic, although many European countries
have begun to report more species-specific data in recent years (ICES 2012). There have been
significant discrepancies between reported catch in databases from ICCAT, FAO and EUROSTAT.
The ICCAT Secretariat consolidated these three data-sources into a unique database, and
currently progress is being made on its validation and the associated data-mining task (analysis
of equivalent data series at various aggregation levels [Palma et al. 2012]). FAO data have been
revised in recent years, and historical catch figures have increased from those previously
reported.
Catch per unit effort data were compiled at the ICCAT assessment in 2004 (ICCAT 2005) and in
2008, and these indicated a declining trend for shortfin mako in the north Atlantic for the years
1975 to 2004. In the 2012 north Atlantic shortfin mako assessment, six catch per unit effort
series from longline fleets were presented, from Portugal, Spain, USA, Uruguay, Japan and
Brazil, and additionally another series was provided from the US Recreational Fishery. The US
pelagic longline logbook program (1986 to 2010) and the US pelagic longline observer program
(1992 to 2010) indexes of abundance showed a concave shape, marked by an initial decline until
the late 1990s, followed by an upward trend to 2010 (Cortés 2012). The Marine Recreational
Fisheries Statistics Survey data (1981 to 2010) showed high variability, with a high in the mid
1990s, followed by a decline, then a stable trend over the last ten years (Babcock 2012).
Standardized catch per unit effort from logbook data of the Japanese tuna longline fishery in the
north Atlantic (1994 and 2010) ranged from 0.07 to 0.1 between 1994 and 2005, then showed a
continuous increasing trend (Semba et al. 2012). A filtering method removed assumed
unreliable data. In general the available catch per unit effort series showed increasing or stable
trends for the final years of each series since the last stock assessment.
Although the relationship between Atlantic and Mediterranean shortfin mako is unclear, Tudela
et al. (2005) estimated catch per unit effort based on driftnetters from Al Hoceima and Nador
fishing in the Alboran Sea. Di Natale and Pelusi (2000) reported on data from the Italian large
pelagic longline fishery in the Tyrrhenian Sea (1998 to 1999) and calculated a catch per unit
effort of 1.1 kilograms per 1000 hooks.
Ferretti et al. (2008) grouped shortfin makos with another Lamnid shark species, Lamna nasus
(porbeagle), for their analysis of elasmobranch population declines in the Mediterranean,
finding the largest declines in the tuna trap of Camogli, with declines of more than 99.99% over
56 years in abundance and biomass. Similar rates of decline were observed in the northern
Ionian Sea, where a large drop in mackerel sharks caught by pelagic longlines was observed in
the early 1980s. Nevertheless, catch rates were very low even at the beginning of the data
series, with an average of 0.2 sharks per 1000 hooks. The meta-analytical estimate of the rate of
decline was more than 99.99% for biomass (IRD: –0.15; CI 95%: –0.21, –0.10; time range: 106
years) and abundance (IRD: –0.12; CI 95%: –0.22, –0.03; time range: 135 years).
Between 2008 and 2011, the average catch per unit effort of shortfin makos in the north
Atlantic Portuguese swordfish longline fishery was 1.3 individuals per 1,000 hooks (Coelho et al.
2012).
ICCAT nominal catch 1999 to 2012: [Filename: ICCAT - TASK1 - I. oxyrinchus]
The shortfin mako is highly susceptible to overexploitation by pelagic longline fisheries in the
Atlantic Ocean, according to a productivity and susceptibility analysis carried out by Cortés et al.
(2010), which revealed the species to be amongst the most vulnerable. The analysis also
revealed that shortfin makos are less productive than previously thought based on more
recently acquired life history variables, and that post-capture mortality for the species was 92%.
Shortfin makos have a very low capacity to withstand fishing mortality, at the same time as
experiencing intense exploitation (Dulvy et al. 2008). For this species, regional variation in the
intensity of fishing mortality is an important factor determining their global threatened status.
Shortfin makos are sought for both their meat and fins; thus unlike many other oceanic pelagic
sharks, they are frequently targeted by the large longline fleets in the Atlantic Ocean. The
shortfin mako comprises up to approximately 7% of total catches (weight) in the Atlantic
swordfish fishery and approximately 10% (weight) of all north Atlantic shark catches (Mejuto et
al. 2006a,b; Hareide et al. 2007). This species is also a highly prized recreational gamefish (Dulvy
et al. 2008). Major declines in shortfin mako abundance have occurred, notably in the eastern
Mediterranean Sea where they are now rarely seen. In the north Atlantic, population declines of
up to 70% have been documented (ICCAT 2005).
Between 1998 and 2000, the shortfin mako was caught more often in the swordfish fishery in
the Mediterranean than any other fishery, with a mean catch per unit effort of 0.07 fish per
1,000 hooks in the “American type” longline and 0.05 fish per 1,000 hooks in the standard
longline. Shortfin makos were more abundant in the Alboran Sea and the Levantine basin than
other areas of the Mediterranean surveyed by Megalofonou et al. (2005). The Azorean fleet
mako landings decreased by almost 50% in numbers from 1987 to 1994 (Castro et al. 1999).
Together with the low catch rates in the Mediterranean Sea, shortfin makos may be one of the
most over-fished pelagic sharks in the Mediterranean Sea (Megalofonou et al. 2005).
Using a novel method of filtering fishery logbooks to obtain catch per unit effort data, as well as
to deduce the levels of underreporting of shark species, Nakano and Clarke (2006) found that
shortfin mako was being under-reported by an average of 33% across all logbooks analysed,
with the worst case scenario for the species revealing an under-reporting rate of 65%.
Trend
Decreasing
Habitat and Ecology
Habitat and movements
The Shortfin Mako is an active, offshore littoral and epipelagic species, found in tropical and
warm-temperate seas from the surface down to at least 500 m, seldom occurring where water
temperature is <16°C (Compagno 2002). It is probably the fastest shark and is among the most
active and powerful of fishes. Like other lamnid sharks, the Shortfin Mako is endothermic using
a heat-exchanging circulatory system to maintain muscle and visceral temperatures above that
of the surrounding seawater allowing a higher level of activity (Carey et al. 1981, Bernal et al.
2001). This shark occurs well offshore but penetrates the inshore littoral just off the surf zone in
some areas such as parts of KwaZulu-Natal, South Africa where the continental shelves are
narrow. Off South Africa, shark meshing data suggests that this species occurs in clear to turbid
water in water temperatures from 17?22°C. In the western north Atlantic it occurs in a similar
range of temperatures, and only moves onto the continental shelf when surface temperatures
exceed 17°C. In the eastern north Pacific, juveniles range into southern Californian waters and
tend to be seen and caught near the surface. They appear to use these offshore continental
waters as nursery areas (Taylor and Holts 2001). It was previously thought that they stay near
the surface above 20 m depth, in waters between 20?21°C, seldom descending into cold
subsurface waters below the thermocline (Holts and Bedford 1992). However, this has been
challenged by more recent tracking studies (summarized below).
Results from a large tagging study in the western north Atlantic show that Shortfin Makos make
extensive movements of up to 3,433 km with 36% of recaptures caught at greater than 420 km
from their tagging site (Casey and Kohler 1992). However, only one fish crossed the mid-Atlantic
ridge suggesting that trans-Atlantic migrations are not as common as in blue sharks Prionace
glauca. Klimley et al. (2002) tracked three shortfin makos near La Jolla, California, for several
days, and their movements were mainly offshore from the surface to 50 m. Holts and Kohin
(2003) deployed pop-up archival tags on eight makos (118?275 cm TL) in June?July 2002 for 2?4
months. Pop-up locations ranged from 20?911 km from deployment locations. The sharks
utilized near-shore and open-water areas off California and Baja California roughly between
23?43°N and out to 125°W. While the records indicate that greater than 90% of the time was
spent above 50 m, several sharks showed a diurnal pattern of vertical excursions to beyond 200
m during daylight hours. Sharks frequently dove into water less than 10°C. These data
demonstrate the range of habitats utilized by mako sharks and begin to shed light on their daily
and seasonal behaviours. Sepulveda et al. (2004) found that seven tagged juveniles stayed near
the surface at night, and went as deep as 200 m, mostly during the day. In addition, stomach
temperatures were measured, indicating feeding occurred during the daytime, with meals taken
during a dive causing stomach temperatures to drop noticeably.
Life History Parameters
The Shortfin Mako reaches a maximum size of about 4 m (Compagno 2001). Initial age and
growth studies in the western north Atlantic suggested that two pairs of growth bands are laid
down each year in their vertebral centra, at least in young shortfin makos (Pratt and Casey
1983). However, recent evidence using marginal increment analysis in Mexico (Ribot-Carballal et
al. 2005) and bomb radiocarbon (Campana et al. 2002, Ardizzone et al. 2006) indicates that the
alternative hypothesis (one pair of growth bands per year; Cailliet et al. 1983) is valid. Age at
maturity has been determined recently in several populations, including New Zealand (7?9 years
for males, and 19?21 years for females Bishop et al. (2006)), and the western north Atlantic
(eight years for males, and 18 years for females (Natanson et al. 2006)). Longevity has been
estimated as 29?32 years (Bishop et al. 2006, Natanson et al. 2006).
There is a large difference in size at sexual maturity between the sexes. In the northwest
Atlantic, males reach maturity at about 195 cm and females at about 265?280 cm (Pratt and
Casey 1983, Stevens 1983, Cliff et al. 1990). In New Zealand, males mature at 198?204 cm and
females at 301?307 cm (Francis and Duffy 2005). Compagno (2001) reports males mature
between 203?215 cm, reaching a maximum size of 296 cm, and females mature between
275?293 cm, reaching a maximum of almost 4 m.
The Shortfin Mako is ovoviviparous and oophagous, but what little is known of its reproductive
cycle indicates the gestation period is 15?18 months, with a three year reproductive cycle
(Mollet et al. 2002). Litter size is 4?25 pups (possibly up to 30, mostly 10?18), which are about
60?70 cm long at birth (Garrick 1967, Compagno 2001). There are comparatively few records of
pregnant females. Among 26 shark species, the Shortfin Mako has an intrinsic rebound potential
(a measure of its ability to recover from exploitation) in the mid-range (Smith et al. 1998). The
annual rate of population increase is 0.046 yr-1 (S. Smith pers. comm.) Cortes (2002) calculated a
finite rate of increase (lambda) of 1.141 (1.098 to 1.181 95% CI, r = 0.13) and the average
reproductive age as 10.1 (9.2 to 11.1 95% CI) years.
Diet
The diet of Shortfin Makos has been reported to consist mainly of teleost fishes (including
mackerels, tunas, bonitos and other scombrids, anchovies, herrings, grunts, lancet fishes, cod,
ling, whiting and other gadids, salmon, yellowtails and other carangids, sea basses, porgies,
swordfish) and cephalopods in studies from the northwest Atlantic and Australia (Stillwell and
Kohler 1982, Stevens 1984), while elasmobranchs were the most common prey category from
Natal, South Africa (Cliff et al. 1990). A daily ration of 2 kg/day (based on an average weight of
63 kg) was estimated for makos in the northwest Atlantic (Stillwell and Kohler 1982). Large
makos (over 3 m in length) have very broad, more flattened and triangular teeth, perhaps better
suited to cutting large prey than the awl-shaped teeth of smaller individuals (Compagno 1984a).
There are several anecdotal accounts of makos attacking and consuming Broad-bill Swordfish
Xiphias gladius. It also eats sea turtles, dolphins, salps and occasionally detritus (Compagno
1984a).
Stevens (2008) suggested that nursery areas would likely be situated close to the coast in highly
productive areas, based on the majority of reports, with nursery grounds off West Africa in the
north Atlantic.
Maia et al. (2007) reported length at 50% maturity of 180 centimetres in male shortfin makos,
but did not deduce this value for females as no mature individuals were caught.
Threats
The Shortfin Mako is an important species for pelagic longline, drifting or set gill nets and on
hook-and-line fisheries wherever it occurs, particularly from nations with high seas fleets (Holts
et al. 1998), because of its relatively high abundance (9.5?10% of pelagic sharks caught in
Spanish long line fleets (Mejuto et al. 2002, 2005, 2006, 2007) and high quality meat. It is taken
as a bycatch from tuna and swordfish longline fisheries worldwide, with carcasses and fins being
retained. Big-game sports angling for mako sharks is widespread, New Zealand and South Africa
being traditional places for offshore sports fishing. The International Game Fish Association lists
the shortfin mako as a record game fish. In the 1980s mako angling became popular in the USA
off southern California, with numerous anglers involved and mako tournaments rivaling
competitive angling for marlin (Compagno 2001). Recreational fishing has also been reported in
the Mediterranean, although there are no official data (A. Soldo pers. comm.).
Fisheries for Shortfin Mako exist or existed in the eastern Atlantic, the Mediterranean Sea, off
Cuba, in the Gulf of Mexico and Caribbean, off southern California, and in the western and
central Pacific (Compagno 2001). This is also a target of pelagic swordfish fisheries in the Atlantic
and Pacific (Mejuto et al. IBID).
Despite the role of Shortfin Makos in worldwide pelagic fisheries, catches have been poorly
reported to FAO. Catch data are incomplete, and the extent of finning in high seas fisheries is
unclear. Brazil, New Zealand and the United States reported very small catches (2?76 t) to FAO
from 1987?1997 (FAO FishStat Plus database 2000). Although it is difficult to accurately assess
the conservation status of this shark because it is migratory and caught in numerous poorly
monitored fisheries worldwide, it is reasonable to assume that decreases may be occurring in
those areas for which there are limited or no data (Castro et al. 1999).
Atlantic
Shortfin Makos have been caught in large numbers particularly on the high seas in pelagic
longline fisheries, but also in other commercial pelagic fisheries and recreational fisheries. The
first longline fisheries were prosecuted by Japan in western equatorial waters beginning in 1956
(Uozumi and Nakano 1996). The fleet expanded rapidly in the 1960s, and covered almost the
entire Atlantic by the late 1960s (Bonfil 1994), including the areas currently fished by the
American fleet. Throughout the Atlantic, the fleet landed mako sharks and fins (Nakano 1993).
In the US and Canadian pelagic longline fisheries, Shortfin Mako is one of the most commonly
caught sharks. The index of abundance in the commercial longline fishery off the Atlantic coast
of US has shown a steady decline (Cramer 1996) and other reports on declines are now available
(Baum et al. 2003, Baum et al., in prep, ICCAT 2005, Cortes et al. in press: see Population section
for details). As for recreational fishing, Casey and Hoey (1985) stated that the recreational catch
of Shortfin Makos along the US Atlantic coast and in the Gulf of Mexico in 1978 was 17,973 fish
weighing some 1,223 t. Between 1987 and 1989, the catch was about 1000 t/year (Casey and
Kohler 1992) taken by longline and gillnet in the Southwest Atlantic (see Population section).
Shortfin Mako shark is a high value bycatch of pelagic longline fisheries on the Atlantic coast of
Canada and therefore retained (Campana et al. 2005).
In 1989, Bonfil (1994) estimated that 5,932 Shortfin Makos were caught by Korean longliners in
the (mainly equatorial) Atlantic and that 763 t of makos were landed in the Spanish swordfish
fishery in the Mediterranean and Atlantic. Mejuto (1985) noted that 304?366 t of mako shark
was landed by longliners operating from northern Spain in 1983?84. More recently, shortfin
mako sharks have comprised about 7% (~2,500 t) of the total catch of the large Spanish pelagic
longline swordfish fleet in the Atlantic (Mejuto et al. 2005). Munoz-Chapuli et al. (1993)
estimated that some 4,500 makos/year are landed from a longline fishery based at Algeciras,
southern Spain (given an average weight of 20 kg this would represent about 90 t). The landings
of Shortfin Makos as bycatch from the swordfish fishery of the Azorean fleet also showed a
decrease (Castro et al. 1999). Shortfin Mako landings reported to ICCAT from Portuguese
surface longline fisheries in the north Atlantic averaged about 698 t during 1993?1996 and 340 t
for the period 1997?2002. Off Namibia, the large pelagic fisheries caught an estimated 123 t in
2001, 399 t in 2002 and 393 t in 2003 by means of pelagic longline. The 2001 catch is an
underestimate as many boats grouped different shark species as ?sharks? (MFMR catch data).
Domingo (2002) records high catches of Shortfin Makos by the Uruguayan fleet in the early-mid
1980s (to a maximum of 144 t in 1984), followed by much lower catches (10?20 t/annum) in the
1990s. This does not necessarily reflect stock abundance because changes in the distribution
and depth of fishing operations and rising mean temperature of water masses in the area had
also occurred.
It has been estimated that in the early 1990s, the Spanish longline fleet caught approximately
750 t/y of Shortfin Mako sharks in the Atlantic Ocean and Mediterranean Sea (Bonfil 1994,
Compagno 2001). The Brazilian longlining fleet based in Santos landed between 13.3 and 138.3 t
annually between 1971 and 1990 (Costa et al. 1996, Compagno 2001). Despite increasing fishing
effort during this period, the CPUE of Shortfin Makos has remained relatively stable with an
initial slight decreasing trend followed by a slight increasing trend (Compagno 2001).
No complete data are available for the northeast Atlantic, but the species is taken as a bycatch
of the pelagic fishery. The area around the Strait of Gibraltar is considered a nursery area for
central Atlantic Shortfin Makos and most specimens caught are juveniles. This area is heavily
fished by the swordfish longline fishery off the western coast of Africa and Iberian peninsula.
There is also evidence that Shortfin Makos are becoming increasingly targeted in the western
Mediterranean. EU vessels fishing for small pelagic species off the western coast of Africa are
also known to take significant elasmobranch bycatch, including Shortfin Makos in unknown
numbers.
Mediterranean
Reports of ?Tonnarella? catches in the Ligurian Sea from 1950 until the 1970s show a rapid
decline and eventual disappearance of the Shortfin Mako (INP 2000). Recent investigations of
the shortfin mako bycatch from the swordfish longline fishery in the western basin show that
catches from this fishery consist almost exclusively of juveniles. It is likely that the western
Mediterranean is a nursery area for the eastern Central Atlantic population Soldo and Jardas
(2002) report that there have been no records of shortfin mako in the eastern Adriatic since
1972 (where they were historically common).
Even though driftnetting is banned in Mediterranean waters, this practise has continued illegally
(WWF 2005). The Moroccan swordfish driftnet fleet in the Alboran Sea operates year round,
resulting in high annual effort levels (Tudela et al. 2005). Even though sharks are a secondary
target or bycatch of this fishery, some boats deploy driftnets 1?2 miles from the coast where the
chance of capturing pelagic sharks is higher. The catch rate for Shortfin Mako is nearly three
times higher in boats actively fishing for sharks (from 0.6 to 1.9 N/fishing operation and 0.06 to
0.14 catch per km net). Both annual catches and mean weights of shortfin mako have fallen as a
result of fishing mortality in the Moroccan driftnet fishery, illustrating the likely impact of this
illegal fishery on stocks in the Alboran Sea and adjacent Atlantic (Tudela et al. 2005).
Megalofonou et al. (2005) reported 321 specimens caught as bycatch in tuna and swordfish
fisheries in the Mediterranean Sea. Of those, 268 specimens were caught in Alboran Sea, 42 in
Balearic Islands area, three in Catalonian Sea, while only eight specimens were caught in central
and eastern Mediterranean area, e.g., Levantine basin. Furthermore, most of caught specimens
were juveniles, with only a few large specimens from Levantine basin. Of 595 specimens caught
in south Spain waters all of them were immature juveniles (Buencuerpo et al. 1998). Reports
from the Ligurian Sea show a significant decline since the 1970s (Boero and Carli 1979).
In the Hong Kong shark fin market, some traders mentioned infrequent mixing with the less
abundant I. paucus (longfin mako) (Clarke et al. 2006). Combining results of the genetic analyses
from this study with the stochastic modeling of trade records allowed the contribution of
individual species to the trade to be determined. Shortfin mako fins showed an 85%
concordance with the market category ‘qing lian’, which equates to a trade proportion for this
species of approximately 2.7% (probability interval 2.3–3.1%). The actual percentage is likely to
be somewhat higher, as there was presence of shortfin mako fins in the ‘wu yang’ (silky shark)
trade category also.
Conservation
A vast improvement in the collection of data is required and effective conservation of this
species will require international agreements. Fishing pressure must be considerably decreased
through reduction in effort, catch limits, measures to enhance chances of survival after capture,
when released and possibly through the implementation of large-scale oceanic non-fishing
areas. Closed areas can only be effective if overall fishing effort is reduced, rather than merely
displacing effort outside of the closed area (Baum et al. 2003).
The Shortfin Mako is listed as a highly migratory species under the 1995 UN Agreement on the
Conservation and Management of Straddling Fish Stocks and Highly Migratory Fish Stocks
(UNFSA). The Agreement specifically requires coastal and fishing States to cooperate and adopt
measures to ensure the conservation of listed species. To date, there has been little progress
(see http://www.unclos.com for further details). Also of relevance is the FAO International Plan
of Action for the Conservation and Management of Sharks (IPOA-Sharks) which recommends
that Regional Fisheries Organisations (RFO?s) carry out regular shark population assessments
and that member States cooperate on joint and regional shark management plans. This is of
particular importance for species such as shortfin mako whose stocks are exploited by many
State on the high seas. Steps are being taken by some RFOs, such as ICCAT, to collect speciesspecific data on pelagic sharks. To date two RFOs, ICCAT and IATTC, have adopted finning bans,
as have several range states (e.g., Canada, USA, EU, Australia, Brazil etc.). More are likely to
follow suit.
A recent shark stock assessment (ICCAT 2005) reported that the overall volume of catch
reported to ICCAT does not represent the total removals of sharks and that the data are also
very limited with respect to the size-, age- and sex- composition of the catch. It is noted that
improvements in the ICCAT shark database can only be achieved if the Contracting Parties
increase infrastructure investment into monitoring the overall catch composition and
disposition of the catch of sharks and by-catch species. The workshop recommended larger
monitoring and research investments directed at sharks in particular and by-catch species. The
group also identified a number of research activities that could provide for improved advice on
the status of these species. This situation applies to all RFOs and is included here as a standard
that needs to be applied internationally.
In 2004 ICCAT requested that management recommendations for this species be developed in
2005 for consideration by the Commission (ICCAT 2004), so far without effect. In the fall of
2008, ICCAT and the ICES Working Group on Elasmobranch Fishes (WGEF) plan to assess shortfin
mako jointly. In 2007, at the data preparatory meeting of ICCAT?s Shark Sub-committee, it was
emphasized that the contribution of CPUE and historical catch data by the member countries
would be very important for the stock assessment of this species to be made at the 2008
meeting.
Within the north Atlantic and Mediterranean this species has been identified as a high priority
for management. Anonymous (2003) suggested that a collaborative stock assessment should be
carried out in the future.
Mediterranean
The draft action plan for the conservation of cartilaginous fishes in the Mediterranean Sea
(Anonymous 2002) highlights that the Shortfin Mako, along with other large pelagic sharks
(whether as target or bycatch), urgently require measures to ensure their sustainable
management in the Mediterranean.
European Commission (EC) Regulation No. 1185/2003 prohibited the removal of shark fins and
subsequent discarding of the body, which was binding on EC vessels in all waters and non-EC
vessels in Community waters. A fin-to-carcass ratio of 5% of the shark’s whole (theoretical)
weight was adopted. This Regulation also allowed Member States to issue Special Fishing
Permits allowing sharks to be processed on board by removing their fins from their bodies. This
loophole was eventually amended in 2013 by Regulation (EU) No. 605/2013 of the European
parliament, which recognises “serious control and enforcement difficulties” associated with the
fin-to-carcass ratio, and states that it “could lead to finning going undetected”. The amendment
states, “The Scientific, Technical, and Economic Committee for Fisheries (STECF) acknowledges
the problem of shark finning, calls for its eradication, without derogations, and advises that all
elasmobranch species should be landed with their fins/wings naturally attached to their bodies.
… Without prejudice to paragraph 1, in order to facilitate on-board storage, shark fins may be
partially sliced through and folded against the carcass, but shall not be removed from the
carcass before landing”.
The shortfin mako is listed in Appendix II of the Convention on Migratory Species (CMS).
Migratory species that need or would significantly benefit from international co-operation are
listed in Appendix II of the Convention. For this reason, the Convention encourages the Range
States to conclude global or regional Agreements.
The shortfin mako is listed in Appendix II of the Barcelona Convention, affording it protection
from fishing activities taking place in the Mediterranean region. All species listed in Appendix II
must be released unharmed and alive to the extent possible, therefore cannot be retained on
board, transshipped, landed, transferred, stored, sold, displayed or offered for sale
(Recommendation GFCM/36/2012/1). The recommendation continues to stipulate that all
vessels encountering these species must record information on fishing activities, catch data,
incidental taking, release and/or discarding events in a logbook or similar document, then all
logged information must be reported to national authorities. Finally, additional measures should
be taken to improve such data gathering in view of scientific monitoring of the species.
Bibliography
Acuña, E., Cid, L., Pérez, E., Kong, I., Araya, M., Lamilla, J. and Peñailillo, J. 2001. Estudio
biológico de tiburones (marrajo dentudo, azulejo y tiburón sardinero) en la zona norte y central
de Chile. Informes Técnicos FIP, FIP/IT Nº 2000-23.
Amorim, A.F., Arfelli, C.A. and Fagundes, L. 1998. Pelagic elasmobranchs caught by longliners off
southern Brazil during 1974-97: an overview. Marine and Freshwater Research 49: 621-632.
Anderson, R.C. and Simpfendorfer, C.A. 2005. Indian Ocean. In: S.L. Fowler, M. Camhi, G.H.
Burgess, G.M. Cailliet, S.V. Fordham, R.D. Cavanagh, C.A. Simpfendorfer and J.A. Musick (eds),
Sharks, rays and chimaeras: the status of the chondrichthyan fishes, pp. 140-149. IUCN SSC Shark
Specialist Group, IUCN, Gland, Switzerland and Cambridge, UK.
Anonymous. 2001. Chapter 07b: Shortfin Mako shark: history of the fishery. California?s living
marine resources: A status report. California Department of Fish & Game, Long Beach,
California.
Anonymous. 2002. Report of the meeting of experts for the elaboration of an Action Plan for the
conservation of Mediterranean species of cartilaginous fish. UNEP, RAC/SPA, Tunis.
Anonymous. 2003. Commission staff working paper report of AD working group elasmobranches
fisheries. Brussels.
Ardizzone, D., Cailliet, G.M., Natanson, L.J., Andrews, A.H., Kerr, L.A. and Brown T.A. 2006.
Application of bomb radiocarbon chronologies to shortfin mako (Isurus oxyrinchus) age
validation. Environmental Biology of Fishes 77: 355-366.
Babcock, E., A. 2010. Updated index of abundance for shortfin mako sharks from the U.S. marine recreational fisheries statistics survey. ICCAT SCRS 2012/077.
Baum, J.K., Myers, R.A. and Blanchard, W. in prep.. manuscript in prep..
Baum, J.K., Myers, R.A., Kehler, D.G., Worm, B., Harley, S.J. and Doherty, P.A. 2003. Collapse and
conservation of shark populations in the Northwest Atlantic. Science 299: 389-392.
Bedford, D.W. 1992. Mako shark. In: W.S. Leet, C.M. Dewees and C.W. Haugen (eds),
California?s living marine resources and their utilization, pp. 51-53. California Sea Grant
Extension Publication.
Bello, G. 1999. The Chondrichthyans of the Adriatic Sea. Acta Adriatica 40(1): 65-76.
Bernal, D., Dickson, K.D., Shadwick, R.E. and Graham, J.B. 2001. Analysis of the evolutionary
convergence for high performance swimming in lamnid sharks and tunas. Comparative
Biochemical Physiology 129: 695-726.
Bianchi, G., Carpenter, K.E., Roux, J.-P., Molloy, F.J., Boyer, D. and Boyer, H.J. 1999. Field guide to
the living marine resources of Namibia. FAO, Rome, Italy.
Bishop, S.D.H., Francis, M.P., Duffy, C. and Montgomery, J.C. 2006. Age, growth, maturity,
longevity and natural mortality of the shortfin mako shark (Isurus oxyrinchus) in New Zealand
waters. Marine and Freshwater Research 57: 143-154.
Boero, F. and Carli, A. 1979. Catture di Elasmobranchi nella tonnarella di Camogli (Genova) dal
1950 al 1974. Boll. Mus. Ist. Biol. Univ. Genova 47: 27-34.
Bonfil, R. 1994. Overview of world elasmobranch fisheries. FAO Fisheries Technical Paper 341.
FAO, Rome.
Buencuerpo, V., Rios, S. and Moron, J. 1998. Pelagic sharks associated with the swordfish,
Xiphias gladius, fishery in the eastern North Atlantic Ocean and the Strait of Gibraltar. Fishery
Bulletin 96: 667-685.
Bustamante, C.; Concha, F.; Balbont_ın, F.; Lamilla, J., 2009: Southernmost
record of Isurus paucus Gitart-Manday, 1966 (Elasmobranchii:
Lamnidae) in the southeast Pacific Ocean. Rev. Biol.
Mar. Oceanogr. 44, 523–526.
Cailliet, G.M. and Bedford, D.W. 1983. The biology of three pelagic sharks from California
waters, and their emerging fisheries: a review. California Cooperative Oceanic Fisheries
Investigations Reports.
Cailliet, G.M., Martin, L.K. Harvey, J.T., Kusher, D. and Welden, B.A. 1983. Preliminary studies on
the age and growth of blue (Prionace glauca), common thresher (Alopias vulpinus), and shortfin
mako (Isurus oxyrinchus) sharks from California waters. In: E.D. Prince and M. Pulos (eds),
Proceedings, international workshop on age determination of oceanic pelagic fishes-tunas,
billfishes, sharks, pp. 179-188.
California Department of Fish and Game. 1999. Ocean sport fishing regulations concerning mako
shark. In: State of California Department of Fish and Game (eds), Shark Tagging News. California
Department of Fish and Game, Long Beach, California, Natural Resources Agency.
California Department of Fish and Game. 2003. Commercial fishing regulations: Marine Mammal
Protection Act section 118.
Campana, S.E., Marks, L. and Joyce, W. 2005. The biology and fishery of shortfin mako sharks
(Isurus oxyrhinchus) in Atlantic Canadian waters. Fisheries Research 73: 341-352.
Campana, S.E., Natanson, L.J., and Myklevoll, S. 2002. Bomb dating and age determination of
large pelagic sharks. Canadian Journal of Fisheries and Aquatic Science 59:: 450-455.
Carey, F.G. and Teal, J.M. 1969. Mako and porbeagle: warm bodied sharks. Comparative
Biochemical Physiology 28: 199-204.
Carey, F.G., Teal, J.M. and Kanwisher, J.W. 1981. The visceral temperature of mackerel sharks
(Lamnidae). Physiological Zoology 54: 334-344.
Casey, J.G. and Hoey,J.J. 1985. Estimated catches of large sharks by US recreational fishermen in
the Atlantic and Gulf of Mexico. Shark catches from selected fisheries off the US East Coast.
NOAA Technical Paper NMFS SSRF. NOAA.
Casey, J.G. and Kohler, N.E. 1992. Tagging studies on the shortfin mako shark (Isurus oxyrinchus)
in the western North Atlantic. Australian Journal of Marine and Freshwater Research 43: 45-60.
Castro, J., De La Serna, J.M., Macias, D. and Mejuto, J. 2000. Preliminary scientific estimates of
by-catch landings by the Spanish surface longline fleet in 1997 and 1998. Collect. Vol. Sci. Pap.
ICCAT/Recl. Doc. Sci. CICTA/Colecc.Doc. Cient. CICAA.
Castro, J.I., Woodley, C.M. and Brudek, R.L. 1999. A preliminary evaluation of the status of shark
species. FAO Fisheries Technical Paper 380. FAO, Rome.
Clarke, S.C., Magnussen, J.E., Abercrombie, D.L., McAllister, M.K. and Shivji, M.S. 2006.
Identification of Shark Species Composition and Proportion in the Hong Kong Shark Fin Market
Based on Molecular Genetics and Trade Records. Conservation Biology 20(1): 201-211.
Cliff, G., Dudley, S.F.J. and Davis, B. 1990. Sharks caught in the protective gillnets of Natal, South
Africa. 3. The shortfin mako shark Isurus oxyrinchus (Rafinesque). South African Journal of
Marine Science 9: 115-126.
An overview of the hooking mortality of elasmobranchs caught
in a swordfish pelagic longline fishery in the Atlantic Ocean
Rui Coelho1,2,a, Joana Fernandez-Carvalho1, Pedro G. Lino1 and Miguel N. Santos Aquat. Living
Resour. 25, 311–319 (2012)
Compagno, L.J.V. 1984. FAO species catalogue. Vol. 4. Sharks of the world. An annotated and
illustrated catalogue of shark species known to date. Part 1. Hexanchiformes to Lamniformes.
FAO, Rome.
Compagno, L.J.V. 1984. Sharks of the World: an annotated and illustrated catalogue of the shark
species known to date. Part 2. Carcharhiniformes. FAO, Rome.
Compagno, L.J.V. 2001. Sharks of the world. An annotated and illustrated catalogue of shark
species known to date. Volume 2. Bullhead, Mackerel and Carpet Sharks (Heterodontiformes,
Lamniformes and Orectolobiformes). FAO, Rome.
Compagno, L.J.V. 2002. Sharks of the World. An annotated and illustrated catalogue of shark
species known to date. Vol. 2. Bullhead, mackerel and carpet sharks (Heterodontiformes,
Lamniformes and Orectolobiformes). FAO, Rome.
Cortes, E. 2002. Incorporating uncertainty into demographic modeling: application to shark
populations and their conservation. Conservation Biology 16: 1048-1062.
Cortés, E. 2012. Standardised catch rates of mako sharks from the US pelagic longline logbook
and observer programs using a generalized linear mixed model. ICCAT SCRS 2012/070.
Cortés, E., Brown, C. and Beerkircher, L.R. 2007. Relative abundance and average size trends of
pelagic sharks in the northwest Atlantic ocean, including the Gulf of Mexico and Caribbean Sea.
Gulf and Caribbean Research 19(2): 37-52.
Cortés, E., Arocha. F., Beerkircher, L., Carvalho, F., Domingo, A., Heupel, M., Holtzhausen, H.,
Santos, M.N., Ribera, M. and Simpfendorfer, C. 2010. Ecological risk assessment of pelagic
sharks caught in Atlantic pelagic longline fisheries. Aquatic Living Resources 23: 25-34.
Costa, F.E.S. 1995. Analise de la pesca e aspectos da biología do anikim Isurus oxyrinchus
Rafinesque 1809, capturado nas regioes sudeste e sul do Brasil (Elasmobranchii ? Lamnidae).
Dissertasao de Mestrado. Universidade Estadual Paulista.
Cramer, J. 1996. Large pelagic logbook indices for sharks. Sharks stock Assessment Workshop.
NOAA/NMFS/SEFSC, Miami.
Crooke, S. 2001. Highly migratory species: Overview. In: W.S. Leet, C.M. Dewees, R. Klingbeil,
and E.J. Larson (eds), California?s Living Marine Resources: A Status Report, pp. 315-316. The
Resources Agency, California Department of Fish and Game.
Dai, X.
1997. A preliminary analysis on the composition of
catches obtained by longline fishing in the Mediterranean
Sea. J. Shangai Fish. Univ. 6:107−111.
De la Serna, J.M., Valeiras, J., Ortiz, J.M. and Macias D. 2002. Large Pelagic sharks as by-catch in
the Mediterranean Swordfish Longline Fishery: some biological aspects. NAFO SCR Doc.02/137
Serial No. N4759.
de Young, C. 2006. Review of the state of world marine capture fisheries management: Indian
Ocean. FAO, Rome.
Di Natale A. and Pelusi P. 2000. Effetti della pesca col palangaro derivante sui grandi pelagici e
sulle specie accessorie nel Mediterraneo Centrale. Rapporto al Ministero per le Politiche
Agricole e Forestali, DGPA, Roma-IV Piano Triennale.
Domingo, A. 2000. Los Elasmobranquios Pelágicos Capturados por la flota de longline Uruguaya.
In: M. Rey (ed.), Consideraciones Sobre la Pesca Incidental Producida por la Actividad de la Flota
Atunera Dirigida a Grandes Pelágicos. ?Plan De Investigación Pesquera?. Inape ? Pnud
Uru/92/003.
Domingo, A., Forselledo, R., Miller, P. and Passadore, C. 2008. Plan de Acción Nacional para la
Conservación de Condrictios en las Pesquerías Uruguayas (PAN - Condrictios Uruguay). Dirección
Nacional de Recursos Acuáticos, Montevideo, Uruguay.
Ferretti, F., Myers, R.A., Serena, F. and Lotze, H.K. 2008. Loss of Large Predatory Sharks from the
Mediterranean Sea. Conservation Biology 22(4): 952-964.
Fischer, W., Schneider, M. and Bauchot, M.L. 1987. Vertebres, Mediterranee et Mer Noire. FAO
ECEE, Rome:.
Francis, M.P. And Duffy, C. 2005. Length at maturity in three pelagic sharks (Lamna nasus, Isurus
oxyrinchus, and Prionace glauca) from New Zealand. Fishery Bulletin 103: 489-500.
Garibaldi, F., Orsi Relini L. 2000.. Abbondanza estiva, struttura di taglia e nicchia alimentare della
verdesca, Prionace glauca, nel santuario pelagico del mar Ligure. Biologia Marina Mediterranea
7((1):): 324-333.
Hanan, D.A., Holts, D.B. and Coan Jr., A.L. 1993.. The California drift gill net fishery for sharks and
swordfish, 1981-82 through 1990-91. Fish Bulletin 175:: 94 p.
Hareide NR, Carlson J, Clarke M, Clarke S, Ellis J, Fordham S, Fowler S, Pinho M, Raymakers C,
Serena F et al.
2007. European Shark Fisheries: a preliminary investigation into fisheries, conversion factors,
trade products,
markets and management measures. European Elasmobranch Association.
Hazin, F., Hazin, H., Carvalho, F., Wor, C. and Travassos, P. In press. Standardization of CPUE
series of Prionace glauca and Isurus oxyrinchus caught by Brazilian longliners in the Western
South Atlantic Ocean, from 1978 to 2006. Data Preparatory Meeting of the ICCAT Shark Species
Group. Punta del Este, Uruguay.
Heist, E.J.; Musick, J.A.; Graves, J.E. 1996.. Genetic population structure of the shortfin mako
(Isurus oxyrinchus) inferred from restriction fragment length polymorphism analysis of
mitochondrial DNA. Canadian Journal of Fisheries and Aquatic Science 53:: 583-588.
Herrera, M., Zarate, P. and Gaibor, N. In press. Tiburones en las Pesquerias del Ecuador. Instituto
Nacional de Pesca.
Holts, D.B. 1988. Review of U.S. west coast commercial shark fisheries. Marine Fisheries Review
50(1): 1-8.
Holts, D.B. and Bedford, D. 1989. Report of the assessment methods workshop for sharks.
National Marine Fisheries Service.
Holts, D.B. and Bedford, D.W. 1993.. Horizontal and vertical movements of the shortfin mako
shark, Isurus oxyrinchus, in the Southern California Bight. Australian Journal of Marine and
Freshwater Research 44:: 901-909.
Holts, D.B. and Kohin, S. 2003. Pop-up archival tagging of shortfin mako sharks, Isurus
oxyrinchus, in the Southern California Bight. Anstract. American Fisheries Society, Western
Division meetings. American Fisheries Society, San Diego, California.
Holts, D. B., Julian, A., Sosa-Nishizaki, O., and Bartoo, N.W. 1998.. Pelagic shark fisheries along
the west coast of the United States and Baja California, Mexico. Fisheries Research 39:: 115-125.
Holts et al. 2004. Trends in CPUE from surveys.
Hurley, P.C.F. 1998.. A review of the fishery for pelagic sharks in Atlantic Canada. Special issue
science and management of shark fisheries. In: Hueter, R.E. (ed). Proceedings of an international
symposium held at the 125th annual meeting of the American Fisheries Society, Ta. Fisheries
Research 39:: 115?125.
ICCAT. 2005. Report of the 2004 Inter-sessional meeting of the ICCAT Subcommittee on
bycatches: shark stock assessment. Col. Vol. Sci. Pap. ICCAT.
ICCAT. 2005. Report of the 2004 Intersessional Meeting of the ICCAT Subcommittee on
Bycatches: shark stock assessment. Collective Volume of Scientific Papers - ICCAT 58(3): 799-890.
ICCAT. 2008. Report of the 2008 shark stock assessments meeting. Madrid, Spain, 1-5
September, 2008. SCRS/2008/017-SHK Assessment, 89 pp.
ICCAT. 2012. Report of the 2012 shortfin mako stock assessment and ecological risk assessment
meeting. Olhão, Portugal, 11-18 June, 2012.
ICES-WGEF. 2006. Report of the ICES Working Group on Elasmobranch Fishes (WGEF). ICES-CM
2006/ACFM:31. ICES-CM.
ICES. 2012. Report of the Working Group on Elasmobranch Fishes (WGEF). 19-26 June 2012,
Lisbon, Portugal. ICES CM 2012/ACOM:19.
INP. 1999. Tiburones del Pacifico Mexicano, Pesqueria Artisanal. Sustentabilidad y Pesca
Responsible en Mexico, Evaluacion y Manejo. INP/SEMARNAP.
INP. 2000. Sustentabilidad y Pesca Responsable en Mexico, Evaluacion y Manejo, 1999-2000.
INP/SEMARNAP, Mexico.
IUCN. 2009. IUCN Red List of Threatened Species (ver. 2009.2). Available at:
www.iucnredlist.org. (Accessed: 3 November 2009).
Jardas, I. 1996.. Jadranska ihtiofauna. ?kolska knjiga,, Zagreb.
Joyce, W.N. 1999. Management of shark fisheries in Atlantic Canada. In: In: R. Shotton, ed. (ed.),
Case studies of the management of elasmobranch fisheries. FAO, Rome.
Kabasakal, H. 2002. Elasmobranch species of the Seas of Turkey. Annales, Ser. Hist. Nat. 12(1):
15-22.
Klimley, A.P., Beavers, S.C., Curtis, T.H. and Jorgensen, S.J. 2002. Movements and swimming
behavior of three species of sharks in La Jolla Canyon, California. Environmental Biology of Fishes
63: 117-135.
Kosic, B. 1903.. Ribe dubrova?ke, Jazu,, Zagreb.
Levesque, J., 2008: International fisheries agreement: review of the
International Commission for the Conservation of Atlantic Tunas
Case study—Shark management. Mar. Policy. 32, 528–533.
Reproductive biology and population dynamics of the shortfin mako,
Isurus oxyrinchus Rafinesque, 1810, off the southwest Portuguese coast,
eastern North Atlantic
By A. Maia1, N. Queiroz2, H. N. Cabral1, A. M. Santos2 and J. P. Correia. J. Appl. Ichthyol. 23
(2007), 246–251.
Megalofonou, P., Damalas, D., Yannopoulos, C., De Metri, G., Deforio, M., De La Serna, J.M. and
Macias, D. 2000. By catches and discards of sharks in the large pelagic fisheries in the
Mediterranean Sea. Comm. Of the European Communities.
Megalofonou, P., Yannopoulos, C., Damalas, D., De Metrio, G., Deflorio, M., de la Serna, J.M. and
Macias D. 2005. Incidental catch and estimated discards of pelagic sharks from the swordfish
and tuna fisheries in the Mediterranean Sea. Fisheries Bulletin 103(4): 620-634.
Mejuto, J., Garcia-Cortes, B and De La Serna, J.M. 2002. Preliminary scientific estimations of bycatches landed by the spanish surface longline fleet in 1999 in the Atlantic Ocean and
Mediterranean Sea. Col. Vol. Sci. Pap., ICCAT 54(4): 1150-1163.
Mejuto, J., García-Cortés, B. and Ramos-Cartelle, A. 2006. An overview of research activities on
Swordfish (Xiphias gladius) and the by-catch species, caught by the Spanish longline fleet in the
Indian Ocean. IOTC 2006-WPB-11.
Mejuto J., García-Cortés B., de la Serna J. M. and Ramos-Cartelle, A. 2005. Scientific estimations
of bycatch landed by the Spanish surface longline fleet targeting swordfish (Xiphias gladius) in
the Atlantic Ocean: 2000?2004 Period. Col. Vol. Sci. Pap., ICCAT 59(3): 1014-1024.
Mejuto J, GarcıÅLa-CorteÅLs B, de la Serna JM, Ramos-Cartelle A. 2006a. Scientific estimations
of by-catch landed by the
Spanish surface longline fleet targeting swordfish (Xiphias gladius) in the Atlantic Ocean: 2000–
2004 period. Collective
Volume of Scientific Papers of the International Commission for the Conservation of Atlantic
Tunas (ICCAT) 59: 1014–
1024.
Mejuto J, GarcıÅLa-CorteÅL s B, Ramos-Cartelle A. 2006b. An overview of research activities on
swordfish (Xiphias gladius)
and the by-catch species, caught by the Spanish longline fleet in the Indian Ocean, Report No.
IOTC 2006-WPB-11.
Indian Ocean Tuna Commission.
Mejuto, J., García-Cortés, B., Ramos-Cartelle, A. and Ariz, J. 2007. Preliminary Overall
Estimations of Bycatch Landed by the Spanish Surface Longline Fleet Targeting Swordfish
(Xiphias gladius) in the Pacific Ocean and Interaction with Marine Turtles and Sea Birds: years
1990-2005. Inter-American Tropical Tuna Commission Working Group on Bycatch, 6th Meeting
BYC-6-INF A. La Jolla, California, USA.
Mendizabal-Oriza, D., Velez-Marin, R., Marquez-Farias, J. F., Milisic, N. 1994.. Sva riba
Jadranskog mora, Niva,, Split.
Ministry of Fisheries Science Group. 2006. Report from the Fishery Assessment Plenary, May
2006: stock assessments and yield estimates. held in NIWA library, Wellington.
Mollet, H.F. and Cailliet, G.M. 2002. Comparative population demography of elasmobranchs
using life history tables, Leslie matrices and stage-based matrix models. Marine and Freshwater
Research 53(8): 503-516.
Mollet, H.F., Cliff, G., Pratt, H.L., Jr. and Stevens, J.D. 2000. Reproductive biology of the female
shortfin mako Isurus oxyrinchus Rafinesque 1810, with comments on the embryonic
development of lamnoids. Fishery Bulletin 98(2): 299-318.
Moreno, J.;Moron, J., 1992: Comparative study of the genus Isurus (Rafinesque,
1810), and description of form (“Marrajo Criollo”) apparently
endemic to the Azores. Mar. Freshwat. Res. 43, 109–122.
Mourato, B.L., Amorim, A.F. and Arfelli, C.A. In press. Standardized catch rate of shortfin mako
(Isurus oxyrinchus) and bigeye thresher (Alopias superciliosus) caught by Sao Paulo longliners off
southern Brazil. Data Preparatory Meeting of the ICCAT Shark Species Group. 25-29 June, 2007,
Punta del Este, Uruguay.
Muñoz-Chàpuli, R., Notarbartolo di Sciara, G., Seret, B. and Stehmann, M. 1994. The status of
the elasmobranch fisheries in Europe. Report of the Northeast Atlantic subgroup of the
IUCN/SSC Shark Specialist Group, 23pp. Annex 2. In: R.C. Earll and S.L. Fowler (eds), Tag and
Release Schemes and Shark and Ray Management Plans: Proceedings of the Second European
Shark and Ray Workshop. London, UK.
Nakano, H. 1993. A review of the Japanese fishery and research on sharks in the Atlantic Ocean.
ICCAT Collective Volume of Scientific Papers. ICCAT.
Filtering method for obtaining stock indices by shark
species from species-combined logbook data in tuna
longline fisheries
Hideki NAKANO1* AND Shelley CLARKE2 FISHERIES SCIENCE 2006; 72: 322–332
Natanson , L.J., Kohler, N.E., Ardizzone, D., Cailliet, G.M., Wintner, S.P. and Mollet, H.F. 2006.
Validated age and growth estimates for the shortfin mako, Isurus oxyrinchus, in the North
Atlantic Ocean. Environmental Biology of Fishes 77: 367-383.
NMFS (National Marine Fisheries Service). 1993. Fishery management plan for sharks of the
Atlantic Ocean. Technical report, NOAA/NMFS. U.S. Dept. of Commerce. Oceans Program, Islip,
New York.
O?Brian, J.W. and Sunada, J.S. 1994. A review of the southern California experimental drift
longline fishery for sharks, 1988?1991. CalCOFI (Calif Coop Ocean Fish Investig) Rep..
?oljan T. 1948.. Ribe Jadrana, Fauna i flora Jadrana, 1, IOR, Split:.
Orsi Relini, L. and Garialdi, F. 2002. Pups of Lamnid sharks from the Ligurian Sea: morphological
and biometrical characteristics of taxonomic value. In: M. Vacchi, G. La Mesa, F. Serena and B.
Seret (eds), Proceedings of the 4th Elasmobranch Association Meeting 199. Livorno, Italy.
PFMC. (Pacific Fishery Management Council). 2003. Draft Highly Migratory Species (HMS)
Fishery Management Plan (FMP).
Palma, C., Ortiz, M., de Bruyn, P., Kell L. and Pallares, P. 2012. Building a consolidated data-base
to crosscheck ICCAT Task-I nominal catch, against EUROSTAT and FAO equivalent statisticS.
ICCAT SCRS 2012/078.
Pons, M. and Domingo, A. In press. Standardized CPUE of mako shark (Isurus oxyrinchus) caught
by Uruguayan pelagic longline fleet (1981-2006). Data Preparatory Meeting of the ICCAT Shark
Species Group. 25-29 June, 2007, Punta del Este, Uruguay.
Pratt, H.L. and Casey, J.G. 1983. Age and growth of the shortfin mako, Isurus oxyrinchus. In: In:
Prince, E.D. and Pulos, L.M. (eds). (eds), Proceedings of the international workshop on age
determination of oceanic pelagic fishes: Tunas, billfishes, and sharks NOAA Tech. Rep. NMFS 8:
175-177.
Queiroz, N.; Araujo, S.; Ribeiro, P.; Tarroso, P.; Xavier, R.; Santos,
A., 2008: A first record of longfin mako, Isurus paucus, in the
mid-North Atlantic. Mar. Biodiver. Rec. 1, e34.
Reardon, M.; Gerber, L.; Cavanagh, R., 2006: Isurus paucus. In:
IUCN 2012. IUCN Red List of Threatened Species. Version
2012.1. Available at: http://www.iucnredlist.org (accessed on 16
October 2012).
Ribot-Carballal, M.C., Galvan Magaña, F. and Quiñonez Velazquez. 2005. Age and growth of the
shortfin mako shark Isurus oxyrinchus from the western coast of Baja California Sur, Mexico.
Fisheries Research 76: 14-21.
Schrey, A.; Heist, E. 2003. Microsatellite analysis of population structure in the shortfin mako
(Isurus oxyrinchus). Canadian Journal of Fisheries and Aquatic Science 60: 670-675.
Semba, Y., Yokawa, K. and Hiraoka, Y. 2012. Standardized CPUE of shortfin mako (Isurus oxyrinchus) caught by the Japanese tuna longline fishery in the Atlantic Ocean. ICCAT SCRS
2012/074.
Sepulveda, C.A., Kohin, S., Chan, C., Vetter, R. and Graham, J.B. 2004. Movement patterns, depth
preferences, and stomach temperatures of free-swimming juvenile mako sharks, Isurus
oxyrinchus, in the Southern California Bight. Marine Biology 145(1): 191-199.
Serena, F. and Vacchi, M. 1997. Attivita di studio sui grandi pesci cartilaginei dell?alto Tirreneo e
Mar Ligure nell?ambito del programma L.E.M. (Large elasmobranchs monitoring). Quad. Civ.
Staz. Idrobiol. 22(3): 17-21.
Simpfendorfer, C.A., Hueter, R.E., Bergman, U. and Conett, S.M.H. 2002. Results of a fisheryindependent survey for pelagic sharks in the western North Atlantic, 1977-1994. Fisheries
Research 55: 175-192.
Smith, S.E., Au, D.W. and Show, C. 1998. Intrinsic rebound potentials of 26 species of Pacific
sharks. Marine and Freshwater Research 49(7): 663-678.
Soldo, A. and Jardas, I. 2002. Large sharks in the Eastern Adriatic. In: M. Vacchi, G. La Mesa, F.
Serena and B. Seret (eds), Proceedings of the 4th Elasmobranch Association Meeting, pp. 141155. Livorno, Italy.
Soriano-Velasquez, S. R. 2000.. Tiburones Oceanicos del Pacifico..
Stevens, J.D. 1983. Observations on reproduction in the shortfin mako Isurus oxyrinchus. Copeia
1983(1): 126-130.
S_t_e_v_e_n_s_,_ _J_._ _2_0_0_8_._ _T_h_e_ _b_i_o_l_o_g_y_ _a_n_d_ _e_c_o_l_o_g_y_
_o_f_ _t_h_e_ _s_h_o_r_t_f_i_n_ _m_a_k_o_ _s_h_a_r_k_,_ _I_s_u_r_u_s_
_o_x_y_r_i_n_c_h_u_s_._ _I_n_ _‘S_h_a_r_k_s_ _o_f_ _t_h_e_ _O_p_e_n_ _O_c_e_a_n_:_
_B_i_o_l_o_g_y_,_ _F_i_s_h_e_r_i_e_s_ _a_n_d_ _C_o_n_s_e_r_v_a_t_i_o_n_’ _(_M_._D_._
_C_a_m_h_i_,_ _E_._K_._ _P_i_k_i_t_c_h_ _a_n_d_ _E_._A_._ _B_a_b_c_o_c_k_,_
_E_d_s_._)_._ _B_l_a_c_k_w_e_l_l_ _P_u_b_l_i_s_h_i_n_g_,_ _O_x_f_o_r_d_,_ _U_K_,_
_8_7_–9_4_._ _
Stevens, J.D. 1984. Biological observations on sharks caught by sports fishermen off New South
Wales. Australian Journal of Marine and Freshwater Research 35: 573-590.
Stevens, J.D. and Wayte, S.E. 1999. A review of Australia?s pelagic shark resources.
Stillwell, C.E. and Kohler, N.E. 1982. Food, feeding habits, and daily ration of the shortfin mako
(Isurus oxyrinchus) in the Northwest Atlantic. Canadian Journal of Fisheries and Aquatic Science
39: 407-414.
Stone, H., and L. Dixon.
2001. A comparison of catches of swordfish, Xiphias gladius,
and other pelagic species from Canadian longline
gear configured with alternating monofilament and multifilament
nylon gangions. Fish. Bull. 99:210−216.
Strasburg, D.W. 1958. Distribution, abundance, and habits of pelagic sharks in the Central Pacific
Ocean. Fisheries Bulletin 58: 335-361.
Taylor, V.B. and Holts, D.B. 2001.. Shortfin Mako Shark. In: In: Leet, W.S, Dewees, C.M., Klingbeil,
R., and Larson, E.J. (Eds.). (eds), California?s Living Marine Resources: A Status Report,, pp. Pages
336-337.. The Resources Agency, California Department of Fish and Game.
Tortonese, E. 1956. Fauna d?Italia: Leptocardia, Ciclostomata, Selachii. Calderini, Bologna.
Tudela, S., Kai Kai, A., Maynou, F., El Andalossi, M. and Guglielmi, P. 2005. Driftnet fishing and
biodiversity conservation: the case study of the large-scale Moroccan driftnet fleet operating in
the Alboran Sea (SW Mediterranean). Biological Conservation 121: 65-78.
Uozumi, Y. and Nakano, H. 1996. A historical review of Japanese longline fishery and billfish
catches in the Atlantic Ocean. Report of the second ICCAT Billfish workshop. Collective volume
of scientific papers. International Commission for the Conservation of Atlantic Tunas, Madrid,
Miami, Florida, USA.
Vacchi, M., Biagi, V., Pajetta, R., Fiordiponti, R., Serena, F. and Notabartolo Di Sciara, G. 2002.
Elasmobranch catches by tuna trap of Baratti (Northern Tyrrhenian Sea) from 1898 to 1922.
Proceedings of the 4th Elasmobranch Association Meeting: 177-183. Livorno, Italy.
Vannuccini, S. 1999. Shark utilization, marketing and trade. FAO, Rome.
White, W.T., Last, P.R., Stevens, J.D., Yearsley, G.K., Fahmi and Dharmadi. 2006. Economically
Important Sharks and Rays of Indonesia. Australian Centre for International Agricultural
Research, Canberra, Australia.
Yatsu, A. 1995. Zoogeography of the epipelagic fishes in the South Pacific Ocean and the Pacific
sector of the Subantarctic, with special reference to the ecological role of slender tuna,
Allothunnus fallai. Bulletin of the National Research Institute of Far Seas Fisheries 32: 145.
Download