10.1007@s11160 019 09589 5

Rev Fish Biol Fisheries
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Longtail tuna, Thunnus tonggol (Bleeker, 1851): a global

review of population dynamics, ecology, fisheries,
and considerations for future conservation and management
Shane P. Griffiths . Duncan Leadbitter . Demian Willette . Farhad Kaymaram .
Mohammed Moazzam
Received: 9 May 2019 / Accepted: 29 November 2019

Ó Springer Nature Switzerland AG 2019
Abstract Longtail tuna (Thunnus tonggol) is a ner- persistently indicated that at least three of the four
itic species that supports commercial, artisanal and stocks defined in this paper are likely to have been, and
recreational fisheries throughout the Indo-Pacific most likely are currently, subject to overfishing and
region. Historically receiving little attention by com- overfished as a result of excess fishing effort on this
mercial fisheries, the global annual catch of longtail relatively slow-growing and long-lived tuna species.
tuna has steadily risen from around 30,000 t in the As the spawning biomass of principal tuna target
early 1980s to exceeding 200,000 t since 2004, species continue to decline in both the Indian and
reaching a peak of 291,264 t in 2007, and was western and central Pacific Oceans, the increasing
281,613 t in 2017. Catches of longtail tuna in the catches of longtail tuna, other neritic tunas, and
Indian Ocean now exceed catches of principal com- seerfishes is worrisome. Few conservation and man-
mercial target species, such as albacore and bigeye agement measures (CMMs) are currently in place
tunas. A sequence of stock assessments undertaken specifically for longtail tuna, although in recent years
throughout the species’ range since the late 1980s some coastal States, Regional Fishery Bodies, and
tuna Regional Fisheries Management Organisations
have begun to develop initiatives to improve the catch
S. P. Griffiths (&) and biological data quality for longtail tuna and
Inter-American Tropical Tuna Commission, 8901 La Jolla sympatric species of neritic tunas and tuna-like
Shores Drive, La Jolla, CA, USA
e-mail: [email protected] species. This paper provides a global review of
biological, ecological and fishery information to
D. Leadbitter provide researchers, fishery managers and policy
Australian National Centre for Ocean Resources and makers with the most current information from which
Security, University of Wollongong, Wollongong,
NSW 2522, Australia to begin to guide future stock assessment and the
development of CMMs for longtail tuna.
D. Willette
Biology Department, Loyola Marymount University, 1
Keywords Coastal Neritic Northern bluefin tuna
LMU Drive, Los Angeles, CA, USA
Indo-Pacific
F. Kaymaram
Iranian Fisheries Research Science Institute, Tehran, Iran
M. Moazzam
WWF-Pakistan, 46-K, PECHS Block 6, Karachi 75400,
Pakistan
123
Introduction Due to the often-poor reporting of neritic tunas in

some countries, available landings data and perceived
The genus Thunnus within the Scombridae family fishing impacts on the species are likely to be
includes eight species of tunas, some of which support underestimated. Furthermore, because longtail tuna
among the largest and most economically important has not been considered a principal target species by
fisheries in the world. In 2017, the total reported global the Indian Ocean Tuna Commission (IOTC), and has
tuna catch of 5.4 million metric tonnes (t) constituted only recently been covered by the Regional Plan of
7.9% of the world’s total marine finfish catch of 67.9 Action on Neritic Tunas in South East Asia, the
million t (FAO 2019). As the global demand for species has attracted little fundamental research
seafood and animal protein is likely to continue to regarding their biology, stock structure, movements
increase to support an estimated global human pop- or extent of fishery exploitation.
ulation of around 9 billion by 2050 (Msangi et al. Previous reviews of the biology and fisheries of
2013), tuna fisheries are likely to concomitantly longtail tuna were conducted more than two decades ago
increase pressure on the populations of target species, (Jones 1963; Yesaki 1987, 1994), which warned fishery
but also other edible or marketable species caught managers of rapidly increasing catches and a paucity of
incidentally. Increasing removals of these high trophic biological data from which population status assess-
level predators has the potential to initiate imbalances ments could be made to develop appropriate conserva-
in trophic pathways that may eventually generate far- tion and management measures (CMMs). Since those
reaching impacts on the structure and functionality of reviews, reported catches of longtail tuna have steadily
marine ecosystems (Daskalov 2002; Polovina et al. increased—as have several other small sympatric
2009; Griffiths et al. 2019a). species of neritic tunas that support many inshore
One such species that has appeared to have become fisheries of developing countries—with the annual
an increasing target, or at least an increasingly utilised global catch almost tripling from 105,910 t in 1993 to
incidental catch, of artisanal and small- and large- a peak of 291,264 t in 2007 and continuing at over
scale commercial tuna fisheries is the longtail tuna, 208,000 t annually to 281,613 t in 2017 (FAO 2019).
Thunnus tonggol (Bleeker, 1851), which is the second In the Indian Ocean in particular, between
smallest Thunnus species and found throughout trop- 1992–2012 longtail tuna catches increased four-fold
ical and subtropical neritic waters of the Indo-Pacific from 40,580 to 170,221 t. Longtail tuna is now the
region (Froese and Pauly 2017). Longtail tuna has fifth most important species in Indian Ocean tuna
apparently avoided significant attention by industrial fisheries, with the 2017 catch of 144,747 t far
fisheries for several reasons. First, this species nearly exceeding catches of principal commercial tuna
exclusively occupies neritic areas close to landmasses species including bigeye tuna (Thunnus obesus)
and is rarely found beyond continental shelf waters (91,078 t) and albacore (Thunnus alalunga) (38,841
where other tunas can be caught simultaneously t) and constituting 8.9% of the total catch of tunas and
(Yesaki 1994). Second, the species is small in size seerfishes (Fig. 1a). The most important tuna species
(often \ 60 cm fork length) relative to other econom- in Indian Ocean tuna fisheries are skipjack (Katsu-
ically-important species of oceanic tunas and tends not wonis pelamis) (496,600 t in 2017) and yellowfin tuna
to form large dense schools that are conducive for (Thunnus albacares) (416,974 t) but a persistent
capture in commercial quantities by purse-seine or decline in standardised longline catch per unit effort
pole-and-line, and fish often dive when approached by (CPUE) and spawning biomass of yellowfin tuna over
vessels (Yesaki 1994). the past decade has resulted in the stock being subject
However, due to their neritic distribution, longtail to overfishing and overfished for at least the last
tuna are exploited by small-scale commercial and 3 years (Langley 2016b). Tuna fishing fleets in the
artisanal multi-species fisheries in many developed Indian Ocean are therefore beginning to retain other
and developing countries throughout the Indo-Pacific species, such as neritic tunas and seerfishes, presum-
and northwestern Indian Ocean that target neritic tunas ably to improve food security and service the nutri-
including kawakawa (Euthynnus affinis), frigate and tional needs of the large and rapidly-growing human
bullet tunas (Auxis thazard and A. rochei) and populations in developing countries such as India, Iran
seerfishes (Scomberomorous spp.) (Yesaki 1994). and Pakistan (Crist et al. 2017; Techera 2018).
123
Fig. 1 Annual reported catches of tunas and seerfishes in a) the Indian Ocean (FAO Area 61 and 71) and b) the western and central
Pacific Ocean (FAO Area 51 and 57) for 1950–2017 (Data source: FAO 2019). Longtail tuna catches are shown in white
In the Western and Central Pacific Ocean (WCPO), established the Working Party on Neritic Tunas
longtail tuna is the seventh most important tuna (WPNT) to formally recognise the information needs
species, although the 136,866 t reported in 2017 to better manage neritic tunas and tuna-like species in
comprised only 3.7% of the total catch of tunas and the Indian Ocean, which longtail tuna has since been
seerfishes due to the larger catches of skipjack identified as a priority species by the WPNT (IOTC
(1,654,308 t), yellowfin tuna (635,483 t) and seerfishes 2012). This recognition was also reflected in the South
(580,013 t) compared to the Indian Ocean (Fig. 1b). China Sea where growing concerns resulted in the
In the Food and Agriculture Organisation of the development of a Regional Plan of Action for
United Nations’ (FAO) 2011 review of the state of the Sustainable Utilization of Neritic Tunas (RPOA-
world marine fishery resources, Majkowski et al. Neritic Tunas) in 2015 (SEAFDEC 2017).
(2011) raised concerns over the increasing exploita- Given the rapidly increasing catches of longtail
tion of neritic tunas and explicitly highlighted that tuna and need for reliable information to facilitate
‘‘Longtail tuna (T. tonggol) is becoming increasingly stock assessment and guide management and policy
important for canning and the object of substantial development in several regions throughout the Indo-
international trade.’’ Coincidentally in the same year, Pacific region, the aim of this paper is to provide a
the Indian Ocean Tuna Commission (IOTC) comprehensive global review of the biology, ecology,
123
population dynamics, and fisheries for longtail tuna Taxonomic description and evolutionary biology
and identify key knowledge gaps that are essential to
being filled before the status of longtail tuna stocks can Longtail tuna is the second smallest of eight Thunnus
be reliably determined through stock assessment or species and grows to a maximum recorded size of
their inclusion in ecosystem models that may be 145.2 cm total length (TL) (Al-Mamari et al. 2014)
increasing used to facilitate emerging initiatives and 35.9 kg (Froese and Pauly 2017). It has a robust
worldwide to implement ecosystem approaches to anterior tapering to an elongated posterior, hence its
fisheries management. common name of ‘‘longtail’’. The first dorsal fin is
similar to, or slightly smaller than, the second dorsal
fin. The pectoral fins are long (22–31% of fork length)
Methods in fish \ 60 cm and proportionally smaller (16–22%)
in larger fish (Collette and Nauen 1983). Longtail tuna
Literature relating to longtail tuna was derived from a is the only species of the Thunnus genus that lacks a
wide range of sources including scientific journal swim bladder as adults (Serventy 1942b). The ventral
abstracts databases (e.g. Aquatic Sciences and Fish- surface of the liver lacks striations (as for yellowfin
eries Abstracts), Web of Science, Google Scholar, tuna, T. albacares) and the first gill arch has 19–27
individual university and scientific journal websites, rakers (Collette and Nauen 1983).
the Commonwealth Scientific and Industrial Research Body colouration ranges from black on the back to
Organisation (CSIRO) and Inter-American Tropical dark blue above the lateral line with an iridescent blue
Tuna Commission (IATTC) libraries, and the websites band extending along the lateral line from the top
of Regional Fishery Bodies (RFBs), RFMOs, and the margin of the gills to the caudal keel. The flanks are
FAO. Reasonably little information on longtail tuna silver, grading to a silvery-white belly with opaque
has been published in the peer-reviewed literature, so horizontal rows of elongated oval spots, which are
many citations in this review are unpublished reports often absent in larger fish. The first dorsal fin ranges
and other forms of ‘grey literature’, theses, and from dark blue to pale yellow, while the second dorsal
conference, workshop and working group proceedings and anal fins are silvery-white to pale yellow. The
of reliable scientific sources such as IOTC’s WPNT finlets are pale yellow with grey to black margins. The
and the Southeast Asian Fisheries Development tail is black to dark blue, occasionally with a pale-
Center (SEAFDEC) member countries that are yellow colouration at the posterior base of the fork.
involved in the RPOA-Neritic Tunas. The caudal keel is black to dark blue—as opposed to
Fisheries statistics summarised in this paper were bright yellow in southern bluefin tuna, Thunnus
taken from official FAO capture production statistics maccoyii.
using FAO’s freely available software ‘‘FishStatJ’’ Despite the economic and ecological importance of
(www.fao.org/fishery/statistics/software/fishstatj/en) Thunnus species, the evolutionary history of the genus
that was released in November 2019, publicly avail- has been poorly understood until recently; information
able data by the IOTC published in each year of the that can be helpful for management and traceability
WPNT website (http://www.iotc.org/science/wp/ efforts. Using morphological characteristics, Collette
working-party-neritic-tunas-wpnt), and SEAFDEC et al. (2001) partitioned these eight tuna species into
through their online statistical bulletins (http://map. two subgenera, the temperate Thunnus (bluefin group)
seafdec.org/fisherybulletin/). For catch statistics, the containing T. alalunga, T. obesus, T. thynnus, T.
Indian Ocean included Major Fishing Areas 51 and 57, orientalis, and T. maccoyii, and the tropical Neothun-
while the WCPO included Major Fishing Areas 61 and nus (yellowfin group) containing T. albacares, T.
71. All references to fish lengths in this paper are atlanticus, and T. tonggol. This partitioning is reflec-
provided as fork length (FL), unless otherwise noted. tive of the geographic distribution of the two groups
Unpublished tagging data for longtail tuna was and being endothermic. More so, members of the
provided by Australia’s New South Wales Department bluefin group share the presence of additional visceral
of Primary Industries Game Fish Tagging Program and cranial heat-exchanger systems (retia mirabil-
(NSW GTP) (NSW DPI 2007). ia) that heat the brain, eyes and viscera to optimise
foraging and digestion, a feature lacking in the
123
yellowfin group. Ciezarek et al. (2018) found this as ‘‘Dawan’’ and ‘‘Ahur’’ in the languages of Sindh
evolution of Thunnus endothermy to be associated and Balochi, respectively.
with parallel selection and went on to use transcrip- In Australia, longtail tuna was, and in many regions
tomics to infer the eight Thunnus are genetically still is, called ‘‘Northern bluefin tuna’’, especially in
distinct species. These results are in contrast to earlier the recreational fishery. This has caused confusion
exploration of the evolutionary relationship among with Pacific bluefin tuna (T. orientalis) that was long
Thunnus species which challenged the two-subgenera considered a subspecies (T. thynnus orientalis) of
organization. Chow et al. (2006) used the rDNA first Atlantic bluefin tuna (T. thynnus). Collectively, the
internal transcribed spacer (ITS1) to identify mito- latter two species were officially known as ‘‘Northern
chondrial introgression among Thunnus species and bluefin tuna’’ before being separated into two species
found almost identical ITS1 sequences between based on morphology (Collette 1999) and genetics
members of the putative Thunnus and Neothunnus (Tseng et al. 2011). In fact, a major Australian-owned
subgroups. These data inferred a largely unresolved cannery appeared to capitalise on the ambiguity in
phylogeny for Thunnus, a conclusion also found in nomenclature and marketed longtail tuna as northern
sequences from mitochondrial Cytochrome oxidase I bluefin tuna that serendipitously implied the product
(COI) (Mudumala et al. 2011), and Control Region was premium sashimi grade T. orientalis. This
D-loop (Kumar et al. 2016) gene regions. Recently an continued until at least 2002 until the issue was
evolutionary tree for Thunnus was derived using 128 resolved by the Australian Advertising Standards
genome-wide nuclear markers resulting in the reposi- Bureau (ASB 2002).
tioning T. obesus from the bluefin group to within the
yellowfin group, further challenging the two-subgen-
era organization (Dı́az-Arce et al. 2016). Due to its Geographic range
unique evolutionary and morphological traits, Gibbs
and Collette (1967) had long ago positioned T. obesus Longtail tuna inhabit tropical and subtropical waters
as an intermediate of the bluefin and yellowfin groups. of the Indo-Pacific region 47° N–37° S and 32° E–154°
The more recent application of molecular tools has E (Fig. 2). On the easternmost side of their distribution
since better resolved the relationship among the the species can be found as far north as the northeast-
Thunnus as a single monophyletic groups inclusive ern Sea of Japan and throughout the southern Yellow
of eight distinct species (Dı́az-Arce et al. 2016; Sea to China and South Korea (Yoon et al. 2013) and
Ciezarek et al. 2018). as far south as Twofold Bay in southeastern Australia
(Serventy 1942b). On the western side, their distribu-
Common and marketing names tion extends from northwestern Arabian Sea—includ-
ing the Gulf of Oman, and Persian Gulf, the Red Sea—
Longtail tuna is known by several common, local and and southward to the Maputo Province in southern
marketing names, often depending on how the meat is Mozambique (Chacate and Mutombene 2014). Their
prepared for human consumption. The official FAO distribution also includes the Comoros archipelago
common names are longtail tuna (English), Thon (Mohamed Toihir 2018) and Madagascar (Fanazava
mignon (French) and Atún tongol (Spanish). In Japan, 2015). In many parts of their distribution, the
longtail tuna is known as ‘‘Koshinaga’’, while in its geographic range of longtail tuna varies seasonally.
sashimi form it is ‘‘Koshinaga maguro’’. In Malaysia For example, along Australia’s east coast when the
the species is known as ‘‘Aya’’, ‘‘Kayu’’, or ‘‘Tongkol warm waters of the East Australian Current (EAC) are
hitam’’, in Indonesia it is known as ‘‘Tongkol abu abu’’ at their most southerly extent during the Austral
or ‘‘Fufu/Ikan asar’’, while in the Philippines it’s summer and autumn longtail tuna can be found over
known as ‘‘Tambakol’’ or ‘‘Tonggol’’. Similarly, in 1000 km south of their usual distribution (Serventy
some Middle Eastern countries such as the United 1942b). In the South China Sea, commercial catches
Arab Emirates, longtail tuna is also known as ‘‘Tong- and tagging data suggests that the distribution of
gol’’, while in Iran it is known as ‘‘Havoor’’ (Heday- longtail tuna moves northward during the northeast
atifard 2011) and along the Pakistan coast it is known monsoon season, and southward to coastal regions for
the remainder of the year (Raja Bidin 2002).
123
Fig. 2 Map showing the worldwide distribution of longtail tuna and the four putative stocks based on best-available information from
genetic, morphometric and tagging studies
Within their broad geographic range, longtail tuna Stock structure

have quite a unique distribution compared to other
Thunnus species in that they nearly exclusively There is great uncertainty regarding the stock structure
occupy the neritic regime close to landmasses and of longtail tuna throughout their range. Serventy
are rarely found far offshore (Yesaki 1994). They also (1956) was one of the first researchers to explore stock
tend to avoid areas near the mouths of large estuaries structure of longtail tuna and suggested that separate
during periods of rainfall where waters have low stocks—and possibly even two subspecies—may exist
salinity and/or high turbidity (Collette and Nauen in Australian waters based on the distinct difference in
1983). Although they occupy shallow neritic waters, gill raker counts and size distributions of fish present
longtail tuna tagged with electronic tags have occu- off the eastern, northern and western coasts of
pied depths of at least 90 m (Griffiths 2011). There- Australia. Wilson (1981a) tested this hypothesis by
fore, in constructing the geographic distribution of using morphometrics and allozyme electrophoresis
longtail tuna (Fig. 2), we confined its distribution from and found no differences between samples collected
the coast out to the 200 m isobath—based on best from Papua New Guinea, Gulf of Carpentaria, More-
available depth preferences (see ‘‘Habitat specificity ton Bay (Queensland) and Shark Bay (Western
and movement’’)—and extended the depth distribu- Australia). However, he warned that the results need
tion to incorporate reliable occurrence records. to be viewed with caution owing to small sample sizes
from the latter three locations.
123
Abdulhaleem (1989) found significant differences throughout the Savu and Banda Seas and the Sunda
in the number of gill rakers in fish sampled from the Trench that concentrates the strong Indonesian
waters off Oman and the southeastern coast of India Throughflow current between northern Australian
and suggests that this may be indicative of at least two and Southeast Asian waters (Lee et al. 2002), the
separate stocks in the Indian Ocean. possibility of separate stocks of longtail tuna being
Molecular markers have been widely used to present throughout its geographical range cannot be
delineate genetic stocks of marine fishes including discounted. In the absence of large-scale tagging data
tuna, albeit data being limited for smaller species like and detailed genetic analyses throughout this region,
longtail tuna. A review of tuna population structure the extent of mixing of fish between countries or water
literature identified 50 genetically distinct tuna pop- masses remains unknown.
ulations worldwide, with two populations inferred for
longtail tuna (Kumar and Kocour 2015). Two recent
studies used advance mitochondrial DNA (mtDNA) Reproductive biology
displacement loop (D-loop) region genetic analyses to
explore stock structure of longtail tuna in the northern Sex ratio
hemisphere. Kunal et al. (2014) sampled from two
regions in northwest Indian waters and found no Longtail tuna is a gonochoristic species exhibiting no
significant genetic differentiation, suggesting a single signs of sexual dimorphism in external morphology
stock throughout Indian waters. Willette et al. (2016) (Griffiths et al. 2019b). In the nine studies that have
used similar mtDNA D-loop analysis on samples reported sex ratio (Table 1), there is little evidence of
collected from Indonesia, Vietnam, and the Philip- departure from the expected male:female ratio of 1:1
pines to conclude that longtail tuna exist as a single across the entire length range. Griffiths et al. (2010a)
stock within the South China Sea. They compared found a (2:1) bias towards males for large ([ 100 cm
their sequences with those of Kunal et al. (2014) and FL) longtail tuna along the east coast of Australia but
found statistically significant genetic differentiation, speculated this may be due to inadequate sampling of
indicating that fish from the South China Sea and larger fish—due to their naturally lower proportion in
Indian waters comprise two distinct stocks. the population—rather than a true departure from the
It is clear from length-frequency data reported in 1:1 sex ratio they documented in smaller size classes.
studies throughout the distribution of longtail tuna, A similar sex ratio bias towards males has been noted
that there is an increase in fish size with increasing for larger size classes of other Thunnus species
latitude. This appears to be most apparent in Aus- (Hurley et al. 1981; Wild et al. 1995, Schaefer 1998;
tralian waters, where several studies (Serventy Gunn et al. 2008). It has been speculated a sex ratio
1942b, 1956; Wilson 1981a; Stevens and Davenport bias towards males in large yellowfin tuna may be due
1991; Griffiths 2010; Griffiths et al. 2010a) have to higher natural mortality in females as a result of
shown fish to be smallest in the Timor and Arafura higher energetic costs of gonad development and
Seas in northern Australia and gradually increasing in spawning (Schaefer 1998). In contrast, a study of 373
size with increasing latitude southward along both the longtail tuna along the northwestern coast of India
east and west coasts. Very few small fish less than (Mohammed Koya et al. 2018) found a male:female
40 cm have been recorded from Australian waters, ratio of 1:2.1, with the proportion of males declining
whereas fish of this size are abundant throughout with increasing size.
Southeast Asia and support large commercial and
artisanal fisheries. This suggests that there may be a Maturity
southward ontogenetic migration from a northern
nursery ground, such as the South China Sea (see Length-at-maturity has been estimated in only a few
‘‘Reproductive biology’’). As a result, longtail tuna studies, with most reporting the length at first maturity
may exist as a single stock throughout Southeast Asia (LM) rather than the length at which 50% of the
and Oceania. However, considering the significant population is mature (L50), which is a more conserva-
geographic and oceanographic barriers evident tive estimate of maturity and allows the development
throughout this region, such as the complex of islands of maturity-at-length (or age) ogives that are generally
123
Table 1 Summary of parameters describing the reproduction staging (HIS), macroscopic staging (MAC), or a gonadoso-
dynamics of longtail tuna estimated in studies from around the matic index (GSI). Months defining the main spawning season
world, including length-at-first-maturity (LM), length-at-50% in each study are shown
maturity (L50) and batch fecundity based on histological
Country Author Gonad Sex Length Female Batch fecundity Spawning
staging ratio type maturity season
M:F LM L50 Fecundity range Mean
fecundity
Oceania
Australia Serventy (1956) MAC – FL 51.0 – – – Sept–Oct
Australia Griffiths et al. HIS 1:1.3 FL 52.7 53.5 600,215–3,468,350 1,516,680 Sept–Mar
(2019b) (± SD
743,980)
Papua New Wilson (1981a) MAC 1:1 FL 60.0 – 768,000–1,900,000 – Sept–Feb
Guinea
Southeast Asia
Japan Itoh et al. (1999) GSI 1:0.88 – – – – – May–Sept
Taiwan Chiang et al. HIS 1:1.18 FL 37.0 – – – Nov–Dec
(2011)
Thailand Chiampreecha MAC – – [47.0 – – – –
(1978)
Thailand Klinmuang MAC 1:1 – 45–50 – 1,200,000–1,900,000 1,400,000 –
(1978)
Thailand Yesaki (1982) MAC 1:1 – 40.0 43.0 – – –
Thailand Cheunpan (1984) MAC – – 34.2 39.6 – – –
Thailand Hassadee et al. MAC 1:0.97 FL 28.8 42.2 99,773–3,165,849 1,438,583 Feb–Aug
(Gulf) (2014) (± SD
920,715)
Thailand Hassadee et al. MAC 1:0.76 FL 41.0 44.0 44,628–240,477 123,966 April
(Andaman) (2014) (± SD
55,470)
Malaysia Raja Bidin and GSI – – 47.8 – – – May–Aug
Rumpet (1990)
Indonesia Hidayat and MAC 1:0.77 FL 41.1 – – – May–Aug
Noegroho
(2018)
Indonesia Wagiyo and MAC 1:0.85 FL 38.9 – – – Apr–May
Febrianti
(2015)
Eastern Indian Ocean
India Abdussamad MAC – FL 48.0 51.1 227,364–1,092,891 – –
et al. (2012)
India Mohammed MAC 1:2.11 TL 48.0 60.7 – – Dec–Apr
Koya et al.
(2018)
preferred for stock assessment. Again, the lack of length data to estimate L50. Furthermore, some studies
studies that have estimated L50 is most likely due to estimated LM using an arbitrary GSI threshold value to
insufficient sampling across the entire size spectrum of define maturity without histological validation (Che-
the stock, particularly large fish, which is important unpan 1984; Raja Bidin and Rumpet 1990). Therefore,
when fitting logistic regression models to maturity-at- many reproductive studies have produced maturity
123
estimates that are probably not representative of the Timing of spawning

wider population and should therefore be viewed with
caution. Several studies have investigated the timing of
Nonetheless, there appears to be a large difference spawning of longtail tuna in a number of countries,
in LM estimates for females between the northern and primarily using a GSI or macroscopic staging of
southern hemisphere, and between regions within each gonads (Table 1). A common result from these studies
hemisphere (Table 1). In the northern hemisphere, is that spawning occurs over a period of several
only one study (Chiang et al. 2011), undertaken in months during the warmest period of the year in each
Taiwan waters, described maturity based on histolog- region. However, there is an apparent difference
ical examination of 231 ovaries. This study estimated between the northern and southern hemisphere as to
LM to be 37 cm but noted that L50 could not be reliably the seasons when spawning takes place.
estimated as fish larger than 80 cm occur in Taiwanese Yesaki (1982) used macroscopic staging of ovaries
waters since they are rarely caught in the commercial to determine that fish in the waters off the west coast of
or artisanal fisheries from which samples were Thailand spawned during two periods; at the begin-
collected. ning and end of the monsoonal period between
Eight studies conducted in the waters surrounding January–April and August–September. Similarly,
Thailand and Malaysia—including the Andaman Sea, Cheunpan (1984) found two spawning peaks slightly
Malacca Strait, the Gulf of Thailand and the south later in the year between March–May and July–
China Sea—used either a gonadosomatic index (GSI) December in the Gulf of Thailand. Further south in the
or macroscopic staging of ovaries to derive LM South China Sea, Hidayat and Noegroho (2018)
estimates of 28.8–44.0 cm (Table 1). No study used macroscopically staged ovaries to determine fish
reliable histological methods, but three studies (Ye- spawned between May and August. At a similar
saki 1982; Cheunpan 1984; Hassadee et al. 2014) used latitude, Mohammed Koya et al. (2018) also used
macroscopic staging of ovaries to produce L50 esti- macroscopic staging of ovaries to determine longtail
mates of 39.6–43.2 cm for the region. tuna spawned off the coast of India between Decem-
Further west, studies using macroscopic staging of ber–April, with a possible second smaller spawning
ovaries produced quite different LM estimates of peak in June–September, although some mature fish
48 cm for Indian waters (Abdussamad et al. 2012, were found year-round.
Mohammed Koya et al. 2018) and L50 estimates of In the southern hemisphere, there appears to be only
51.1 cm TL and 60.7 cm FL, respectively. one spawning peak in the waters of Australia and
In the southern hemisphere, three studies have Papua New Guinea, primarily during the spring and
examined the reproductive biology of longtail tuna summer, although the spawning period slightly dif-
using either macroscopic staging (Serventy 1956; fered among studies. Serventy (1956) used macro-
Wilson 1981a) or histology (Griffiths et al. 2019b). scopic staging of ovaries to suggest a spawning period
Each study revealed that female longtail tuna mature of September–October in southeastern Australia.
at larger sizes than in the northern hemisphere Wilson (1981a) used macroscopic staging of ovar-
(Table 1). In the waters of Australia and Papua New ies and a GSI to determine longtail tuna had a
Guinea, macroscopic staging of ovaries by Wilson protracted spawning season in Papua New Guinea
(1981a) led to the conclusion that fish first matured at between October and April. Similarly in northern and
51 cm and 60 cm in each respective region. Using eastern Australia, Griffiths et al. (2019b) used histol-
histological analysis of longtail tuna samples from ogy and GSI to determine that longtail tuna have an
northern and eastern Australia, Griffiths et al. (2019b) extended spawning season between October and
found the smallest mature female was 52.7 cm February.
(2.3 years of age), while L50 and the age at 50%
maturity (A50) was estimated to be 53.6 (± 95% CI Spawning locations
46.3–57.0) cm and 2.51 (± 2.14–2.79) years, respec-
tively. However, due to the small sample size of fish The specific spawning locations of longtail tuna is
less than 50 cm, there is the potential for LM and L50 to poorly understood throughout most of its range.
be overestimated. However, based on the capture of ripe females and/
123
or the presence of larvae, possible spawning grounds sample of 106 mature females suggests that the
have been proposed for the southern Andaman Sea majority of longtail tuna may spawn elsewhere. It is
(Puewkhao et al. 2000), the western Sea of Japan and possible they may move to offshore waters to spawn,
the northern region of the East China Sea (Itoh et al. which has been suggested to occur off Malaysia and
1999, Yoon et al. 2013), and the outer neritic zone in Japan (Yesaki 1989; Itoh et al. 1999). In October–
the Gulf of Thailand (Yesaki 1982). Furthermore, the November 2015, twenty small (\ 40 cm) longtail tuna
high proportion of juveniles \ 20 cm in fishery were captured by a commercial fisher in waters of
catches in these regions (Yesaki 1982, 1989; Itoh 100 m depth off Fraser Island, Queensland and fish of
et al. 1996) indicate they are nursery habitats and that a similar size are reputed to occur at the location at a
spawning probably takes place nearby. similar time each year (Dr Julian Pepperell, pers.
In Oceania, the evidence to identify spawning comms.). Further studies are required to explore this
locations is less convincing. Wilson (1981a) hypoth- issue, including larval and catch surveys in offshore
esised that spawning may take place in the vicinity of waters during the spawning season, possibly in areas
Aru Island in the northern Arafura Sea. This was based known to be spawning locations for other Thunnus
on the presence of smaller size classes of fish in this species, such as the Coral Sea (McPherson 1991).
region, compared to the east coast of Australia, and
elevated sea surface temperatures (24–28 °C) that are Batch fecundity
conducive for spawning among Thunnus species
(Schaefer 2001). However, this spawning hypothesis Only five studies have published fecundity estimates
is questionable since no ripe females were captured for longtail tuna (Table 1), all of which describe batch
during 6 years of monthly sampling. Furthermore, the fecundity as opposed to total fecundity. In the southern
smallest fish recorded in his study was 46 cm—around hemisphere, Wilson (1981a) estimated female fish
1–2 years of age—suggesting that these fish may have (75–98 cm) from Papua New Guinean waters produce
been spawned elsewhere and moved to the study 768,000–1,900,000 oocytes per spawning. However,
region. he was unable to obtain samples of ripe females and
Along the southeastern coast of Australia Serventy therefore, these fecundity estimates are probably
(1956) observed longtail tuna having spent ovaries underestimates.
during April and suggested that fish most likely In eastern and northern Australian waters, Griffiths
recently spawned further north—possibly in the et al. (2019b) estimated the fecundity of 15 mature
vicinity of the Great Barrier Reef in the Coral Sea— female fish (68.5–106.3 cm) with ripe ovaries (histo-
before travelling southward with the seasonally logical Stage V) to be 600,215–3,468,350 oocytes,
expanding EAC that reaches its southernmost extent with the average batch fecundity being 1,352,760
in around May each year (Ridgway and Godfrey (± SD 47,642) oocytes. This study also determined
1997). that there was a strong positive relationship between
Recent histological analysis of longtail tuna ovaries fecundity and fish length, indicating that larger
provided conclusive evidence of at least three spawn- females produce a higher number of oocytes. How-
ing locations in Australian waters (Griffiths et al. ever, mean relative fecundity (163.25 ± SD 52.36)
2019b). In this study, monthly sampling was under- oocytes per gram of BW per spawning showed no
taken over a 15-month period across a large region relationship with fish length, implying that all spawn-
incorporating the Timor, Arafura, Coral and Tasman ing females, regardless of size, make a similar
Seas—including the Gulf of Carpentaria—to central contribution to the oocyte production biomass.
New South Wales. Post-ovulatory follicles (POFs) In the northern hemisphere, Klinmuang (1978)
were present in ovaries from fish caught in tropical examined the ovaries of four fish between 44–49 cm
waters in the western Arafura Sea, waters offshore of from the western Gulf of Thailand and off the east
the Edward and Holroyd rivers in the eastern Gulf of coast of Malaysia, and estimated batch fecundity to be
Carpentaria, and in coastal waters inside the Great between 1.2–1.9 million oocytes. In a more recent
Barrier Reef along the central Queensland coast. study in the same region, Hassadee et al. (2014)
Although spawning was confirmed in these regions, examined 14 fish (38.4–49.2 cm) and estimated
the capture of only 8 fish containing POFs from a fecundity to be 99,773–3,165,849 oocytes. However,
123
this study found fecundity estimates to be lower younger and faster growing fish may be missed in
(44,628–240,477 oocytes) in 12 fish (43.0–49.5 cm) subsequent sampling events, leading to inaccurate
sampled from the adjacent Andaman Sea. growth parameters being estimated. Furthermore,
To the west of the Andaman Sea in coastal Indian even where the entire population is represented in
waters, Abdussamad et al. (2012) produced batch samples, older cohorts begin to become increasing
fecundity estimates of 227,364–1,092,891 oocytes for difficult to identify through time, as they begin to
fish between 53.7–79.4 cm. However, the number and merge with other cohorts as their growth rate
the histological stage of specimens used was not decreases and their relative abundance in the popula-
reported and therefore their results should be viewed tion declines.
with caution. Not surprisingly, the VBGF parameters (L?, K and
t0) estimated in studies analysing length-frequency
data vary widely, even for studies conducted in the
Age, growth and longevity same region. For example, in the Gulf of Thailand,
Supongpan and Saikliang (1987) estimated L? and
There are marked regional differences in maximum K to be 58 cm and 1.44, respectively, implying that
size of longtail tuna. For example, fish found through- longtail tuna are fast-growing and short-lived, reach-
out Southeast Asia from the Sea of Japan to the South ing 58 cm by age 5. In the same region, Yesaki (1989)
China Sea appear to be smallest, with a maximum estimated L? and K to be 108 cm and 0.55, respec-
recorded length of 58 cm (Itoh et al. 1999, Mohri et al. tively, suggesting the species is longer-lived, reaching
2010). The species then tends to attain a larger 101 cm by age 5 (Table 2). It is likely that this
maximum size towards the western and southern variability is an artefact of size selectivity of the gear
boundaries of their distribution, with fish larger than from which fish were sampled and the time periods
120 cm frequently recorded in the Gulf of Oman and when sampling was undertaken, as fish move season-
the Persian Gulf (Prabhakar and Dudley 1989; Kay- ally through Southeast Asia (Raja Bidin and Rumpet
maram et al. 2011), and along the southeastern coast of 1990; Chiang et al. 2011).
Australia (Serventy 1942a, 1956; Griffiths Four studies aged longtail tuna using a more
2010, 2012). accurate method of quantifying growth increments in
A number of growth studies have been undertaken sagittal otoliths. These were undertaken in the Sea of
on longtail tuna although their entire geographic Japan (Itoh et al. 1999), the Gulf of Oman (Brothers
distribution, with the majority of studies undertaken 1990), the waters from northern Australia to Papua
in countries where significant commercial and/or New Guinea (Wilson 1981a), and the coastaal waters
artisanal fisheries exist for the species (e.g. Thailand, off the northern and eastern coasts of Australia
Malaysia, India, Iran and Oman). Tracing modal (Griffiths et al. 2010a) (Table 2). The former three
length progressions of cohorts over time using length- studies estimated age by counting presumed daily
frequency data has been the primary method used to increments and suggested that longtail tuna are fast-
estimate growth of longtail tuna. These studies have growing and short-lived. Itoh et al. (1999) aged 33
generally relied on fishery dependent methods, such as small fish (12–49 cm), and estimated L? and maxi-
market sampling or scientific observers onboard mum age as 55 cm and 434 days, respectively.
commercial vessels (Table 2). Unfortunately, length- Brothers (1990) aged only 22 fish (30–82 cm) and
frequency analyses can be an unreliable method for estimated L? to be 78.5 cm. In contrast, Wilson
estimating growth parameters for use in fisheries stock (1981a) aged 26 fish from a broader size range
assessment (Fournier et al. 1998). This is because the (45.3–110.9 cm), estimated an L? of 131.8 cm and
size selectivity of the fishing gear does not allow all concluded that the oldest fish was 1700 days
cohorts present in the population to be properly (4.7 years) old at 110.9 cm. However, the results of
represented in the sample through time, as cohorts these three studies should be viewed with caution.
grow in size. Consequently, growth rates are biased Apart from the small sample sizes, the growth model
towards prominent size classes susceptible to capture parameters would have been biased in each case since
by the gear. A further problem is if the collection of substantial components of the total size spectrum of
samples is not sufficiently frequent, cohorts of the populations were not sampled.
123
Table 2 Summary of von Bertalanffy growth parameters, longevity and length-at-age (in cm) estimated in studies of longtail tuna
where growth was characterised using otoliths or length-frequency (LF) analysis
Area References Ageing Length Sample von Bertalanffy growth Length-at- Longevity
method range (cm) size parameters age (years)
L? K (year-1) t0 1 3 5
(year-1)
Oceania
Australia Serventy (1956) LF – – – – – 38 62 – –
Australia Griffiths et al. Otoliths 24–125 FL 461 135.4 0.233 - 0.020 27 66 91 18.7
(2010a)
Papua New Wilson (1981a) LF 46–103 FL 1477 122.91 0.410 - 0.032 42 87 107 –
Guinea
Papua New Wilson (1981a) Otoliths 45–111 FL 26 131.8 0.395 - 0.035 44 92 114 4.7
Guinea
Southeast Asia
Japan Itoh et al. (1999) Otoliths 12–49 FL 33 55.0 1.700 - 0.089 46 54 55 1.2
Thailand Chiampreecha LF – – – – – 27 45 – –
(1978)
Thailand Klinmuang (1978) LF – – – – – 31 – – –
Thailand Yesaki (1982) LF 20–58 – – – – 30 – – –
Thailand Supongpan and LF – – 58.2 1.440 - 0.027 45 57 58 –
Saikliang (1987)
Thailand Yesaki (1989) LF – – 108.0 0.550 – 46 87 101 –
Malaysia Chiampreecha LF – – – – – 30 – – –
(1978)
Malaysia Raja Bidin and LF 31–50 – 73.5 0.440 – 26 54 65 –
Rumpet (1990)
Indonesia Wagiyo and LF 29–51 168 55.7 1.500 – 43 55 56 –
Febrianti (2015)
Indonesia Restiangsih and LF 18–81 168 85.0 0.400 0.046 27 59 73 –
Hidayat (2018)
Eastern Indian Ocean
India Silas et al. (1986) LF – – 93.0 0.490 - 0.240 42 74 86 –
India James et al. (1993) LF 16–92 – 94.0 0.480 – 36 72 85 –
India Pillai et al. (2003) LF 36–100 – 108 0.550 – 46 87 101 –
India Ghosh et al. (2010) LF 30–98 2976 107.4 0.180 - 0.073 19 46 64 –
India Abdussamad et al. LF 23–111 FL – 123.5 0.510 - 0.032 51 97 113 4.5
(2012)
India Kumar et al. (2017) LF 22–86 – 98.7 0.390 0.335 31 68 84 5.0
Western Indian Ocean
Pakistan Ahmed et al. LF – 300 69.9 0.934 - 0.09 45 66 69 –
(2016)
Iran Kaymaram et al. LF 26–128 4313 133.7 0.350 – 40 87 111 –
(2011)
Iran Yasemi et al. LF 27–107 111.2 0.300 - 0.380 38 71 89 –
(2017)
Iran Darvishi et al. LF 25–124 4383 129.6 0.390 - 0.280 51 94 113
(2018)
Oman Prabhakar and LF 24–118 12333 133.6 0.228 – 27 66 91 –
Dudley (1989)
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Table 2 continued
Area References Ageing Length Sample von Bertalanffy growth Length-at- Longevity
method range (cm) size parameters age (years)
L? K (year-1) t0 1 3 5
(year-1)
Oman Brothers (1990) Otoliths 30–82 22 78.5 0.679 - 0.490 50 71 77 7.0

Yemen Anon (1989) LF – – 104.0 0.250 – 23 55 74 –
In an attempt to resolve some of the sampling In the Gulf of Mannar—between India and Sri
artefacts in previous ageing studies, Griffiths et al. Lanka—Silas (1967) reported that the most important
(2010a) examined the otoliths from 497 fish prey items, in terms of biomass, of 40 longtail tuna
(24–125 cm) and validated annual growth increments (39.5–77.5 cm) were squids (Ommastrephidae), fol-
using daily ageing and edge type analysis. Their study lowed by crustaceans (stomatopods, mysids and
estimated that longtail tuna live for at least 18 years megalopa) and a range of pelagic and demersal fishes
and obtained a similar estimate of L? (135.4 cm) as representing families such as Engraulidae, Clupeidae,
Wilson (1981a) from the adjacent waters of Papua Syngnathidae, and Lutjanidae.
New Guinea. However, their estimate of In contrast, two recent studies conducted in the
K (0.223 year-1) was nearly half that of Wilson coastal waters of the Arabian Sea off western India
(1981a) (K = 0.395 year-1), implying the species is (Abdussamad et al. 2012, Mohammed Koya et al.
slow-growing, and having comparable growth dynam- 2018) found that longtail tuna primarily consumed
ics to bigeye tuna (T. obesus) and southern bluefin teleosts (Sardinella sp., Thryssa sp., Decapturus sp.,
tuna (T. maccoyii), particularly in that these species Selar sp., exoceotids, hemiramphids, Megalaspis
live in excess of 18 years (Griffiths 2010). Earlier cordyla, and Auxis spp.), followed by crustaceans
work by Prabhakar and Dudley (1989) in Omani (penaeids, portunids, and stomatopods), and
waters who analysed length data for 300–500 fish cephalopods.
collected every 10 days per month for nearly 2 years In the Sea of Japan, Kobayashi (2005) examined
also suggested longtail tuna are relatively long-lived, 242 fish (41–60 cm), of which 147 stomachs con-
reaching at least 10 years of age at around 116 cm. In tained prey, and found them to primarily consume
a recent ageing study conducted in adjacent Irani small schooling pelagic fishes (Trachurus japonicus,
waters, Darvishi et al. (2018) analysed monthly Engraulis japonica, and Etrumeus teres), and to a
length-frequency samples (n = 4383) comprising a lesser extent cephalopods.
wide size range (25–124 cm) and also determined that In the Gulf of Papua, Wilson (1981a) found 26 fish
longtail tuna are slow-growing and long-lived, possi- (\ 70 cm) to primarily feed on small pelagic fish from
bly reaching 12 years of age. the families Engraulidae and Scombridae, crustaceans
(Alima, Decapoda and Penaeidae) and cephalopods.
One of the most comprehensive examinations of the
Trophic ecology feeding ecology of longtail tuna was a 2-year study of
497 fish sampled from northern and eastern Australia
Diet composition (Griffiths et al. 2007a). They recorded 101 prey taxa,
with most the common taxa (in terms of biomass)
Longtail tuna plays an important role as both a being: small pelagic fishes (Engraulidae, Clupeidae,
predator and as prey in coastal ecosystems. Although Scombridae, Belonidae and Hemiramphidae), demer-
relatively few studies have quantified the diet compo- sal fishes (Carangidae, Leiognathidae and Sil-
sition of longtail tuna, available data indicates they are laginidae), cephalopods (Teuthoidea and Sepia spp.)
an opportunistic predator consuming a wide range of and crustaceans (Portunidae, Penaeidae and Squilli-
prey types. dae). The study showed a distinct ontogenetic shift in
123
feeding behaviour and diet after fish attained about Foraging behaviour and ecological role as prey
100 cm.
The diets of longtail tuna frequently contain From two major longtail tuna research programs
demersal and benthic prey, most likely facilitated by undertaken in Thailand (Yesaki 1982) and Australian
their occupation of shallow coastal waters, that waters (Griffiths et al. 2007a), a number of observa-
provides an important trophic linkage between pelagic tions were made to describe the species’ feeding
and demersal components of the ecosystem, which behaviour and their predation by larger predators. For
differs from the role that larger tuna play in the example, juvenile and small adult fish (\ 70 cm)
trophodynamics of open ocean systems. For example, tended to form large schools and occupied an inter-
longtail tuna consumed fishes representing the fami- mediate trophic level by primarily consuming small
lies Leiognathidae, Mullidae, Gerridae, Gobiidae, schooling baitfishes such as Sardinella spp., Stole-
Nemipteridae and Callionymidae in the waters of phorous spp. and Thyrssa spp., and a wide variety of
Australia and Papua New Guinea (Serventy epipelagic crustaceans such as penaeids and Por-
1942a, 1956; Wilson 1981a; Griffiths et al. 2007a), tunus spp. Fish of this size often form ripples on the
while 76% of the diet biomass of 112 fish (32–61 cm) surface when feeding, but they rarely leap from the
examined from Malaysian waters in the South China water (Yesaki 1987). At these sizes, longtail tuna were
Sea comprised Monacanthus spp. (Bachok et al. often associated with other small neritic tunas such as
2004). Off the coast of India, longtail tuna also kawakawa and frigate and bullet tunas. Together,
consumed benthic species including Platycephalus these neritic tunas are important prey for larger
spp. (Mohammed Koya et al. 2018), while in the Sea predators. Although there are apparently no docu-
of Japan they often consumed the demersal Apogon mented scientific identifications of longtail tuna in the
semilineatus (Kobayashi 2005). diet studies of other predators, there are numerous
Interestingly, plastic materials have been docu- anecdotal accounts from scientists and fishers who
mented in the diets of longtail tuna sampled from the have witnessed the predation of longtail tuna. For
waters around Australia, Indonesia, and India. Grif- example, small (\ 60 cm) longtail tuna were observed
fiths et al. (2007a) found three plastic drinking straws in northern Australia to be predated upon by Spanish
in the stomachs of longtail tuna in Moreton Bay, mackerel (Scomberomorus commerson), billfish (Is-
Australia—located adjacent to one of Australia’s tiompax indica and Istiophorus platypterus) and
largest cities. Wagiyo and Febrianti (2015) and various carcharhinid sharks.
Mohammed Koya et al. (2018) did not report the type Larger longtail tuna ([ 100 cm) are less commonly
of plastic materials ingested by longtail tuna in observed as surface aggregations and appear to feed in
Indonesian and Indian waters, but in terms of smaller schools (15–20 fish), and often as solitary
frequency of occurrence they comprised 1% and 8% individuals, attacking their prey in arrow formation
of the diet, respectively. However, Mohammed Koya with fish being equidistant to each other. In these
et al. (2018) described floating plastic debris aggre- cases, some fish have been observed leaping from the
gating small fish species that were commonly found in water. Fish of this size are near-apex predators in
the stomachs of longtail tuna, and therefore ingestion coastal ecosystems, such as Australia’s Gulf of
of plastics is probably accidental whilst pursuing prey. Carpentaria where they have an estimated trophic
Nonetheless, with the increasing incidences of plastics level of 4.62 (Okey et al. 2007). By comparison,
in the marine environment (Cózar et al. 2014), oceanic tunas that support commercial tuna fisheries in
primarily from land-based sources, neritic predatory a similar region of the western Pacific Ocean occupy
species such as longtail tuna, may be increasingly higher trophic levels, such as bigeye (4.93) and yel-
vulnerable to ingesting these plastics as they are lowfin (4.78) tunas (Griffiths et al. 2019a).
transported by rivers and estuaries into coastal habi-
tats, and further offshore where they have been found Prey consumption rates
to be ingested by oceanic tunas such as yellowfin tuna
(T. albacares) (Chagnon et al. 2018) and southern Longtail tuna are visual predators and primarily feed
bluefin tuna (T. maccoyii) (Young et al. 1997). during the day, but feeding has been documented to
occur during the night, but to a far lesser extent
123
(Griffiths et al. 2007a). Feeding intensity has also been (86–122 cm) with miniature pop-up archival tags
shown in both Australian (Griffiths et al. 2007a) and (‘‘miniPAT’’, Wildlife Computers) along the northern
Indian (Mohammed Koya et al. 2018) waters to have half of Australia’s east coast. From the 494 days of
an inverse relationship with reproductive activity, data collected by the tags, fish dived to a maximum
indicating a possible energy investment for gonad depth of 90 m and occupied water temperatures of
development. 17.1–28.9 °C. However, tagged fish primarily occu-
Prey consumption rates and daily ration of longtail pied depths of less than 30 m and water temperatures
tuna have been estimated by one study in Australian of 20–28.5 °C (Fig. 3). Geolocation of tagged fish—
waters (Griffiths et al. 2007a), based on stomach estimated from the tag’s archived light data—revealed
content biomass and estimated prey evacuation rates. fish did not move beyond the continental shelf,
The study found that daily ration averaged 2.36% body confirming the species’ presumed preference for
weight (BW) day-1 but decreased with increasing fish neritic waters. The data highlights the marked differ-
size from 2.26% to 1.3% BW day-1 for fish \ 100 cm ence in movement and behaviour compared to other
and [ 100 cm, respectively. In a concurrent study, Thunnus species, particularly the absence of diving at
Griffiths et al. (2007b) used commercial gillnet catch dawn to remain at depth during the day and moving to
data to estimate a biomass of 12.07 kg km-2 for surface waters during the night (Schaefer and Fuller
longtail tuna in the Gulf of Carpentaria (397,700 km2). 2002; Schaefer et al. 2007, Patterson et al. 2008). The
Using the longtail tuna biomass estimates coupled restricted vertical movements of longtail tuna may
with the daily ration estimates, Griffiths et al. (2007a) explain its lack of a swim bladder as a possible adap-
estimated that 148,178 t year-1 of prey were con- tation for living in shallow neritic waters.
sumed annually in the Gulf of Carpentaria. Together, Very little information is available on the horizon-
these studies demonstrated the important ecological tal movements of longtail tuna. The historical low
role that longtail tuna play in Australian neritic commercial value of longtail tuna in comparison to
ecosystems and are likely to play a similarly important other commercially important tunas is likely the
role in other neritic ecosystems throughout their global primary reason for the lack of a dedicated long-term
distribution. tagging programs, and as a result, little is known of
their movements throughout their distribution, with
the exception of a few of opportunistic tagging studies.
Habitat specificity and movement Raja Bidin (2002) tagged 8842 longtail tuna
(15–47 cm) in Malaysian waters in the South China
Longtail tuna have been documented occupying water Sea over a period of 8 years. Unfortunately, only 19
temperatures of 17–30 °C (Griffiths 2011, Mohri and tagged fish (23–46 cm) were recaptured, all of which
Yoritake 2014). The optimal water temperature range were recaptured less than a year later within 180 nm of
for fish \ 50 cm has been suggested to be 24–25.6 °C the release locations. In adjacent Indonesian waters,
based on modelling of commercial catch data in Kaltongga (1998) tagged 75 small longtail tuna
relation to remotely-sensed sea surface temperature (52–58 cm), but no recaptures were reported.
(SST) data in the Sea of Japan (Mohri et al. Wilson (1981a) recaptured 25 of the 414 fish tagged
2005, 2008). In Australia, Griffiths (2012) showed a in the Gulf of Papua over a 2-year period. Although no
seasonal spatial shift in recreational fishing effort for information was reported on distances moved, all
large longtail tuna ([ 80 cm), which coincided with recaptures were made within the Gulf (* 350 km
the southward movement of the EAC when SSTs were wide) after being at liberty for 76–103 days.
18–22 °C, indicating that large longtail tuna probably Australia’s NSW GTP has recapture information
prefer this range of SSTs. for 57 of the 3838 longtail tuna tagged by recreational
In the first study of the habitat utilisation and fishers since 1974 (NSW DPI 2007). The movement
movement of longtail tuna using electronic tags, data reveals that the species can move large distances
Griffiths (2011) tagged nine longtail tuna in short periods. For example, the fish moving the
123
Fig. 3 Percentage of time that nine longtail tuna (86–122 cm) tagged with pop-up archival satellite tags off eastern Australia spent at
temperature (°C) and depth (m). Data summarised from Griffiths (2011)
longest recorded distance was tagged in Moreton Bay, distances, with two fish occupying Hervey Bay for
Queensland and was recaptured * 600 km to the the entire 3-month duration of the tag deployments.
south 24 days later (Fig. 4). The tagging data reveals
that there may be a seasonal movement of fish as most
fish tagged in Queensland waters that moved south Fisheries
into NSW waters were recaptured between March and
May when the EAC is at its maximum southern extent Longtail tuna is an important resource that is exploited
(Ridgway and Godfrey 1997). Interestingly, the data in the waters of mainly developing countries bordering
also reveals that movement of fish tagged at some the western Pacific Ocean across to the northern Indian
locations can be very limited, particularly within large Ocean. As a result of their coastal distribution, longtail
marine embayments such as Moreton Bay and Hervey tuna are caught by small-scale commercial and
Bay. For example, of the 36 fish recaptured after being artisanal fisheries in at least 21 countries throughout
tagged in Moreton Bay, only four fish were recaptured these regions (FAO 2019) primarily using purse-seine,
outside of the bay, despite being at liberty for up to gillnet, and hook and line (e.g. trolling), but also a
3833 days. This may either reflect permanent resi- variety of other minor methods including beach seine,
dency within the bay or that the region is an annual stake traps, and set nets (Boonragsa 1987). Longtail
visiting site during their presumed southward move- tuna are caught in two broad fishery types differenti-
ment during summer and autumn (see Serventy 1956). ated by geography, the WCPO and the Indian Ocean,
In contrast, all nine fish tagged with miniPAT where the predominant gear used differs. Although
electronic tags by Griffiths (2011) moved north for longtail tuna comprise the majority of catches in the
linear distances of up to 650 km. Two tagged fish - coastal tuna fisheries of both regions, they are not
moved 450 km and 650 km north to a common area often targeted specifically. In the WCPO, longtail tuna
inside the Great Barrier Reef at times where spawning constitutes a major component of the catches from
has been previously presumed to take place in multi-species fisheries for small neritic tunas and
October–March (Serventy 1956; Griffiths et al. seerfishes, which primarily include kawakawa, frigate
2019b). The remaining fish moved much shorter and bullet tunas, and Spanish (or King) mackerels
123
Fig. 4 Release and recapture locations of seven tagged longtail Industries’ Gamefish Tagging Program in Australian waters.
tuna that showed the largest movements among 55 recaptured Month of recapture and number of days at liberty is shown
fish from the New South Wales (NSW) Department of Primary adjacent to each movement path
(Scomberomorous spp.) (Yesaki 1994). In the Indian WCPO fisheries

Ocean, longtail tuna are generally larger and are
caught in other multi-species drift gillnet fisheries that The major fisheries for longtail tuna in the WCPO
target larger oceanic tunas (e.g. yellowfin tuna) and operate in Southeast Asia, primarily the Gulf of
sharks—particularly in Iran, Pakistan and India Thailand and the South China Sea where small fish
(MRAG 2012; Gerami and Dastbaz 2013). (15–55 cm) are primarily caught by purse-seine and
drift gillnet, and to a lesser extent by hook and line
(primarily trolling, but also includes handline and
123
small longlines), particularly in Malaysia, Thailand, by up to 10 crew members, who undertake trips of
Vietnam, and Indonesia (Raja Bidin and Rumpet 1–4 days in duration (Yonemori et al. 1995). Gillnets
1990; Kamarruddin and Raja Bidin 1991; Chullasorn usually consist of 60–120 mm stretched monofilament
1995; Yonemori et al. 1995, Lewis 2006; Nootmorn mesh of 1–12 km in length and around 7–50 m in
2015; Siriraksophon 2017). depth, depending on the location (Boonragsa 1987;
The purse-seine vessels and gear used in this region Chullasorn 1995; Yonemori et al. 1995).
range in their sizes depending on the types of sets The hook and line (i.e. trolling) vessels are much
intended to be made, such as those on fish aggregating smaller in size (7–18 m LOA)—often fitted with
devices (FADs) or free-swimming tuna schools during outboard engines (16–55 hp)—and make trips of
the day, or in association with luring light devices (e.g. 1–15 days in duration (Yonemori et al. 1995, Tam-
electric lamps) during the night (Yonemori et al. pubolon et al. 2015). Given the small size of the
1995). vessels, fish are kept on ice until returning to port.
In general, vessels range from 18–24 m length Each boat has a crew of 1–5 fishers who generally
overall (LOA) are crewed by 25–45 people and make deploy up to 12 trolling lines, some with branchlines,
short trips of around 1–4 days in duration (Boonragsa fitted with various types of lures or baits (Merta 1987).
1987; Merta 1987; Yonemori et al. 1995). The vessels Some trolling techniques such as the ‘‘Rintak’’ (trans-
are generally equipped with radar, echo sounders, and lating to ‘‘thousand lines’’) used in southern Indonesia,
satellite navigational instruments (Chullasorn 1995). appear more like a longline, with 100–120 branchlines
These vessels deploy nets having a stretched mesh size fitted with lures that are attached to a mainline
of 25–97 mm and range in length and depth of (Tampubolon et al. 2015).
500–1600 m and 50–150 m, respectively (Chullasorn It is also noteworthy that longtail tuna are caught in
1995; Chantawong 1999). The smaller mesh sizes are reasonable quantities in unique multi-species set net
primarily used for targeting small pelagic fishes (e.g. fisheries in coastal waters of Japan (Nakamura 1969;
Indian mackerel, Rastrelliger kanagurta), whereas the Mohri et al. 2010, 2013) and Taiwan (Chiang et al.
larger mesh sizes are used to target tunas, including 2011), where complex net arrangements set adjacent
longtail tuna. In recent years, many purse-seine to a major current corral migrating fish into a central
vessels have installed power blocks to reduce labour net pocket (Chiou and Lee 2004).
and to enable the use of longer and deeper nets The composition of the WCPO longtail tuna catch
(Chullasorn 1995). by gear type has changed considerably over the history
In Thailand, small size classes of longtail tuna are of the fishery. When disaggregating the catch into
caught in purse seine operations that target small three separate time periods, each of 15 years
pelagics, but they comprise a very minor component (1972–1986, 1987–2001, 2002–2016), purse-seine
(0.39–1.9%) of the catch (Khemakorn and Vibunpant attributed 60.6%, 81.3% and 88.0% to the WCPO
2008). The main catches are in tuna nets using larger catch, respectively, while the contributions by gillnet
mesh sizes that take larger fish (Nootmorn and and hook and line decreased considerably (Fig. 6).
Tossapornpitakkul 2013). Interestingly, longtail tuna
do not associate with FADs in the way that oceanic Indian Ocean fisheries
tuna species such as skipjack and yellowfin tuna do.
Consequently, longtail tuna is not a target species as In contrast to the WCPO fisheries that catch longtail
this fishery sets on FADs targeting small pelagics. tuna, the fisheries in the Indian Ocean generally catch
Instead, longtail tuna are targeted in the early morn- larger fish (60–80 cm) primarily using drift gillnets
ings when they generally school near the surface and to a lesser extent purse-seine—especially in the
where they can be detected by sonar. waters of Thailand and Malaysia—and hook and line
The second method that accounts for a large (trolling). These catches are usually made in coastal
proportion of the longtail tuna catch in the WCPO is regions of the northwestern Indian Ocean from the
drift gillnet, often known as Spanish (or King) west coast of India to Somalia, particularly in coun-
mackerel drift gillnets, since the primary target species tries bordering the Gulf of Oman, Persian Gulf, Red
are Scomberomorus spp. (Chullasorn 1995). Drift Sea and the Gulf of Aden (Prabhakar and Dudley
gillnet vessels are often 14–18 m LOA and operated 1989; Kahraman et al. 2011). Off the west coast of
123
Malaysia, longtail tuna is primarily a bycatch in the In contrast to the WCPO, purse-seine has accounted
small pelagics fishery that uses lights and/or FADs, as for comparatively little (18–19%) of the longtail tuna
is the case in Thailand. Between 2006 and 2011, catch over the history of the neritic tuna fishery, while
neritic tunas accounted for 9% of the catch in the Strait hook and line (primarily trolling) has also accounted
of Malacca, with longtail tuna accounting for just over for a minor component (9–12%) of catches (Fig. 6).
half of the landings (Basir and Jamon 2012). Over the There appears to be little country-specific published
history of the neritic pelagic fishery in the Indian information on the configuration of vessels and the
Ocean, the methods used that account for the majority gear used for the purse-seine and hook and line
of the longtail tuna catch has remained very consistent, fisheries in the neritic tuna fisheries in the western
with drift gillnet being responsible for 18.8%, 19.0% Indian Ocean. However, there is some basic informa-
and 18.2% of catches for three time periods tion available on Iran’s and Oman’s purse-seine and
(1972–1986, 1987–2001, 2002–2016), respectively. troll fisheries that operate in the Gulf of Oman and the
Much of the drift gillnet catch of longtail tuna is Persian Gulf that account for a significant component
contributed by Iran, Pakistan and India, from vessels of the longtail tuna catch in the western Indian Ocean.
of 10–30 m LOA and 1–1.5 t capacity that are Thai purse-seining is carried out in the same manner as
generally equipped with iced fish holds, hydraulic for the South China Sea, from purse-seine vessels
net haulers, GPS, and echo sounders (Moazzam and ranging in length from 55–99 m LOA (Moradi 2015)
Nawaz 2014). The gillnets are made of polyamide or and equipped with modern electronics including GPS
polyethylene monofilament material of 90–170 mm and echo sounders (Parsa et al. 2018). The vessels are
stretched mesh with a length and depth of crewed by 30–35 people undertake voyages of
4.83–11.27 km and 14 m, respectively (Pillai et al. 2–4 weeks in duration (RECOFI 2013b). The typical
2003, Gerami and Dastbaz 2013; Moazzam and configuration of the purse-seine net is similar to gear
Nawaz 2014). The nets are generally set in the evening used in the WCPO in terms of length (1886 m) and
and retrieved during the early morning. depth (210 m), but the net has a much smaller
The drift gillnet fishery has a long contentious stretched mesh size of 16–20 mm (Hosseini and
history with conservation groups since the relatively Ehsani 2014; Parsa et al. 2018).
unselective gear has very high catch rates of a variety The hook and line fishery for neritic tunas in the
of bycatch species, including turtles, cetaceans, and northwestern Indian Ocean almost exclusively
elasmobranchs. In a recent study of the bycatch of involves trolling from outboard-powered vessels of
Pakistan’s tuna drift gillnet fishery, it was estimated less than 15 m LOA, where 3–4 crew operate single
that 28,000 sea turtles and 12,200 cetaceans were day trips in coastal waters and fish kept in ice until
caught annually between 2011 and 2013 (Moazzam returning to port (RECOFI 2013c; Kakoolaki 2017).
and Nawaz 2014). However, in order to reduce the Some of the larger vessels venture further offshore,
entanglement and mortality of cetaceans and sea equipped with freezers or ice holds to facilitate trips of
turtles, WWF-Pakistan promoted the introduction of 1–2 weeks in duration (RECOFI 2013a). The specific
subsurface gillnetting in 2015—wherein the gillnets configuration of the troll gear is not well documented,
are set 2 m below the surface—that is now adopted by however, it is likely to be similar to trolling methods
the entire gillnet fleet of Pakistan. This has resulted in used to target neritic tunas in Southeast Asia, which
major decrease in entanglement of sea turtles to consist of several lines and hooks. Yesaki (1994)
around 2,700 per year in 2015–2016, while for suggested that one line with a single hook, most likely
cetaceans, entanglement decreased to 480 and 280 a lure, is fished from the stern of the smaller vessels in
individuals in 2016 and 2018, respectively (Moazzam the north Arabian Sea.
and Nawaz 2017; Kiszka et al. 2018). By comparison,
Iran’s drift gillnet fisheries were estimated by Ander- Commercial catches
son (2014) to catch between 24,694 and 101,345
cetaceans per year—based on the catch rates reported India, Indonesia, Madagascar and the United Arab
by Yousuf et al. (2009) and Leatherwood (1994), Emirates were the first countries in FAO records to
respectively. report catches of longtail tuna, with the total reported
catch being 796 t in 1950 and increasing to 2,038 t by
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1960. By 1987, 11 countries reported landings of longtail tuna in their EEZs since at least 2009
longtail tuna, taking a combined annual catch of (Akhondi 2012; Al-Kiyumi et al. 2014, Moazzam
90,122 t (Fig. 5a). The year 1988 marked the begin- 2014), since Iranian and Pakistani vessels have dual
ning of a significant change in the catch trend that registration in both countries (Moazzam 2012b). In
indicated a possible focus on targeting longtail tuna, Iran alone, this resulted in an increase in longtail tuna
with the annual reported catch almost doubling to catch from 25,000 to 81,000 t between 2006 and 2011
168,330 t in just 1 year. This was primarily due to (Akhondi 2012). Over the past 5 years, Indonesia,
significant increases in the reported catches by the Iran, Malaysia, Thailand, Oman, Pakistan and India
large-scale purse-seine fisheries in both Taiwan have collectively contributed 98% to the global
(7,080 t to 26,120 t) and Thailand (39,180 t to reported catch (Fig. 5a).
92,925 t) in the WCPO, which persisted until 1992 In Oceania there is no significant commercial
(Fig. 5c). These catch increases roughly coincided fishery for longtail tuna, despite their abundance and
with steady increases in annual catches in the Indian large size. Although FAO statistics reveal no com-
Ocean, particularly by Iran and Oman, that averaged mercial longtail tuna catches being reported for Papua
around 46,000 t (Fig. 5b). New Guinea—the easternmost extent of the distribu-
The total global catch continued to increase through tion of longtail tuna—Wilson (1981a) reported a small
the late 1990s but rapidly began to accelerate from commercial fishery consisting of up to 26 vessels
2002 to in excess of 208,000 t per year and reaching a existed in the Gulf of Papua in the mid-1970s that
peak 291,264 t in 2007 (Fig. 5a). Again, this was due generally caught less than 100 t per year. He also
to a combination of increased catches in both the described a small and seasonal (November to April)
WCPO and Indian Ocean. In the WCPO, this was artisanal and subsistence fishery that operates in the
mainly due to the commencement of catch reporting Gulf of Papua that was estimated to take less than
by Indonesia catching 70,735 t in 2004 and annually about 10 t of longtail tuna annually. No commercial or
averaging 72,005 t (± SD 24,719) thereafter until artisanal catches have been reported for other nearby
2017 (Fig. 5c), which was primarily caught by purse- Pacific Island Nations, such as the Solomon Islands,
seine (Fig. 6). However, it is important to note that Vanuatu, Fiji, New Caledonia, or New Zealand.
Indonesia’s sudden catch increase may be a possible In Australia, longtail tuna were fished since at least
artefact of misidentification and misreporting. In 1897 (Serventy 1956), but catches between
Malaysia and Indonesia, longtail tuna is locally known 1950–2014 have averaged about 34 t and peaking at
as ‘‘Tongkol abu abu’’, while small-sized tunas— just 138 t in 1985 (FAO 2019). These low catches in
including neritic species such as longtail tuna, Australia generally reflected a historic low domestic
kawakawa, frigate and bullet tunas, as well as juvenile market demand for tuna. Catches by Australian
oceanic tunas including yellowfin, bigeye and skipjack flagged Taiwanese gillnet vessels—managed by the
tunas—are collectively referred to as ‘‘Tongkol’’ Australian Fisheries Management Authority (AFMA)
(Ingles et al. 2008). under a temporary bilateral agreement in the Aus-
In the Indian Ocean, the dramatic increase in tralian Fishing Zone (AFZ)—off northern Australia
catches from the early 2000s was primarily due to between 1974–1986 ranged between 200–2000 t per
increasing drift gillnet catches by Iran and a number of year whilst primarily targeting sharks and narrow-
coastal States in the western Indian Ocean, but also barred Spanish mackerel (Scomberomorus commer-
India, Malaysia and Thailand fishing in the Andaman son) (Stevens and Davenport 1991). Although there
Sea (Fig. 5). The increase in catches by Iran and has been various proposals over the past few decades
Pakistan has been attributed to the increase in Somali to develop a commercial neritic tuna fishery in
piracy that intensified from around 2008 in the western northern Australian waters (Robins 1975; Wilson
Indian Ocean, primarily in Somali waters and adjacent 1981b; Lyle and Read 1985), the high incidental
Areas Beyond National Jurisdiction (ABNJ). This mortality of cetaceans caught by the Taiwanese gillnet
forced a significant proportion of the effort by large- operations in the AFZ—estimated to be * 14,000
scale industrial tuna fleets that would normally fish animals in 1981–1985 (Harwood and Anderson
beyond their national EEZs in the northwestern Indian 1987)—has since halted any subsequent
Ocean to retreat to more coastal waters to target
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Fig. 5 Annual reported catches of longtail tuna by country Ocean (Data source: FAO 2019). Catches for all other countries
having the highest catches for the years 1950–2017 a) globally, are combined in the ‘‘other’’ category
b) in the Indian Ocean, and c) in the western and central Pacific
123
Fig. 6 Percentage of total catches by gear type during three Central Pacific Ocean (Data source: SEAFDEC 2018). Number
time periods (1976–1986, 1987–2001, 2002–2016) for the of gear types included in the ‘‘other’’ category shown in
Indian Ocean (Data source: IOTC 2018) and the Western parentheses
considerations for the development of a similar exceeded the official landings reported to the FAO
domestic drift gillnet fishery. for all fisheries combined (29,800 t) in 2010. They
It is important to note that the landings data estimated that 5% of the artisanal catch was composed
presented in this paper, sourced from RFMOs, RFBs of longtail tuna (* 1,600 t), which is not currently
or national fisheries agencies, are likely to suffer from included in FAO catch statistics. Similar catch recon-
a number of significant shortcomings. Firstly, because structions for Pakistan’s commercial tuna drift gillnet
longtail tuna comprise important artisanal and subsis- fishery revealed that the reported catch of longtail tuna
tence fisheries in many developing countries, where in 2015 (* 7000 t) was 58% lower than the recon-
vessel logbook reporting is either not in place or structed catch (* 15,000 t) (Moazzam and Ayub
strictly enforced, reported landings from these coun- 2017).
tries are likely to be underestimated. This issue is Catches may also be underestimated in some
significant for countries such as Somalia, where no countries due to non-reporting or under-reporting for
compulsory reporting of catches is required and various political reasons. For example, Taiwan his-
validation of catches by domestic or international torically reported annual catches of several thousand
fisheries agencies is difficult and dangerous (IOTC tonnes to SEAFDEC and the FAO since at least 1986,
2015; Persson et al. 2015). In a recent reconstruction but annual catches abruptly dropped to zero after
of Somali catches, Persson et al. (2015) estimated that 2003, with the exception of 2980 t being reported in
the catch by the artisanal fleet alone (32,730 t) 2009 (Fig. 5c), suggesting Taiwan may no longer
123
completely report their catches. In a reconstruction of bluefin tuna (T. orientalis) and/or yellowfin tuna (T.
Tawian’s fishery catches from 1950–2007, Kuo and albacares) and the species complex is commonly
Booth (2011) estimated that around 400,000 t of catch reported in different regions as ‘‘Shibi’’ (Itoh et al.
was unreported since 2003. The majority of this 1996) or ‘‘Yokowa’’ (i.e. ‘‘juvenile tuna’’) (Mohri
unreported catch was believed to be from coastal et al. 2008). Interestingly, no catches of longtail tuna
fisheries, including the set-net, gillnet, and small appear in official FAO statistics for Japan, despite the
purse-seine fisheries that are responsible for the fact they are frequently caught throughout the East
majority of the catch of neritic tunas in Taiwan (Chiou China Sea and the Sea of Japan in quantities in the
and Lee 2004). order of at least hundreds of tonnes in the set net
Similarly in Australia, despite the Australian- fishery alone (Itoh et al. 1996, Kawatsu et al. 2011). It
flagged Taiwanese gillnet fleet reporting annual land- is possible that similar reporting and species identifi-
ings of up to 2000 t between 1976 and 1984 (Stevens cation issues are evident in other coastal States where a
and Davenport 1991), the catch statistics for this mixture of neritic and oceanic tunas are caught, or at
period are still not appear to be accurately reflected in least brought together at central landing sites.
FAO statistics. More recently, longtail tuna are still
captured in large numbers in northern Australia’s Recreational fisheries
State-managed offshore and inshore shark and mack-
erel (Scomberomorous spp.) gillnet fisheries, however Longtail tuna are important to recreational (sport)
they are generally an unreported discarded bycatch. fisheries in a small number of countries throughout
Misidentification and mis-reporting of longtail tuna their range, particularly in Australia, Oman, Kenya,
has been a fundamental problem in some countries, Mozambique, Pakistan, Thailand, Malaysia and
especially throughout Southeast Asia. It has only been Indonesia (Wekesa and Ndegwa 2011; Griffiths et al.
recently determined that catches of ‘‘Tongkol’’—a 2013, Chacate and Mutombene 2014; Hornby et al.
species complex of small sized tunas—in Indonesia, in 2014). In countries such as Australia and Oman, they
particular West Sumatra, has primarily been attributed are highly regarded for their large size and fighting
to longtail tuna and has therefore probably resulted in ability, and because they can generally be targeted
significant overestimates of the catch (Geehan 2016). from small vessels in relatively sheltered inshore
Similar reporting issues appear to be evident in Japan waters. In contrast, in other countries in Southeast
where generic names such as ‘‘Shibi’’ (Itoh et al. 1996) Asia, longtail tuna are generally much smaller
and ‘‘Yokowa’’ are used for reporting species com- (\ 50 cm FL) and are used as bait (alive or dead) to
plexes of small-sized tunas, which include longtail target large billfishes (Istiophoridae spp.), tunas and
tuna (Itoh et al. 1996, Mohri et al. 2008). This likely sharks.
explains the dramatic increase in longtail tuna catches In Australia in particular, longtail tuna has grown in
in the WCPO region from 2004 and the subsequent popularity in recent years, with several annual catch
rapid decline since around 2011 (Fig. 5c) following and release tournaments being established to solely
improvements in port sampler training and reporting target the species, particularly by saltwater fly anglers.
protocols (Geehan 2016). Although longtail tuna is a This is probably helped by the presence of large
commonly caught species throughout parts of Indone- longtail tuna in Australia waters, with all 33 Interna-
sia, there is now great uncertainty over the extent of tional Game Fish Association (IGFA) world line class
over-reporting of longtail tuna in previous years, and records for the species—including the all-tackle world
whether further improvements in observer training can record of 35.9 kg—primarily coming from Australia’s
overcome these fundamental issues that severely east coast (IGFA 2017).
compromise the data available for future assessments, In recognition of the importance of longtail tuna to
for not only longtail tuna, but the several other recreational fisheries in Australia, the species was
principal commercial tuna species that have been declared a ‘recreational-only’ species by the Com-
incorporated into the ‘‘Tongkol’’ taxonomic monwealth government in December 2006, with an
aggregation. annual bycatch quota of 70 t allocated to multi-species
Similarly in Japanese waters, small longtail tuna Commonwealth commercial fisheries and a 10-fish
(\ 50 cm FL) occur sympatrically with small Pacific trip limit to allow for incidental catches in fisheries
123
that may have difficulty in eliminating longtail tuna fishers (Chacate and Mutombene 2014), and that the
captures due to the regions fished and the selectivity of surveys did not sample all fishing clubs or fishers who
the fishing gears (e.g., gillnet fisheries targeting sharks are not members of clubs, these catch estimates are
and Scomberomorus spp.) (Borthwick 2012). The most likely underestimated.
recreational catch of longtail tuna is managed by
individual State government fisheries agencies. Each Utilisation and marketing of longtail tuna
State imposes slightly different regulations, but gen-
erally longtail tuna are managed through minimum Despite being lesser known than some of the oceanic
size limits and daily catch limits. For example, in New tunas, such as skipjack and yellowfin tuna, longtail
South Wales, there is a per person possession limit of 2 tuna has established markets in canned and fresh forms
and 5 fish for longtail tuna above 90 cm TL and less both in domestic and export markets. Longtail tuna is
than 90 cm TL, respectively. most widely used in canned form, being processed in a
Very few data sources exist to accurately determine small number of plants primarily in Iran, India,
the magnitude of catches of longtail tuna by recre- Indonesia, and Thailand, which also process other
ational fisheries, mainly due to the species being principal tuna species such as skipjack, albacore and
targeted by specialised fishers who are infrequently yellowfin tuna. The canned product is then exported to
intercepted in recreational fishing surveys that are countries including the United States, Australia,
usually designed to sample generalist fishers (Griffiths Finland and Sweden (Asia–Pacific Fishwatch 2019).
et al. 2013). The specialised land-based gamefish In Thailand, longtail tuna destined to be canned for
fishery along eastern Australia was surveyed by export are delivered to the cannery directly from purse
Griffiths (2012) to estimate catch rates for species seine vessels unloading fresh fish at local ports, or they
such as longtail tuna. Catch rates were found to be very are imported frozen. Once at the cannery, the fish are
low, with fishers catching one longtail tuna for every graded for size and quality before being cooked. The
62.5 h of effort expended (0.016 fish hr-1). This was meat is removed manually and placed in cans prior to
due to a high average trip effort (9.44 ± SD 6.65 h), vacuum sealing and labelling (Asia–Pacific Fishwatch
which contributed AU$1735 (± 788) per fisher annu- 2019).
ally to local economies. In Bitung, Indonesia, some longtail tuna is packed
In a subsequent Australian nation-wide survey of in catering-sized cans destined for the food service
longtail tuna catches by boat-based and land-based market in the USA. However, most is packed in
recreational fishers, 8 t were reported to have been consumer-sized cans under a variety of labels and sold
caught in 2009 by the 1182 sport fishers surveyed. to small retailers. Some of the larger tuna companies
Given there are an estimated 3.6 million recreational that focus on processing skipjack and yellowfin tunas
fishers in Australia, of which approximately 5% target also produce small volumes of longtail tuna as an
tunas (Henry and Lyle 2003), it may be conceivable alternative for consumers who prefer ‘white meat’
that the recreational catch may exceed 1000 t per year. albacore tuna, particularly in some Middle Eastern
In Kenya and Mozambique, longtail tuna comprise countries, such as longtail tuna is a popular as a canned
a suite of near-shore tuna and tuna-like species that are white meat tuna. In the United Arab Emirates about
targeted by recreational fishers in small boats (3–9 m) 30% of the canned market is for white meat (albacore
(Wekesa and Ndegwa 2011; Chacate and Mutombene and longtail tunas) (Asia–Pacific Fishwatch 2019).
2014). The recreational fisheries in either country have In addition to canning and fresh fish, in some
not been the subject of detailed scientific surveys. In countries longtail tuna is smoked or used as sashimi. In
Kenya, Wekesa and Ndegwa (2011) used the volun- Indonesia, longtail tuna is hot smoked—known as
tery catch records from a single sport fishing club in ‘‘Fufu/ikan asar’’—in the same way in which skipjack
Kenya to estimate a recreational catch of 3.3 t for is prepared. Longtail tuna is used for sashimi in Japan,
longtail tuna. Similarly, Chacate and Mutombene and most likely other countries in Southeast Asia,
(2014) used voluntary submission of catch cards from although Japan’s reasonably small catches of longtail
the * 2700 licenced recreational fishers in Mozam- tuna mean that it is generally not widely or consis-
bique and estimated a total catch of 125 t. Given the tently available to consumers (Zennic 2016).
high frequency of non-reporting by Mozambique
123
In Pakistan, longtail tuna are generally not con- None of the stock assessments defined whether the
sumed locally but are salted and dried and exported to reported BRPs used were target or limit references
Sri Lanka (Moazzam 2012a). However, since 2003, all points. To be precautionary in this paper, BRPs were
longtail tuna and other tropical tunas (yellowfin and assumed to be limit reference points. Therefore, values
skipjack) are transported either by land or carrier of C 1 for F/FMSY or F/F0.1 indicate the stock is
vessels to Iran (Moazzam 2012a). It is believed that subject to overfishing (i.e. growth overfishing),
the majority of the tuna product sent from Pakistan to whereas values of B 1 for B/BMSY or SB/SBMSY
Iran is reflected in the annual landings of Iran, which indicate the stock is overfished (i.e. recruitment
may explain the relatively smaller annual catches by overfishing). For the more rudimentary assessments,
Pakistan (see Fig. 5b). that are often undertaken using ELEFAN or FiSAT
software, E has been used as the BRP. In these
assessments, values greater than 0.5 were assumed to
Stock assessment indicate the stock was subject to overfishing, since
E = 0.5 has been considered a proxy for MSY
There have been at least seventeen stock assessments (Gulland 1985). The BRP Fx% is the value of F at
of longtail tuna conducted in various regions through- which the spawning potential ratio (SPR) has been
out their geographic distribution. Despite available reduced to x% (usually in the range of 20–40%) of the
genetic and length-frequency data from fishery spawning stock in the absence of fishing (F = 0)
catches providing some evidence of the possible (Gabriel and Mace 1999).
existence of at least four main putative stocks— Hereafter, we summarize the results of these stock
Oceania, Southeast Asia, western Indian Ocean, and assessments by ocean basin, although the validity of
eastern Indian Ocean (Fig. 2)—the majority of assess- the outcomes need careful consideration as some of
ments did not define the spatial extent of the stock, but the data inputs are highly uncertain. For example,
instead generally used political boundaries. Of the assessments that only produce a value for E from
assessments that are hereafter summarised, ten used length-frequency analysis are likely to have higher
yield per-recruit (YPR) models—with length- or age- uncertainly in determining stock status than statistical
frequency data to derive biological and mortality models that produce estimates of F, FMSY, BMSY, and
parameters—one assessment applied a stock produc- SBMSY from time series of catch and effort data.
tion model to standardized CPUE data for the drift
gillnet fishery, while four assessments of the entire Eastern Indian Ocean
Indian Ocean used a range of data-poor assessment
models based on annual catches for all fisheries There have been at least six stock assessments of
combined (Table 3). longtail tuna in the eastern Indian Ocean all being
The assessments defined stock status using a range conducted off the west coast of India in the Arabian
of biological reference points (BRPs), making direct Sea and apparently none being undertaken off the east
comparisons between assessments difficult in many coast in the Bay of Bengal (Table 3). All of these
cases. Some of the more rigorous assessments used assessments used YPR models, and generally analysed
FMSY, BMSY, and SBMSY, which define the instanta- length-frequency data to derive biological and mor-
neous fishing mortality rate (F year-1), annual stock tality parameters, and so their results should be used
biomass (B) and spawning stock biomass (SB) at with caution. In an assessment of the stock in western
maximum sustainable yield (MSY), respectively. Indian waters using a YPR model for the period
Some assessments used the precautionary F0.1 BRP, 1984–1988, James et al. (1992) estimated an E of
which is the F where the slope of the YPR curve is 0.397 year-1 for commercial and artisanal fisheries
10% of the slope of the curve at its origin. The YPR and concluded the stock was approaching full
BRPs of FMAX and EMAX is the value of F and exploitation. A subsequent YPR analysis for the
exploitation rate (E) where yield is maximised. Here, period 1997–2002 (Pillai et al. 2011) indicated that
E is defined as E = F/(F ? M), where M is the E had steadily increased to 0.81 year-1 in 2002, which
instantaneous natural mortality rate. was in the vicinity of EMAX. They noted a progressive
decline in the mean size of fish in catches and
123
Table 3 Summary of stock assessments conducted on longtail (NO), Overfished (OF). Assessment models were: Stock
tuna categorised by putative stock (see Fig. 2), including Reduction Approaches (SRA), Stochastic Stock Reduction
countries involved, study source, assessment year(s), model Approaches (SSRA), Optimised Catch Only Method (OCOM),
and data type used, fishing mortality (F year-1), exploitation A Stock Production Model Incorporating Covariates (ASPIC),
rate (E), biological reference point(s) (BRP) used and their and yield per-recruit (YPR). Assessment parameters that were
values, and the status of the stock shown as: Overfishing not not reported, or unable to be calculated using other reported
occurring (ON), Overfishing occurring (OO), Not overfished parameter values, are shown as ‘‘NR’’
Stock Country/region References Assessment Model Data F year-1 E year-1 BRPs BRP Status
period type type values
Pan-Indian Ocean-wide Zhou and Sharma 2012 SRA Catch NR NR F/FMSY 1.08 OO
Ocean (2014)
B/BMSY 1.12 NO
Ocean-wide Martin and 2013 OCOM Catch 0.433 0.419 F/FMSY 1.11 OO
Sharma (2015)
B/BMSY 1.02 NO
Ocean-wide Martin and 2014 OCOM Catch 0.402 0.401 F/FMSY 1.03 OO
Robinson
(2016)
B/BMSY 0.99 OF
Ocean-wide Martin and Fu 2015 SSRA Catch 0.818 0.577 F/FMSY 3.27 OO
(2017)
SB/ 1.02 NO
SBMSY
Western Oman Prabhakar and 1987–1988 YPR LF 1.355 0.760 EMAX 0.73 OO
Indian Dudley (1989)
Ocean Oman, Pakistan, Al-Kiyumi et al. 2012 ASPIC CPUE 1.3 NR F/FMSY 1.38 OO
Yemen, Iran (2014)
B/BMSY 1.01 NO
Iran Yasemi et al. 2014 YPR LF 0.720 0.630 E 0.50 OO
(2017)
Iran Darvishi et al. 2015–2016 YPR LF 1.090 0.690 FMAX 0.85 OO
(2018)
F0.1 0.47 OO
F20% 0.17 OF
Eastern India James et al. 1984–1988 YPR LF 0.584 0.397 EMAX 0.79 ON
Indian (1992)
Ocean India Ghosh et al. 2003–2006 YPR LF 0.720 0.640 EMAX 0.64 OO
(2010)
India Pillai et al. (2011) 2001–2002 YPR LF 3.414 0.810 EMAX 1.00 OO
India Abdussamad et al. 2008–2010 YPR LF 2.940 0.790 EMAX 0.80 OO
(2012)
F10% 0.09 OF
Andaman Sea, Nishida et al. 2014 ASPIC CPUE 0.566 NR F/FMSY 1.11 OO
Malacca Strait (2016)
B/BMSY 0.89 OF
Malacca Strait Wagiyo and 2014 YPR LF 2.07 0.51 E 0.50 OO
Febrianti (2015)
Southeast Malaysia Raja Bidin and 1998–1992 YPR LF 1.439 0.630 EMAX 0.49 OO
Asia Rumpet (1990)
South China Sea Nishida et al. 2014 ASPIC CPUE 0.193 NR F/FMSY 0.18 ON
(2016)
B/BMSY 2.22 NO
Indonesia Restiangsih and 2014 YPR LF 1.01 0.62 E 0.50 OO
Hidayat (2018)
Oceania Australia Griffiths (2010) 2002–2004 YPR AF 0.209 0.369 FMAX 0.59 ON
F0.1 0.33 ON
F40% 0.39 NO
123
recommended caution be exercised in any further therefore the results need to be viewed with caution.
consideration of fishery expansion. A third assessment An assessment in Omani waters—primarily in the
undertaken in the same region for the period Gulf of Oman—was undertaken by Prabhakar and
2003–2006 (Ghosh et al. 2010) estimated E to be Dudley (1989) produced an E of 0.760 year-1, which
0.64 year-1, which exceeded EMAX by around 40%. was slightly less than the estimated EMAX of
Based on the assessment results, the authors recom- 0.727 year-1, leading to the conclusion that the stock
mended a reduction in fishing effort, or at least was probably fully exploited and recommended no
diversion of effort to deeper offshore waters, espe- further increase in fishing effort.
cially by large factory purse-seine vessels. The most In a similar region encompassing the Persian Gulf
recent assessment in Indian waters was conducted for and the Gulf of Oman, Al-Kiyumi et al. (2014) used
the years 2008–2010 (Abdussamad et al. 2012) and standardized CPUE data from the Omani drift gillnet
indicated that E remained high (0.80 year-1), exceed- fishery in A Stock Production Model Incorporating
ing the estimated EMAX of 0.63 year-1, which was Covariates (ASPIC). The model indicated that the
substantially lower than previous assessments due to stock in 2012 was subject to overfishing (F/FMSY-
the inclusion of updated biological parameters. Their = 1.38) and was in bordering on an overfished state
analysis indicated that the stock biomass had declined (B/BMSY = 1.01). In a subsequent assessment for 2014
to just 9.4% of the unexploited biomass and recom- using a YPR model, Yasemi et al. (2017) estimated
mended management action be taken to reduce the F (0.720 year-1) for the commercial drift gillnet
fishing mortality on small fish (\ 50 cm FL). fishery to be half that of Al-Kiyumi et al. (2014), but it
Further to the east, Wagiyo and Febrianti (2015) still produced a high E (0.626 year-1) that far
undertook a stock assessment of longtail tuna in the exceeded their estimated E BRP of 0.50 year-1. These
Malacca Strait using a YPR model based on monthly results led the authors to conclude that ‘‘the population
length-frequency samples taken from the port of of longtail tuna in the waters of northern Persian Gulf
Langsa during 2014. They estimated a high and Oman Sea is being heavily exploited and
F (2.07 year-1) and E of 0.51 year-1, which exceeded overfished at a higher level than the optimum and a
their E BRP of 0.50 year-1, implying the stock is better management policy is necessary in this area’’.
subject to overfishing. In the most recent assessment, conducted in Irani
Two stock assessments were initiated by SEAF- waters including the Persian Gulf and the Oman Sea,
DEC by Nishida et al. (2016) to characterise two Darvishi et al. (2018) determined that F had further
putative stocks for 2014; the region of the Indian increased to 1.090 year-1 since previous assessments
Ocean including the Andaman Sea and Malacca Strait in the region. They estimated the YPR BRPs of FMAX,
and the Pacific region incorporating the South China and F0.1 to be 0.85 and 0.47, respectively, and that the
Sea (see Southeast Asia). The Indian Ocean assess- population biomass was 17.2% of the unexploited
ment is one of the most reliable assessments available biomass at the current F value. From these results the
for longtail tuna that utilised CPUE data in ASPIC authors concluded the stock is subject to overfishing
models to conclude that the stock was subject to and is overfished (Table 3).
overfishing (F/FMSY = 1.11) and overfished (B/BMSY-
= 0.89). The authors recommended a reduction in Pan-Indian Ocean
catch and fishing effort by around 10%, despite the
catch being around one third of the catch in the In 2014, the IOTC—in conjunction with their Work-
adjacent stock in the South China Sea. ing Party on Neritic Tunas established in 2012—began
to solicit the development of data-poor methods for
Western Indian Ocean assessing the stock status of neritic tunas and seer-
fishes across the entire India Ocean. Although the
There have been at least four stock assessments of input data and stock status in the assessments
longtail tuna in the western Indian Ocean, all using described herein contain significant uncertainty, this
YPR models based on length-frequency data. Again, uncertainty is quantified statistically, and therefore
the reliability of the input data and subsequent stock these assessments are among the most reliable for the
status for the four assessments is highly uncertain and species to date.
123
Zhou and Sharma (2014) used traditional Stock subject to overfishing, primarily due to high catches by
Reduction Analysis (SRA) approaches and a Posterior gillnets that are most commonly used in the region.
Focussed Catch Reduction Approach (PFCRA) that The second stock assessment initiated by SEAF-
produced complementary results. Using the SRA DEC for longtail tuna was undertaken by Nishida et al.
results, they concluded that in 2012 the longtail tuna (2016) for the South China Sea, and represents one of
stock—assumed for modelling purposes to be a single the more reliable assessments for the species. They
panmictic stock throughout the Indian Ocean—was used CPUE data in ASPIC models to characterise the
subject to overfishing (F/FMSY = 1.08) and was near- stocks for 2014 and determined that the stock was
ing an overfished state (B/BMSY = 1.12). exposed to extremely low fishing mortality
Since this first assessment, the IOTC secretariat has (F = 0.193 year-1), resulting in the stock not being
conducted an updated stock assessment every year for subject to overfishing (F/FMSY = 0.18) and was not
longtail tuna in the Indian Ocean, but slightly refining overfished (B/BMSY = 2.22). In a review of the model,
the assessment models to cope with the generally poor Siriraksophon (2017) advocated for an increase in
catch and effort data available. Using an Optimised fishing effort, despite major uncertainties in the model
Catch-Only Method (OCOM), the stock continued to resulting from the use of data only from Thai fisheries
be subject to overfishing and in an overfished state in (mainly purse-seine).
2013 (F/FMSY = 1.11; B/BMSY = 1.02) (Martin and
Sharma 2015), and subject to overfishing and in an Oceania
overfished state in 2014 (F/FMSY = 1.03; B/BMSY-
= 0.99) (Martin and Robinson 2016). In the most Only one stock assessment has been conducted for
recent assessment for 2015 using OCOM, Stochastic longtail tuna in Oceania—in Australian waters (Grif-
Stock Reduction Approaches (SSRA) and Catch-MSY fiths 2010)—that used an age-structured YPR model
(C-MSY) models, Martin and Fu (2017) showed that based on a combination of fish aged directly from their
F/FMSY and B/BMSY have slightly reduced to around otoliths and length-frequency data. Therefore, the
1.0, due to a reduction in catches from 175,459 t in assessment is based on the most reliable biological
2012 to 136,849 in 2015. However, considering the data available and probably represents one of the most
results from the three models, and the large uncer- reliable assessments for the species. The assessment
tainty in parameter values, they concluded that the characterised the period 2004–2006 where the esti-
stock was still subject to overfishing and overfished. mated average annual F and E was 0.24 year-1 and
0.353 year-1, respectively. At this level of exploita-
Southeast Asia tion, the stock was not subject to overfishing relative
to both FMAX and F0.1. However, when considering
Three assessments of longtail tuna have been con- the spawning biomass, the current fishing mortality—
ducted in Southeast Asia, where some of the most where commercial exploitation is negligible by global
significant fisheries exist for the species, particularly standards—was in the vicinity of F40%, indicating that
for small fish (\ 50 cm FL), presumably juveniles. An the stock was probably fully fished and that any
assessment conducted for Malaysian waters for the increase in fishing effort would likely result in the
period 1998–1992 using a YPR analysis (Raja Bidin stock being overfished. The assessment explored the
and Rumpet 1990) was based on landings from two potential benefits of increasing minimum retention
adjacent landing sites in the southwest South China lengths and handling practices to reduce post-release
Sea and estimated an E of 0.630 year-1. This value mortality to reduce fishing mortality by the recre-
exceeded the estimated EMAX value of 0.490 year-1, ational fishery, but these measures proved to provide
indicating that the stock was most likely subject to marginal benefits to the stock since the majority of the
overfishing. catch was composed of large ([ 60 cm FL) fish.
Restiangsih and Hidayat (2018) used using a YPR
model based on monthly length-frequency samples
taken in the Java Sea during 2014 and estimated
longtail tuna to have an F of 1.01 year-1 and an E of
0.62 year-1. They also concluded that the stock was
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Discussion Indian Ocean longtail tuna stock. Although Somali

piracy appears to have abated in recent years, which
Tunas are among the most productive and economi- has contributed to the re-establishment of more tradi-
cally important fishes in the world’s oceans, annually tional tuna and billfish fishing grounds in ABNJ in the
supplying over 4 million tonnes of product to domestic Arabian Sea (Moazzam and Ayub 2015; Nishida
and export markets since 1998 (FAO 2019). For the 2015), the extent to which the additional post-2009
species that comprise the majority of the world’s effort has impacted the spawning stock of longtail tuna
capture production, namely skipjack and yellowfin and the population’s potential to recover whilst being
tunas, the sustainability of their catches can be exposed to current levels of fishing effort remains
attributed to their biology—being short-lived unknown.
(\ 10 years), fast-growing, maturing early in life Similar to the Indian Ocean, declines in the
(\ 2 years) and highly fecund—and their diffuse spawning biomass of target species in the WCPO
geographic distribution across ABNJ. In contrast, this have been well documented for yellowfin tuna
review has shown longtail tuna to be relatively long- (Tremblay-Boyer et al. 2017), bigeye tuna (McKech-
lived (up to 18 years), slow-growing, and have a nie et al. 2017), swordfish (Takeuchi et al. 2017) and
geographical distribution that is confined to relatively striped marlin (Davies et al. 2012) and pose a risk that
shallow continental shelf waters that make them fisheries may begin to increasingly target neritic tunas
vulnerable to capture by a range of coastal fisheries. in order to remain profitable and provide food security
Consequently, longtail tuna form an increasingly for coastal States, particularly those bordering the
important component of a range of artisanal, recre- South China Sea. There is certainly growing evidence
ational, and small and industrial scale commer- of the increased retention of formerly discarded
cial fisheries in many countries throughout the species (e.g., wahoo, rainbow runner, dolphinfish)
species’ geographic range. However, non-reporting, with the increasing shift by the WCPO industrial
and misidentification or taxonomic aggregation of purse-seine fishery to set on floating objects (i.e.,
small tunas—e.g., ‘‘Tongkol’’ in Indonesia (Ingles FADs), which attract high diversity and biomass of
et al. 2008) and ‘‘Shibi’’ in some regions of Japan (Itoh tunas, sharks and tuna-like species (Leroy et al. 2013,
et al. 1996)—have contributed to limitations in Griffiths et al. 2019a). It is possible at some point that
accurately aligning catch data with consumer demand coastal States throughout Southeast Asia may begin to
(Willette et al. 2017) and our current poor understand- withdraw their industrial vessels from the current
ing of the true extent of fishing impacts on their stocks. distant water purse-seine and longline fishing grounds
In the face of the declining spawning biomass of located in the western Pacific Warm Pool Province
several economically important species of pelagic (Williams and Reid 2018) to target longtail tuna and
fisheries in the Indian Ocean since at least the 1990s, other neritic tunas and tuna-like species within more
including bigeye tuna (Langley 2016a), yellowfin tuna local waters, such as the South China Sea.
(Langley 2015), albacore (Langley and Hoyle 2016), Despite the widespread adoption of the principals
swordfish (Nishida and Yokoi 2017), blue marlin of the precautionary approach to fisheries manage-
(Wang and Huang 2016) and striped marlin (Nishida ment (FAO 1995) by numerous national fisheries
2015; Wang 2015)—and in some cases, overexploita- agencies worldwide and the more recent pursuit of
tion—longtail tuna and other neritic tuna and tuna-like more holistic ecosystem-based approaches to fisheries
species including kawakawa, frigate and bullet tunas management (FAO 2003) by RFBs such as the Asia–
and seerfishes have become an increasingly attractive Pacific Fishery Commission (FAO 2017a), it is
alternative for the fisheries of coastal States. worrisome that such obvious increases in longtail
Moreover, the increase in Somali piracy in the tuna catches have failed to capture the attention of
western Indian Ocean that resulted in a dramatic shift fishery managers until very recent years, particularly
in the effort by large-scale commercial tuna fleets, in the Indian Ocean. Although many aspects of the
such as drift gillnet fisheries of Iran and Pakistan, to biology, movements and stock structure of longtail
fish coastal waters to target neritic species since at tuna are little understood, this review has demon-
least 2009 (Al-Kiyumi et al. 2014, Moazzam 2014) is strated there is certainly sufficient information avail-
likely to have greatly exacerbated pressure on the able to undertake reasonably rigorous integrated
123
assessments—beyond that of simplistic YPR mod- objectives, licencing controls, catch limits, consulta-
els—for at least each of the four putative stocks tion arrangements, monitoring/control/enforcement
(Fig. 2) using new data-poor assessment methods that measures and catch monitoring in accordance with
have been applied recently to the Indian Ocean stock both established best practices and the requirements of
(see Martin and Fu 2017). any agreed CMM (Cochrane 2002; FAO 2003;
Even more alarming is that several independent Cochrane and Garcia 2009).
lines of evidence, albeit from rudimentary YPR Fortunately, the recent stock assessment outcomes
models based on biological and fishery data of have generated the impetus for the IOTC Scientific
varying degrees of reliability, have indicated that Committee in 2017 to recommend the introduction of
longtail tuna has likely been subject to overfishing and CMMs for longtail tuna; including the development of
overfished for a number of decades, especially where limit reference points and the reduction in catches by
intense and poorly regulated fisheries exist. For 10% of 2015 levels (to 136,849 t) to not exceed a
example, even as far back as the late 1980s in Omani provisional MSY of 140,000 t (IOTC 2017). Although
waters where the most reliable biological and fishery this is a positive move forward to develop interim
data was available for longtail tuna at the time, precautionary management measures, these recom-
Prabhakar and Dudley (1989) warned that the stock mendations have not yet been accepted and imple-
was probably fully exploited and recommended no mented into formal CMMs by the IOTC.
further increase in fishing effort. Unfortunately, some Another positive initiative by the IOTC that should
30 years later, such scientific advice has not yet benefit longtail tuna was the development of a CMM
initiated the development or implementation of mean- (Resolution 17/07 ‘‘On the prohibition to use large-
ingful CMMs, neither within the coastal States or at scale driftnets in the IOTC area’’) that entered into
the level of RFMO. force in 2017, prohibiting the use of large-scale drift
gillnets ([ 2.5 km in length) in the IOTC area of
Developing a management framework competence by 2022. This measure is likely to reduce
the fishing mortality on longtail tuna, as well as other
The fisheries management frameworks in place across neritic tunas and a range of bycatch species with
the distribution of longtail tuna differ markedly, and vulnerable life histories, including turtles, cetaceans,
have probably contributed to the slow progress in the and elasmobranchs (Northridge 1991; Moazzam and
species being formally recognised as requiring specific Nawaz 2014). Additionally, although there is no
CMMs. At a macro level, different management specific CMM in place for longtail tuna in Pakistan,
arrangements are in place in the Indian Ocean versus there is a two-month closure (June–July) that provides
the Western Pacific Ocean as the Indian Ocean has a some respite for the longtail tuna stock. During the
single organisation—the IOTC—that has ulti- winter (January–March) the Pakistan gillnet fleet
mate management responsibility for longtail tuna. moves to offshore waters of the EEZ and in ABNJ,
Based on evidence of overexploitation of longtail and thus longtail tuna are not harvested during this
tuna from recent exploratory stock assessments under- period.
taken annually by the IOTC Secretariat between In contrast to the IOTC, longtail tuna is not an
2013–2017, there is an immediate opportunity for explicit responsibility of the Western and Central
the IOTC to promulgate CMMs aimed at reducing Pacific Fisheries Commission (WCPFC), since its
pressure on the species. However, CMMs developed Convention includes only ‘‘highly migratory species’’
by the IOTC that may afford some protection to listed under Annex I of the 1982 United Nations
longtail tuna, either directly or indirectly, only become Convention on the Law of the Sea (UNCLOS) (UN
mandates for those members that agree to a CMM at 1982), which does not include longtail tuna. A further
the time of its adoption by the Commission. Therefore, complication is that the WCPFC Convention area does
this would require the countries that contribute most to not include the waters throughout Southeast Asia and
catches to agree on measures to reduce fishing effort, the South China Sea (see Annex 4 of SPFFA 2000),
especially amongst the gillnet fleets. Each nation where the majority of longtail tuna catches are made
should move to prepare a comprehensive management outside of the Indian Ocean. However, FAO catch
plan for these fleets, which set out management statistics attribute longtail tuna catches to Major
123
Fishing Areas 61 and 71 that encapsulate a large western Pacific is that there is no fishery body that has
proportion of the WCPFC Convention area. Fortu- responsibility for coordinating management across the
nately, RFBs within Southeast Asia and the South countries that catch longtail tuna. Moreover, there
China Sea are beginning to make progress to improve seems to be a current dispute amongst ASEAN
knowledge on the biology and fisheries for longtail members as to where responsibility for initiatives on
tuna, mostly as part of a wider complex of neritic tunas neritic tunas should lie (SEAFDEC 2018). To date
and seerfishes. For example, in 2014 SEAFDEC there has been little political appetite across the
established the Scientific Working Group on Neritic various smaller RFBs to commit the required
Tuna Stock Assessment in the Southeast Asian Waters resources to initiate a comprehensive research pro-
and subsequently developed the RPOA-Neritic Tunas gram and stock assessments for longtail tuna in the
in the ASEAN (Association of Southeast Asian western Pacific Ocean, despite catches throughout
Nations) Region in 2015 (SEAFDEC 2017). Southeast Asia contributing 56% to the global catch of
The RPOA-Neritic Tunas has focused on data longtail tuna in 2017 (FAO 2019). Much of the current
collection, with work plans being established for funding has been sourced from foreign aid (e.g. via the
research, catch monitoring, and capacity building for Swedish International Development Cooperation
Member States, which should greatly improve the Agency). The management of longtail tuna—despite
quality of data available to undertake future assess- attempts to push for multijurisdictional management
ment of the important Southeast Asian longtail tuna approaches—has to compete with a plethora of issues
stock(s). confronting Southeast Asian nations such slave labour
The earliest known management initiative in the (Marschke and Vandergeest 2016; Fischman 2017)
WCPO was the ban on commercial targeting of and IUU fishing (Li and Amer 2015).
longtail tuna as part of its declaration as a ‘‘recre- The aforementioned complexities mean there is no
ational only’’ species in Australian waters in 2006 clear mechanism for establishing CMMs that parallel
(Borthwick 2012). This was essentially implemented those available in the IOTC. Given the large-scale
to prevent any future large-scale commercial targeting movements of longtail tuna, there may be merit in
of longtail tuna, where there has been negligible seeking a change to the relevant articles of the WCPFC
commercial exploitation historically. Convention or inclusion of the species in Annex I of
The growing interest in management has resulted in UNCLOS that underpins the WCPFC Convention.
the development of fisheries management plans that However, under Article 5 of the WCPFC Convention
cover longtail tuna in both Thailand and Indonesia. In it states the Commission will ‘‘assess the impacts of
2015, Thailand implemented a new fisheries manage- fishing, other human activities and environmental
ment plan that includes all pelagic species (DOF factors on target stocks, non-target species, and
2015), which is a valuable step forward. The Thai plan species belonging to the same ecosystem or dependent
aims to regulate overall fishing effort rather than seek upon or associated with the target stocks’’. Therefore,
to implement a large number of individual species it stands to reason that the WCPFC should assume
management measures. Whether this will benefit responsibility for longtail tuna as they belong to the
longtail tuna has not yet been demonstrated. Indone- same ecosystem, and are associated with, several
sia’s tuna plan (2014–2019) (MAFRI 2015) is detailed exploited species listed under Annex 1, in particular E.
but in terms of the neritic tunas, is mainly focused on affinis. A. thazard and A. rochei. It should also be
data gathering rather than putting into place effective noted that the absence of particular species in Annex 1
management controls. Part of the reason for this may has not precluded the WCPFC from developing
be the optimistic stock assessment but history would specific CMMs for some of these species, such as
suggest that the best time to put in place catch controls sea turtles (CMM 2008-03).
is when stocks are healthy. Alternatively, a specific management body for
However, even if all countries had management neritic tunas could be developed, possibly via the
plans in place there would continue to be a need for ASEAN. Such an approach has been used in the
coordination across countries in order to keep the western Indian Ocean, where the Regional Commis-
cumulative fishing mortality at a biologically sustain- sion for Fisheries (RECOFI) developed a Working
able level. Therefore, the largest challenge in the Group on Fisheries Management in 2001, which is
123
currently developing improved data reporting and data their fisheries to improve the reliability of stock status
harmonization among its Member States, and plans to determined by stock assessment models, a number of
coordinate stock assessments on the region’s most fundamental studies are recommended and discussed
economically important neritic species, such as King- hereafter to be undertaken in at least the four major
fish (S. commerson) (FAO 2017b). Nonetheless, the putative stocks of longtail tuna (Fig. 2). Until such
aforementioned approaches are unlikely to result in times as more reliable data become available to better
suitable catch controls being developed in a timely assess longtail tuna stocks, precautionary management
manner to curb the overexploitation of longtail tuna in measures need to be exercised, such as setting interim
recent years. input and/or output control measures such as intro-
ducing or reducing quotas or fleet capacity (e.g. IOTC
2017), or changing gear configurations, such as
Research and management priorities increasing mesh size of nets to reduce the mortality
on juvenile tuna.
Longtail tuna has attracted the attention of scientists at
least since the work of Serventy (1942b), but there has Improved quality and reporting of catch and effort
been surprisingly little research conducted in the for conservation and management
intervening 76 years, and much of the work appears to
have been undertaken in an opportunistic or ad hoc As has been discussed throughout this review, our
manner, or using methodologies or sampling designs poor level of understanding of longtail tuna population
that are not of sufficient reliability to characterise dynamics, fisheries exploitation and stock status
population dynamics parameters to be informative for primarily stems from the historically lower catches
stock assessment. Much of the reasoning for this is and economic importance of longtail tuna, compared
twofold. First, many of the countries that contribute to to principal target species, that has failed to garner
the global capture production of longtail tuna are sufficient interest from national fisheries agencies,
developing nations, and therefore do not have the RFBs and RFMOs to invest resources into monitoring
resources to undertake the required biological research and research. Although the tuna catch composition has
and fisheries monitoring or employ adequately trained certainly changed in recent years, particularly in the
staff to undertake these activities. Second, until the Indian Ocean, a major obstacle for data collection is
late 1980s, longtail tuna had not been established as a that a large proportion of the longtail catch is derived
bona fide species of economic or social importance, as from small-scale commercial and artisanal fisheries—
its magnitude of catches were generally dwarfed by and primarily recreational fisheries in Australia.
catches of principal species such as skipjack and Quantifying catch and effort from these fisheries can
yellowfin tunas, especially in the WCPO. While be an enormous challenge as there can be large
managers have directed their attention, and often numbers of operational vessels, but many do not need
limited resources, to rectify the situations for principal to be registered or require a fishing licence or permit—
target species, longtail tuna appears to have been or at least appear on any type of official registration
overlooked, until recent scientific evidence has shown list—or require their catch and/or effort to be reported
that the species is slow-growing and long-lived (living (e.g. Somalia; Persson et al. 2015). Consequently, the
for at least 18 years)—similar to the life histories of magnitude of participants in these fisheries and their
the larger Thunnus species bigeye and southern bluefin impacts on stocks are poorly understood. Even fishery-
tunas that have contributed to their overexploitation independent surveys of these fleets are logistically
(Griffiths et al. 2010a)—and is now in an overex- difficult and often cost-prohibitive as vessels or fishers
ploited state throughout the majority of its worldwide need to be intercepted at an enormous number of
distribution. diverse access points (e.g. public and private docks,
Fisheries scientists and resource managers must beaches, rivers, motherships) that are dispersed across
recognise that urgent action is required to ensure the thousands of kilometres of coastline and open ocean
long-term sustainability of longtail tuna stocks. In (Griffiths et al. 2010b; Geehan 2016).
order to make meaningful progress in better under- It is of paramount importance that coastal States,
standing the population dynamics of longtail tuna and with the support of RFBs and RFMOs, develop, or
123
improve, data collection systems that are reliable and international trade and the markets they feed into.
validated periodically. In Indonesia for example, data Understanding the trade can help generate a better
collection systems have been in place since the 1970s understanding of volumes, especially in the context of
and comprise obligatory reporting from fishing com- growing trade requirements for more accurate trace-
panies, monitoring at landing sites, and periodic ability. Importing countries, such as the United States,
fishery-independent surveys of fishing villages. can impose data collection requirements on exporting
Recent advances in, and broader availability of, nations if traceability is unclear or if the fishery in
molecular genetic methods present new opportunities question is in violation of other requirements such as
to rapidly and accurately examine fish stocks and the Marine Mammal Protection Act of 1972. In
catches (Willette et al. 2014). DNA barcoding is an January 2017, the United States introduced the
increasingly common method used in both developed Seafood Import Monitoring Program (SIMP) that
and developing countries to detect mislabelling of requires data on fish products to be verified at each
market-sold tuna (Maralit et al. 2013, Abdullah and step in the supply chain from harvest to final impor-
Rehbein 2014; Khaksar et al. 2015), and has recently tation into the US (Willette and Cheng 2018).
been shown effective in assessing the biodiversity and The gillnet fisheries for tunas in Iran and Pakistan
biomass of marine fish, including tuna, in seawater would be among the most affected under the SIMP,
samples (Takahara et al. 2012, Kelly et al. 2014, Port which would also apply to intermediary nations that
et al. 2016, Willette 2017). Of particular value is a pair may be importing and processing longtail tuna prod-
of Thunnus-specific primers (MiFish-tuna-F and ucts prior to arrival into the US. The SIMP gives
MiFish-tuna-R), which can unambiguously distin- exporting nations a 5-year period to demonstrate to the
guish longtail tuna from other Thunnus species (Miya US government that they have a fishery-level marine
et al. 2015). This may be one plausible validation mammal bycatch that is no different from a compa-
process that may be periodically applied to ensure rable US fishery. Failure to provide this assurance will
correct species reporting, or at the least, better allow result in the products being denied entry to the US.
the extent of misidentification to be understood and
corrected for in final catch statistics. Stock structure and movement
Although improved data collection processes will
greatly improve the quality of catch and effort data for The definition of stock boundaries is critically impor-
fisheries that catch longtail tuna, it is clear, even from tant in the assessment of species impacted by fishing
the limited data currently available, that implementa- (Cadrin and Secor 2009). Stock boundaries have been
tion of management measures aimed at controlling defined using a variety of techniques including
catches is needed. Capacity reductions, for example, morphometrics (Serventy 1956; Abdulhaleem 1989),
may not only benefit longtail tuna but other species conventional tagging and biological markers (e.g.,
known to be under pressure (e.g. yellowfin tuna). parasites, otolith microchemistry) (Begg and Wald-
Moreover, as stocks of some principal species decline man 1999), and mtDNA D-loop genetic analysis
fishing effort is likely to shift to other species, such as (Willette et al. 2016). However, each of these methods
longtail tuna, which may result in serial depletion of infer potential stock boundaries across different time
new target species, and ultimately the degradation of scales from days to months (tagging), years (otoliths),
the structure and function of the ecosystem. In the to generations (morphometrics and genetics).
South China Sea, the focus needs to be on developing a Genetics has become an increasingly powerful and
more robust mechanism for agreeing and implement- inexpensive method used to define stock structure for
ing coordinated management across jurisdictions. This tunas, particularly single nucleotide polymorphism
could take place in the absence of any formal RFMO, (SNP) markers (Grewe et al. 2015). Genetic analyses
if the political will is sufficient. are ideal in situations where a strong genetic gradient
can be used to differentiate stocks. However, minimal
Markets and market demand generational gene flow between true sub-populations
can result in the appearance of a single panmictic
There is currently insufficient information on the stock. Therefore, stock structure studies for longtail
usages of longtail tuna and, in particular, their tuna may benefit from the use of complementary
123
techniques used in concert (see Begg and Waldman Ferreri et al. 2009) and allowing the identification of
1999), such as genetics to establish an evolutionary spawning locations using the presence of post-ovula-
scale baseline coupled with, for example, conven- tory follicles in the ovaries.
tional tagging that will allow the shorter term direction Therefore, it is recommended that growth and
and magnitude of movement of fish to be determined reproductive studies be undertaken within each stock
in order to identify independent reproductive stocks. using, at a minimum, routine methodologies applied to
Such work would involve the collection and samples representing the entire size spectrum of fish
tagging of specimens at sufficient spatial scales across found within each stock. For ageing studies, this
the entire distribution of the stock, for example the should involve the analysis and quantification of daily
entire Indian Ocean. With the rapid advancement of and/or annual growth increments in sectioned sagittal
genetic techniques, a genetic study may serve two otoliths that have been validated by chemically-
important purposes. First, it can define stock bound- marked (e.g., oxytetracycline, bomb radiocarbon) fish,
aries, and the second may be to use genetic material for or at least by otolith edge-type analysis, of fish
new cost-effective close-kin mark-recapture tech- representing a wide range of age classes (Campana
niques (Bravington et al. 2016b) to estimate the 2001). Reproductive studies ideally should be under-
absolute abundance of longtail tuna for stock assess- taken simultaneously with growth studies using the
ment, which has been achieved for southern bluefin same sampled fish, thus optimising cost-effectiveness
tuna (Bravington et al. 2016a) and the white shark of the sampling. If pilot studies or previous work
(Hillary et al. 2018). Such an estimate of abundance cannot determine the spawning period, monthly
would greatly reduce the current uncertainty in stock histological samples should be processed, or a GSI
size and the estimated level at which the stock can be be closely monitored, to detect changes in reproduc-
sustainably harvested using biological reference tive condition. Intensive sampling is then recom-
points such as MSY (see Martin and Fu 2017). mended during the spawning period across the widest
possible size range of fish in order to optimise the
Growth and reproductive dynamics precision of an L50 estimate and representativeness of
the maturity ogive.
As shown in Tables 2 and 3, there are wide-ranging If stocks were to be prioritised for this biological
estimates of growth and reproductive parameters for research, we suggest to first examine the Indian Ocean
longtail tuna, even within the same region, which is stocks, since this is the region that contributes most to
likely a result of inadequate sampling and/or unreli- the global catch of longtail tuna and where the stock is
able analysis techniques (e.g. modal length analysis). overfished and overfishing is known to be occurring.
These effects can have significant impacts on esti- Given that the putative stocks of longtail tuna
mates of growth and mortality parameters, and thus, straddle several political boundaries of varying spatial
stock status. Although population parameters and scales from coastal States, to RFBs, to RFMOs
stock status can be estimated reliably using length- (Fig. 2), it is imperative that entities responsible for
frequency data, as is routinely done using MULTI- the management of a particular stock(s) of longtail
FAN-CL for principal tuna species in the WCPO tuna closely collaborate to ensure consistency in
(Fournier et al. 1998), these models require a great collection and analysis methods, which may be
deal of data from all fleets and time and spatial strata, achieved through forums such as the IOTC’s WPNT,
which is unlikely to be feasible for several coastal RECOFI’s Working Group on Fisheries Management
States throughout Southeast Asia and the Indian Ocean (WGFM), and SEAFDEC’s RPOA-Neritic Tunas.
where resources for long-term monitoring may be Improved data quality will allow more reliable stock
limited. assessments to be undertaken and allow fishery
With respect to reproductive dynamics, macro- managers to be better equipped to develop appropriate
scopic staging and gonadosomatic indices have been CMMs to safeguard the long-term sustainability of
used in the majority of published studies over reason- longtail tuna stocks.
ably inexpensive histological methods, which have far
higher precision for estimating L50 and developing Acknowledgements The authors would like the thank Meryl
Williams for allowing some of the information researched by
maturity at length (or age) ogives (Vitale et al. 2006,
123
some of the authors for the longtail tuna AsiaPacific-Fishwatch Terengganu waters, east coast of Peninsular Malaysia.
profile to be used in the production of this paper. D. Willette NAGA, ICLARM Q 27:41–47
thanks M. Joaquin for logistical support. Basir S, Jamon S (2012) Catch performance of the purse seines
for the neritic tuna fishing in the Strait of Malacca. Second
meeting of the IOTC Working Party on Neritic Tunas,
Penang, Malaysia, 19–21 November 2012. Document
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Rev Fish Biol Fisheries

Longtail tuna, Thunnus tonggol (Bleeker, 1851): a global

Received: 9 May 2019 / Accepted: 29 November 2019

Introduction Due to the often-poor reporting of neritic tunas in

Within their broad geographic range, longtail tuna Stock structure

estimates that are probably not representative of the Timing of spawning

Oman Brothers (1990) Otoliths 30–82 22 78.5 0.679 - 0.490 50 71 77 7.0

(Scomberomorous spp.) (Yesaki 1994). In the Indian WCPO fisheries

Discussion Indian Ocean longtail tuna stock. Although Somali

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