Current update: Understanding the King of River, ASEAN Mahseers, Tor spp.

Faizul Jaafar1 and Ishwar Parhar1

1
Brain Research Institute Monash Sunway (BRIMS), Jeffrey Cheah School of Medicine and
Health Science, Monash University Malaysia, Bandar Sunway, 47500, Malaysia

1
Brain Research Institute Monash Sunway (BRIMS), Jeffrey Cheah School of Medicine and
Health Science, Monash University Malaysia, Bandar Sunway, 47500, Malaysia.

*Corresponding author
Prof. Dr. Ishwar Parhar
Address: Brain Research Institute Monash Sunway (BRIMS), Jeffrey Cheah School of Medicine
and Health Science, Monash University Malaysia, Bandar Sunway, 47500, Malaysia
Email address: ishwar@monash.edu
Tel: +603-5514 6304
Fax: +603-5514 6341
Abstract

Mahseer has been nominated as King of River by the fisher is comprised of three genus which
are Tor, Neolissochilus and Naziritor under classification of Cyprinidae family and Cyprininae
subfamily. Amongst these three genus Tor genus has been classified as trues mahseer due to the
presence of median lobe which mainly present in this genus. Tor species is widely distributed
across Asian region. In Southeast Asia (ASEAN) there are nine valid species and one considered
as invalid. Furthermore, out of these nine valid species eight species have been listed in IUCN
Red List. Each ASEAN country has their owned or shared Tor species. Tor species that has been
reported that inhibit ASEAN countries are Tor tambroides, Tor tambra, Tor douronensis
(invalid), Tor ater, Tor laterivittatus, Tor putitora, Tor sinensis, Tor dongnaiensis, Tor
mekongenesis and Tor tor. This review would provide current update and would discuss about
the ASEAN’s Tor species in term of distribution, identification and specification and future fate
of this genera or species. Currently, there are limited number of publications on ASEAN’ Tor
species studies particularly on ichthyology to determine the current population size and
distribution. Furthermore, current ichthyology study shows the disappearance of Tor species in
particular river system. Different and unstandardized approach such as COX1, 16S rRNA, Cyt b,
ATPase 6 and microsatellites in identification and specification further widen the gap and
ameliorates the instability of Tor species taxonomy. Therefore, synchronized and strategized
research among Tor species researcher would been an alternative way to improve and fill the gap
of knowledge. In addition, DNA that carries an organism data base should be fully sequenced for
deep understanding in term of sex maturation, growth, feeding and breeding process. Current
understanding would help to update our current knowledge about ASEAN’s Tor species and
open up the new direction of ASEAN’s Tor research.

(300 words)
Introduction

The king of river or mahseers are amongst the largest scale carp and valuable group of
freshwater fish in Asia region which referred to three genera: Tor, Neolissochilus and Naziritor
under the classification of Cyprinidae family and Cyprininae subfamily (Nguyen et al. 2008).
The word mahseers represent two Bengal root words which are; Maha means Greatness and Seer
means Mouth or Head (Ng 2004). Mahseers are considered as important species in the
ecosystem, attractive in sport fish and commercially valuable as high esteemed food and high
demand in aquaculture (Pinder et al. 2019). These king of river are endemic to Asia which
originated from southwest China and widely distributed to the west of Asia across trans-
Himalayan region encompassing the river in Pakistan, India, Sri Lanka, Bangladesh and
Myanmar. These mahseers further extended down to the most of southeast of Asia nations
(ASEAN) across Thailand, Laos, Vietnam, Malaysia and Indonesia (Nguyen et al. 2008). The
habitat of mahseers are mainly found in the fast flowing of large river and lake and they are
migrating to the upstream river with clear water streams, pristine and rocky bottom for breeding
(Azuadi et al. 2011). Generally, mahseers are characterized by large sized freshwater fish with
compressed and elongated body (Stone 2007). The mahseers body is covered with strong and
large cycloid scale. The head of this fish is strong and large. Furthermore, these mahseers have
large mouth and tail with the addition of strong muscle and fin (Ng 2004).

ASEAN comprised dozens of rivers and one of it is the twelfth longest river in the world
that run thorough from Tibetan Plateau across Myanmar, Laos, Thailand, Cambodia and end in
Vietnam before its meets the South China Sea. ASEAN’s rivers has been described as the most
productive river for wild-captured freshwater fish and contributed to the 20.6% from total
worldwide capture fish produced by ASEAN in year 2013 (Chan et al. 2017). The percentage of
wild-captured fish in ASEAN is projected to increase1.8% annually from 2014 until 2030 (Chan
et al. 2017). Besides that, ASEAN also is an important producer for aquaculture that produce
10.9 million ton of aquaculture product in 2014 that account 14% of world total aquaculture
product (Chan et al. 2017). The main group of freshwater fish cultured in captivity are catfishes,
tilapias and cyprinids which include mahseers species (Pinder et al. 2019). With increased
demand of freshwater fish yearly encourage overfishing of wild mahseers and push the
aquaculture farmer to have more mature mahseers for broodstock and sustained the increased
demand of these species. Despite high productivity of rivers for wild-capture fish, the number of
polluted ASEAN’s rivers also increased tremendously. Rapid deforestation for development and
several anthropogenic factors contribute to the increased number of polluted rivers. Furthermore,
increased population in ASEAN demands for large volume water supply which required more
dams to be built would restrain the migration and isolation of mahseers. All of these factors
would contribute to the rapid decline of mahseers in their natural population (Pinder and
Raghavan 2012).

Amongst three genera of mahseers, the genus Tor has been considered as the ‘true
mahseers’ based on the morphological structure of the median lobe which mainly present in this
genus but not in others two genus (Nguyen et al. 2008; Pinder et al. 2019). There are 11 Tor
species that have been reported previously in ASEAN (Table 1). However, the morphological
features such as shape, size, fin ray and the length of the medium used previously to distinguish
the Tor species has caused confusion and debate among the biology researchers (Walton et al.
2017). These morphological features of Tor species showing high variability intra- and inter-
species and influenced by the environmental factors and locality (Ng 2004; Walton et al. 2017).
Recent review by Pinder et al. 2019 has concluded that there are only 9 species of Tor which
have been considered as valid species. Among these valid species, 8 species have been listed
under IUCN criteria (IUCN 2019). There is 1 species classified under each “vulnerable” and
“endangered” categories, 2 species classified under “near threatened” and 4 species classified
under “data deficient” (Table 2). Furthermore 4 species of ASEAN’s Tor show decreasing trend
of their population number (IUCN 2019). Therefore, an extensive research is required to improve
the ASEAN’s Tor species data particularly on specific identification of each species, genetic
diversity and variety and also sex determination. In this review paper, we will focus and discuss
on the current update of distribution, locality, identification and taxonomy of ASEAN’s Tor
comprehensively. These would provide another points of strategies in conserving wild Tor
species, maintaining the productivity of Tor aquaculture and preservation of their habitat and
surrounding biodiversity.
No Species Validity Reported Geography References
. Status by distribution
2019
1. Tor tambroides Valid Malaysia, Thailand, Vietnam, (Pinder et al.
Indonesia, Myanmar 2019)
2. Tor tambra Valid Malaysia, Thailand, Indonesia (Pinder et al.
2019)
3. Tor douronensis Invalid Malaysia, Thailand, Indonesia (Nguyen et al.
2008)
4. Tor ater Valid Laos (Roberts 1999)
5. Tor laterivittatus Valid Laos (Kottelat 2011)
6. Tor putitora Valid Myanmar (Shahi et al.
2014)
7. Tor sinensis Valid Vietnam, Laos, Cambodia (Hoang et al.
2015)
8. Tor dongnaiensis Valid Vietnam (Hoang et al.
2015)
9. Tor mekongenesis Valid Vietnam, Thailand (Hoang et al.
2015)
10. Tor tor Valid Myanmar (Shahi et al.
2014)
Table 1: List of valid Tor spp in ASEAN countries and its geographical distribution.
No. Species Listed in IUCN Species Status Population status
1 Tor tambroides Listed Data Deficient Unknown
2 Tor tambra Listed Data Deficient Decreasing
3 Tor ater Listed Near Threatened Unknown
4 Tor laterivittatus Listed Data Deficient Decreasing
5 Tor putitora Listed Endangered Decreasing
6 Tor sinensis Listed Vulnerable Unknown
7 Tor dongnaiensis Listed Near Threatened Unknown
8 Tor mekongenesis Not listed N/A N/A
9 Tor tor Listed Data Deficient Unknown

Table 2: Current status of ASEAN’s Tor species by 2019; (IUCN 2019)

Geographical distribution Tor species habitat in ASEAN region

Malaysia

Malaysia consists of peninsular of Malaysia and Borneo Island (Sabah and Sarawak) and home
to tropical rainforest with many mountains that rise more than 500 above from the sea level.
These mountains served as starting point of the 189 major river basins in Malaysia. In Peninsular
Malaysia there are 89 major rivers basin present in Peninsular Malaysia, mean Borneo Island
comprised of 78 major river basins in Sabah and 22 major river basins in Sarawak. Amongst
these river are the place where the Tor species begins their lifecycle (Ng 2004). For Peninsular
Malaysia this Tor species is named as ‘Kelah’ fish. Differently in Borne Island, Tor species in
Sabah is locally known as ‘Pelian’ fish. Meanwhile, in Sarawak, Tor species is known as
‘Empurau’ fish that gives the meaning of ‘unforgettable taste’ in Chinese character. Riverine in
Peninsular Malaysia is divided into the East Coast (Kedah, Perak, Selangor and Negeri Sembilan
State), West Coast (Kelantan, Terengganu and Pahang State) and Southern State (Johor State).

Peninsular Malaysia is the home to Tor tambroides and Tor tambra. Currently, both Tor
species are considered as valid species despite controversial taxonomy classification (Pinder et
al. 2019). Most of the Tor species in Peninsular Malaysia found in upstream of the main river
basin which the surrounding area of river remained conserved and untouched by any
development activities. In West Coast, the northern state, which is Kedah, Ulu Muda River
serves as home to a large population of Tor species. Another rivers in Kedah that inhibited by
Tor species are Telian and Gawi River (Ng 2004). Moving south next to Kedah, Perak has the
most extensive riverine and home to second longest river in peninsular Malaysia. Previous report
has shown that Tor species inhibit Perak, Bernam, Karawang, Kejar, Mangga, Sungkai, Sara,
Sigor and Tiang River and also Temenggor Lake (Ng 2004). Selangor is the most advanced and
rapidly developing state in Malaysia with high population. Hence, increased number of forest
land transformation into residential or industry area to compete with the rapid growth of
population and economy (Aisyah et al. 2015). These lead to deprivations of river basin number
and increasing number of polluted rivers in Selangor which caused depletion of Tor species
population in this state. Previously Tor species has been reported that they inhibit Kancing,
Langat, Semenyih and Selangor River (Ng 2004). Lastly in Negeri Sembilan state, Tor species
has been reported that this fish inhibit Kampung Esok (Esa et al. 2011) and Serting River, Negeri
Sembilan (Esa et al. 2008).

The East Coast of Malaysia comprising of Kelantan (northern state of East Cast),
Terengganu and Pahang. These three states remain undeveloped and in last two decade the
number of Tor population is relatively high. In Kelantan, Tor species is reported inhibit Galas,
Nenggiri and Lebir River that meet together and form Kelantan River. However, recent report
has shown the failure in capturing Tor species especially Tor tambroides in Kelantan River
(Ambak and Zakaria 2010; Hashim et al. 2015). However, recent report have shown that Tor
species also found to inhibit the Pergau Lake, Jeli, Kelantan State (Alias et al. 2019). Further
south of Malaysia East Coast, Tor species in Terengganu mainly can be found in Kenyir Lake
(Farinordin et al. 2017; Mohamad et al. 2019). Tor species also found to inhibit the rivers that
flow into Kenyir Lake which include Tembat (Farinordin et al. 2017) Cacing, Terengganu and
Petuang River (Ng 2004). Recent study has shown that the population size of Tor species in
Kenyir Lake particularly Tor tambra is decreasing and rather alarming (Mohamad et al. 2019).
Rivers in Pahang State flown down to the South China Sea from Two range, Tahan Range and
Titiwangsa Range. Most of the upstream rivers remained conserved as these areas have been
gazetted as National Park. Pahang state also serves as major habitat for Tor species and recent
finding reported the presence of T. tambroides and Tor tambra in Keniyam River of National
Park Pahang (Mohd-Azham and Singh 2019; Norfatimah et al. 2014). Besides that, Tor species
also inhibit the longest river in Peninsular Malaysia, Pahang River (Asaduzzaman et al. 2017)
and Tembeling River, Pahang (Esa et al. 2008), Jelai, Lipis, Rompin and Merchong River (Ng
2004).

Previously several rivers in Johor, the Southern State of Peninsular Malaysia have been
reported inhibited by Tor such as Endau, Kinchin, Kemapan , Jasin and Marong River (Ng
2004). However, the study on Tor species in Johor remained limited and current study only
reported the presence of Tor spp population in Endau River (Ahmad et al. 2016). The two states
of East of Malaysia in Borneo Island, Sabah and Sarawak have large area of untouched and
conserved forest area. In Sabah, Moyog and Tenom River (Ng 2004) have been reported with the
presence of Tor species. The most common species found in this state are Tor tambra and Tor
tambroides. Meanwhile, in Sarawak State, the most common Tor species been reported is Tor
douronensis and Tor tambroides. Currently, in year 2019, Tor douronensis is been classified
synonym of Tor tambra (Pinder et al. 2019). This will be discussed further in the following
chapter. Generally Tor species in Sarawak can be found in most major rivers such as Trusan,
Limbang, Rejang, Layar, Sadong, Sarawak and Batang Lupar River. Despite Sarawak state has
22 river basins, this river basins comprised of large rivers and streams and remained unexplored.
Furthermore, Sarawak has the longest river in Malaysia. Most of the Tor species habitat reported
in the previously found in major rivers. However, due to large river system and the upstream
remain unexplored the exact number of Tor species population in Sarawak remains elusive.

Currently, all Tor species available in Malaysia is facing decreasing trend in population
size and deficient data. In order to prevent further extinction of Tor species in Malaysia
particularly in Peninsular Malaysia, several rivers and lakes have been gazetted as preservation
area for Tor species conservation. Amongst these rivers and lakes listed are Chilling River
(Tributary of Selangor River) which have been gazetted as Tor species sanctuary, Kenyir Lake in
Terengganu and Tiang and Kejok River and Temenggor Lake in Perak (Fuad Hadinata and
Amrullah 2017). Furthermore, there are limited number of study related to Tor species in
Malaysia. This will add on to data deficiency which limit the current update of Tor species
geographical distribution, taxonomy identification that become prolonged debate particularly to
resolve Tor douronensis validity and other components such as feeding, diseases and genetics
references.

Thailand

Freshwater fish aquaculture in Thailand has been practiced for an almost one decade and
continue to growth rapidly. In ASEAN region, Thailand is one of the countries contributed to the
major supply of freshwater fish either wild-captured or aquaculture. Mahseer species is one the
freshwater fish that contributes to the freshwater fish supply and recreational activity particularly
fishing game. Despite rapid growth of freshwater fish demand, aquaculture industry and fishing
activity, the number of publication on mahseers species particularly on Tor species remains
limited. The lake and river listed in the following paragraph are taken from previous findings for
over one decade and there is no current study of this species in Thailand. Therefore, from this
review we hope this will open an opportunity of new research and collection of more Tor species
to avoid uncertainty and further decreasing of Tor species population. Several Tor species has
been reported that inhibit Thailand lakes and rivers are Tor tambroides, Tor tambra, Tor
sinensis, Tor putitora and Tor douronensis. Amongst these species, Tor tambroides is widely
distribute in Thailand Peninsular.

Starting from northwestern part of Thailand, Tor tambroides found to inhibit rivers in
Chiang Mai and Mae Hong Son Province (Monkolprasit et al. 1997). Meanwhile northeastern
part of Thailand, from Nong Khai to Bueng Kan, Nakhon Phanom and Kaeng Tana National
Park serve as river basins to the 10 th longest river in world, Mekong River and home to Tor
tambroides species (Vidthayanon et al. 1997).In the Central Thailand, Tor tambroides inhibits
rivers in Chao Phraya, Phetchaburi, Uthai Thani, Kanchanaburiand and Maeklong
(Monkolprasit et al. 1997). Further down to south region of Thailand, Tor tambroides found to
inhibit rivers in Surat Thani province and Cheow Lan Lake, Khao Sok National Park
(Monkolprasit et al. 1997). Besides Tor tambroides, Tor tambra and Tor sinensis have been
reported that these two species inhibit the Mekong basin which lie in the northeastern part of
Thailand. Despite taxonomy conflict and considered as invalid, Tor douronensis has been
reported to inhibit several area in peninsular Thailand that include Chao Phraya and Mekong
basins. Furthermore this species also found to inhibit Si Sawat District where lies the Sinakharin
Lake, Chiang Mai and also Pattani River (Monkolprasit et al. 1997).

Vietnam

Vietnam is one of ASEAN countries that are among top 10 countries in fish producing globally
(Chan et al. 2017). Furthermore, fishing industry is a major industry in Vietnam that become
main source of income. There are more than five million people of Vietnam working in fishing
industry. Meanwhile, part of fish industry which is wild capture fishery in either marine or
freshwater fish employs more 1.4 million fishers. Vietnam geographically can be divided into
northern and southern region. The riverine system in Vietnam has been considered as most
complex and dense river network. Despite complex and dense riverine system, the number of
publication on Tor species in Vietnam remained limited. There are two studies mentioning the
presence of Tor species in Vietnam and both are focusing on the southern region. Several species
of Tor have been reported that inhibit the riverine system in Vietnam which are Tor tambra, Tor
tambroides, Tor sinensis, Tor mekongenesis and Tor dongnaiensis (Hoang et al. 2015; Van Sung
1995). Recent study by Hoang et al. 2015 shows the presence of Tor mekongenesis and Tor
sinensis in Krong No River. This river lies between two provinces Dak Lak and Lam Dong of
southern Vietnam region (Hoang et al. 2015). Besides that, his study also found that Tor
tambroides and Tor dongnaiensis inhibit the Đồng Nai River that lies in Đồng Nai Province of
southern Vietnam region (Hoang et al. 2015).

Myanmar

Besides Vietnam, Myanmar is also one of ASEAN country that majorly contributes in supplying
fish to the world fish market. The freshwater fish is the major source of fish farming which
provide an employments as full-time or part-time to the Myanmar people. Myanmar is the
second larger country in ASEAN that stretch from eastern tip of Himalayan Plateau to the south
and meet the Andaman Sea. Myanmar comprised of six major river basin which are; i)
Ayeyarwady (Irrawaddy)-Chindwin river basin, ii) Sittaung river basin, iii) Thanlwin (Salween
in Thailand, Nujiang in China) river basin, iv) Mekong (Lankang in China) river basin, v)
Rakhine (Arakan) coastal basin and iv) Tanintharyi (Tenasserim) coastal basin that flown from
north to south. Most of the reports mentioning the presence of Tor species in Myanmar’s river is
in the Thanlwin River. This originate from Himalayan Plateau and pass through China, Thailand
and Myanmar and flows into Andaman Sea. Locally, Tor species is called as Nga-dauk. There
are several Tor species reported that inhibit Myanmar’s river particularly Thanlwin River which
are Tor putitora (golden mahseer), Tor tambroides and Tor tor (Shahi et al. 2014). Despite
Myanmar is well known as the major source of fish globally, there is no current study that look
into the availability and population size of Tor species. The limitation of Tor species study raise
a big issue in the number of Tor species which might end up with the disappearance or rapid
decrease of this species in Myanmar.

Laos PDR

Geographically Laos PDR country is a land-locked country and the only ASEAN country
without sea. Therefore, the fish industry in Laos PDR is mainly depended on freshwater fish that
comes from river, reservoir, pond and lake. The fish industry in Laos PDR contribute around
13% of the country GDP and 60% of the fish sources are from the Mekong River and its
tributaries. Most of the main rivers which are Nam Ou River, Nam Suang River, Nam Khane
River, Nam Ngum River, Nam Sane River, Nam Theun/Kading River, Sebangfay River,
Sebanghieng River, Sedone River and Sekong/Xekong River in Laos PDR are the tributaries to
the Mekong River. These tributaries river in Laos PDR contributes the 35% of the water flow of
Mekong River. There are only two rivers namely Nam Ma and Song Chu rivers flows from Laos
PDR to the South China Sea through Thanh Hóa Province of Vietnam. Amongst these rivers,
previous study back in 1999 have found the present of Tor species which are Tor tambra, T.
sinensis, and T. ater that inhibit Nam Theun/Kading River (Roberts 1999). In recent study,
Sekong River tributaries, Se Kaman/Xe Kaman river is home to the several Tor species which
are Tor tambra, tor tambroides and Tor laterivittatus (Kottelat 2011). Up-to-date, there is no
other study that reporting the number of Tor species population in Laos PDR despite 75% of
protein source of Laos PDR people population are depending on the freshwater fish either wild-
capture or aquaculture (Phonvisay 2013).

Cambodia

Cambodia is a lowland country that lies between Thailand and Vietnam in western and eastern
border of Cambodia respectively and bordered with Laos PDR in the northern region.
Geographically, Cambodia is surrounded by the mountain range in north, east and southwest
region that serve as drainage origin and tributaries mostly primarily or secondary for the Mekong
River. The Mekong River is flows through into this country from Laos PDR and enter into Stung
Treng Province, in north-east Cambodia. This river bisecting the Cambodia and flows into
Phnom Penh, the Capital City of Cambodia before it enter Đồng Tháp Province in the Mekong
Delta region of Vietnam. As the earth surface of Cambodia are a lowland and floodplain,
Cambodia has the largest freshwater lake in ASEAN, Tonle Sap Lake that stretch form Phnom
Penh to Angkor Wat during raining season and the lake surface can expand up to 10,000 square
kilometers. During dry season the lake surface may shrink into 3,000 square kilometers.
Meanwhile, the southern region Cambodia is the coastal area that stretch along Thailand Gulf for
443 km. Despite Cambodia comprise of inland and marine water system, the fishing industry in
Cambodia mainly focused on freshwater fish fishing. Furthermore, freshwater fish is the main
animal protein source for Cambodia people (Vilain et al. 2016). Even though freshwater fish
have been captured and consume largely in Cambodia, the study focusing on the population size
of Tor species is almost unavailable. Previous study have shown that the Tor tambroides, Tor
tambra and Tor sinensis inhibit the Mekong River in Cambodia (Rainboth 1996; Rainboth et al.
2012). Unfortunately, there is no study to show the presence of Tor species in other river of
Cambodia or even in Tonle Sap Lake.

Indonesia

Indonesia is the largest island country in the world that comprise more than seventeen thousand
islands and separated from the mainland of Southeast Asia by the seas. The major island in
Indonesia comprised of Sumatra Island, Jawa Island, Borneo Island (Kalimantan), Sulawesi
Island, Nusa Tenggara Island and Papua Island. With large riverine system and surrounded by
sea, Indonesia is amongst the top countries globally in supplying fish for both freshwater and
marine fish. Despite Indonesia is separated from the mainland of Southeast Asia, Indonesia
somehow share a similar Tor species with some of ASEAN countries such Malaysia, Thailand,
Vietnam and Cambodia. The Tor species reported that inhibit Indonesia’s rivers are Tor
tambroides, Tor tambra and Tor douronensis (Tamba et al. 2019). The widely distribution of Tor
species across Southeast Asia mainland across into Borneo Island, Sumatra Island and Java
island of Indonesia have been postulated that the Mekong River system is connected with the
river system of these island during Pleistocene time as Sunda Shelf (Hanebuth et al. 2011). After
the rising of sea level, the area are separated and forming into on an Island as it is now.

In Indonesia, the mahseers or Tor species is locally known or called as Keureling Fish.
Starting from Sumatra Island, the Tor species reported that inhibit riverine system in this island
are Tor tambroides, Tor douronensis, Tor tambra and Tor soro (De Silva et al. 2004). Currently
Tor soro has been classified as Neolissochilus soro (Khaironizam et al. 2015). In Sumatra,
several studies have reported the presence of Tor species. Current study shows the presence Tor
tambroides in found to inhibit the upstream of Wampu Waters, north region of Sumatra Island
(Tamba et al. 2019). Another study also show that Tor tambroides are presence in the Manna and
Tarusan River of western region of Sumatra Island (Wibowo and Dwirastina 2016) and also ,
Bahorok River and Berkail River. Meanwhile, study in Batang Toru River System, South
Tapanuli, North have shown the presence of several Tor species which are Tor tambra, T.
douronensis, and T. tambroides (Roesma et al. 2016). Furthermore study in Kreuang Sabee
River, Aceh Jaya District, Indonesia shows the presence of Tor tambra species only (Timorya et
al. 2018). Despite Tor douronensis has been classified as invalid species, this species has been
widely studied in Indonesia and found widely distributed in several rivers in Sumatra Island
which are Batang Gumanti River, Batang Antokan River, Batang Malalo River, Batang
Matur River, Batang Sinuruik River and Lubuk Mangkuih River (Roesma et al. 2017).

Moving toward south of Sumatra, Java Island is also a home to Tor species (Roberts
1993). Recent study shows that Tor tambroides present in West Java river particularly in
Cimanuk River that flows from Pappandayan Mountain into Java Sea (Herawati 2019). Another
recent study shows that Tor douronensis inhibit riverine system surrounding Cereme Mountain
located in Kuningan District, West Java (Abidin et al. 2018). The river system area that inhibited
by Tor douronensis are Balong Dalem, Darma Loka, Cigugur, Cibulan and Bale Kambang
(Abidin et al. 2018). In Kalimantan of Borneo Island, Tor tambroides have reported that this
species presence in riverine system of Muller Mountain in Central Kalimantan (Akmal and
Rahardjo 2018; Haryono 2008). Another previous study shows that Tor tambroides and Tor
tambra are present in Mendalam river system, Betung Kerihun National Park, West Kalimantan.
As compared to Sumatra Island, the number of study on Tor species in Java Island and
Kalimantan are limited. However, in Bogor, South Jakarta, Java Island, the Research Institute for
Freshwater and Fisheries Extension is actively study on Tor douronensis and Tor tambroides for
aquaculture and preservation purpose (Cahyanti et al. 2019).

Brunei, Singapore & Philippines

Brunei is a small country located in Borneo and surrounded by Malaysia. Brunei comprised four
major rivers which are Belait River, Tutong River, Temburong River and Brunei River and
the Brunei people are a river-based civilization which they depends on river resources
particularly freshwaterfish for animal protein (Bac-Bronowicz and Becek 2009). Recent study
has shown that Tor tambroides and Tor tambra inhibit Brunei riverine system (Sulaiman et al.
2018). However the population size of these two species remained unknown. There are two
ASEAN countries which are Singapore and Philippines have no recent study on Tor species. In
Singapore, the Tor species particularly was present in Singapore riverine system back in 1966
(Alfred 1966). However, in year 1997 the Tor tambroides has been classified as extinct species
in Singapore (Ng and Lim 1997). Meanwhile, for Philippines, this country is not part of Sunda
Shelf during Pleistocene time. Therefore the river system in Philippines is not connected with
any mainland of Southeast Asia and Tor species are unavailable.

Morphological features of ASEAN’s Tor species

Tor tambroides

Amongst Tor species in ASEAN countries, Tor tambroides is broadly distributed throughout
Myanmar, Thailand, Laos PDR, Cambodia, Peninsular Malaysia, Sumatra, Java, and Borneo
(Pinder et al. 2019). Generally, the color of Tor tambroides are silver-bronze and reddish. The
unique morphological feature specifically represent Tor tambroides is the presence of upper
median projection. Beside that, another features that commonly share with other Tor species are
Tor tambroides have sub-terminal mouth position, long lower median lobe, pointed rostrum hood
and equal caudal fin lobe. Previously, Tor tambroides has been suggested that this species is a
synonym of Tor tambra. However, short lower median and absence of upper median projection
in Tor tambra distinguishing this species from Tor tambroides. Furthermore, another argument
has been stated that Tor dongnaiensis and Tor tambroides are similar species. However, the
absence of upper median projection in Tor dongnaiensis separate this species from Tor
tambroides. Distinct morphological features from Tor tambra and Tor dongnaiensis indicates
that Tor tambroides is a valid species.

Tor tambra (Valenciennes, 1842)

Similar like Tor tambroides, Tor tambra can be found in many riverine system of ASEAN
countries including the Mekong River system, Thailand, Peninsular Malaysia, Sumatra Island,
Java Island and Borneo Island (Walton et al. 2017). The colour of this species varies depending
on the locality which are reddish, olive, dark or slightly olive (Hoang et al. 2015; Walton et al.
2017). The morphological features that differentiates Tor tambra from Tor tambroides are
various size of lower median lobe and blunt rostral hood. Another feature of Tor tambra closely
share with other Tor species are absent of upper median projection and equal caudal fin.

Tor douronensis

Taxonomy of Tor douronensis is the most complex taxonomy amongst Tor species which which
has been debated since 1999 and prolong until now (Kottelat 2013). According to the California
Academy of Science's Catalog of Fishes website, Tor douronensis has been classified as valid
species. However, several researchers including Roberts (1999) and Kottelat (2013) have debated
about validity of Tor douronensis as there is no specific morphological feature that can
distinguish between this species and Tor tambra. Hence, Kottelat (2013) has postulated that Tor
douronensis found in Peninsular Malaysia and Borneo Island of Sabah and Sarawak is a
synonym of Tor tambra. Despite unstable taxonomy, researchers in Indonesia particularly in
Sumatra Island postulated that Tor douronensis is still a valid species. Furthermore, there are
several recent studies that have been done and reporting about this species. Morphologically, the
Tor douronensis has short lower median lobe, absent of upper median projection, blunt rostral
hood. In addition, the color of Tor douronensis is silvery with back darkish. These
morphological features of Tor douronensis is almost similar to Tor tambra. Furthermore (Wei
and Cui 1996). However, study done by Haryono and Tjakrawidjaja 2006 shows that three Tor
species, Tor tambroides, Tor douronensis and Tor tambra from one source are distinguishable
from each other through body measurement (Haryono and Tjakrawidjaja 2006). From this study
the body measurements found to differentiate Tor species are interorbital width, caudal peduncle
length, caudal peduncle depth, head width and body depth (Haryono and Tjakrawidjaja 2006).
Interestingly, Tor douronensis is different with Tor tambra with small interorbital with and large
caudal peduncle length. However, for further confirmation of validity of this species
morphological and genetic comparison between Tor tambra and Tor douronensis from Mekong
riverine system, Peninsular Malaysia, Borneo Island, Sumatra Island and Java Island should be
done to dissolve this unstable taxonomy.

Tor dongnaiensis

Tor dongnaiensis ia one of the Tor species mainly found in Vietnam. As mentioned above, this
species has been classified under Tor tambroides specification due to similar morphological
features. However, the finding has shown that Tor dongnaiensis has yellow to grey body color,
the mouth morphological features with absent of median projection of upper lips and unequal
caudal fin distinguished this species from Tor tambroides. Generally, the appearances of this
species are conical head shape, pointed snout, subterminal mouth with fleshy lips, long lower lip
of median lobe and straight with pointed rostral hood.

Tor tor
Most of the Tor species found in Myanmar is originated from Himalayan Plateau. Tor tor which
found inhibited Myanmar riverine system has been reported that this species presents in India
river system that include Ganga, Brahmaputra and Narmada river system. The prominent
characteristics that differentiate Tor tor from others Tor species in Myanmar are big head and
large scales, subterminal mouth with interrupted fold of lower lip, two pairs of large barbel and
lateral-line scales.

Tor mekongenesis

Mekong riverine system inhibited by several Tor species. Furthermore, amongst these Tor
species, Tor mekongenesis is the only Tor species specifically present Mekong River. General
appearance of Tor mekongenesis are as follow; longer head, blunt snout, blunt rostral hood,
subterminal mouth with fleshy lips and short median lobe of lower lip.

Tor sinensis

Tor sinensis is another Tor species predominantly inhibit Mekong riverine system. This species
has its own unique characteristics that differ from others Tor species in the same riverine system.
First, to differ from Tor ater, Tor sinensis present with having a smaller number of lateral,
predorsal, transverse row scales, long median lobe in lower lip and absent of strip alongside the
body. As Tor sinensis have similar features of Tor tambroides, large ratio of standard length over
body depth and the presence median projection of upper lip distinguished between these two
species. In comparison with Tor mekongenesis, Tor sinensis uniquely having prolong rostral
hood and various number of lateral scales between 23-28. The color of Tor sinensis usually dark
on head and back with grey-silver or yellowish color side of the body. Furthermore, unlike other
Tor species, Tor sinensis shows the presence of longitudinal stripe appears from head until
caudal fin.

Tor putitora

In India, this Tor putitora is known as golden mahseer. The color of this species appears greenish
and silvery on side of the body which then turn reddish yellow or gold color on anal and pectoral
fish that represent its name. Similar like Tor tor which originated from Himalayan Plateau, this
species is endemic only inhibit in Myanmar riverine system. The unique characteristics of this
species are having elongated and nearly straight body, small mouth with lower jaw slightly short
than upper jaw and deeply forked caudal fin.

Tor laterivittatus

Tor laterivittatus is predominantly inhibit in Laos PDR riverine system. The characteristic of Tor
laterivittatus that unique from others Tor species are elongated median lobe of lower lip, the
upper lip rolled upward and backward, the presence of longitudinal stripe along the body side
and deeply concave dorsal.

Tor ater

Another unique Tor species that predominantly inhibit Laos PDR is Tor ater. This species is the
smallest Tor species as compared to other ASEAN’s Tor species. Generally, Tor ater is
characterized with having short median lobe, the presence of longitudinal dark stripe on the body
and all fins appear in dark color or black color.

Genetic identification and specification of ASEAN’s Tor species

The evolution and adaptation process that can cause genetic changes results in phenotypical and
morphological changes of a species. There are several factors that contribute to the evolution and
adaptation process which include temperature, pH, food, environment such difficulty in finding
source, water sources, and many more. Solely depends on morphological features create
confusion and question in species specification. Generally, animal mtDNA comprised of 16-17
kbps in size and consist of 22 tRNA-encoding and 13 protein-encoding gene which are NADH-
ubiquinone oxidoreductase chain 1-6 (ND1-6), NADH-ubiquinone oxidoreductase chain 4L
(ND4L), cytochrome complex III (Cytb), Cytochrome c oxidase complex IV subunit;
cytochrome c oxidase subunit I-III (COX 1-3) and lastly complex V subunits; ATPase 6 (ATP6)
and 8 (ATP8) (Yang et al. 2014). The characteristic of animal mtDNA are uniparental
inheritance through maternal lineage, non-recombining and higher mutation rate as compared
with nuclear DNA (DeSalle et al. 2017). In addition, mtDNA is postulated that it is polymorphic
at the intraspecific level (Ladoukakis and Zouros 2017). With these characteristics, mtDNA has
used as phylogenetic analysis for species identification and specification for over than two
decades.
 For more than two decades identification and specification of Tor species particularly in
amongst ASEAN’s countries remains unstable until now. With the advancement of molecular
techniques, genetic phylogenetic analysis helps to improve the species specification with
combination morphological features. None of the ASEAN’s Tor species even besides ASEAN
countries have complete whole genome DNA sequence. However, in 2014 research group from
Malaysia has successfully sequenced the full mitochondrial DNA (mtDNA) sequence of Tor
Tambroides from Malaysia (Norfatimah et al. 2014). Despite complete mitochondrial DNA been
sequenced in 2014, some of the gene sequence of present in mitochondria such as COX1, 16S
rRNA and ATP6 has been sequenced before year 2014 and used as phylogenetic analysis for Tor
species identification and specification. Besides mtDNA, another DNA sequence which is
microsatellite region of nuclear DNA also been used in identification and specification of Tor
species. Despite various genetic identification and specification, up-to-date there is no study on
comparison of all ASEAN’s Tor species.
COX1

Cytochrome c oxidase subunit I (COX1) is a gene that encode for one of three subunit of
Cytochrome c oxidase complex IV. COX1 gene has been used in Tor identification and
specification in ASEAN countries. Study done by Hoang et al. 2015 and Wibowo et al. 2013 in
Southern Vietnam and Sumatra Island, Indonesia, respectively were using the 5’ region of COX1
gene (Ivanova et al. 2007; Ward et al. 2005) for identification and specification of Tor
tambroides, Tor dongnaiensis, Tor mekongenesis and Tor sinensis (Hoang et al. 2015; Wibowo
et al. 2013). The selection of the 5’ region of COX1 gene was due to availability of primer aiding
recovered from broad range of species (Hebert et al. 2003). In addition, this 5’ region of COX1
gene has been used for identification and speciation widely in Indonesia (Walton et al. 2017).
Meanwhile, study by Esa et al. 2013, 2008 have used 3’ region of COX1 gene for identification
and speciation for Tor tambroides and Tor douronensis in Malaysia (Esa et al. 2013; Esa et al.
2008).

16S rRNA

COX1 gene previously has been proposed to be the powerful universal marker for animal
particularly vertebrate identification and specification. However, another gene sequence in
mitochondria which is 16S rRNA gene sequence has been introduced for vertebrate identification
and specification. Generally, 16S rRNA gene sequence is encoded for ribosomal RNAs that
involved in translation of mRNA into mitochondrial protein. Previous study have shown that 16S
rRNA gene sequence has fulfill the criteria for marker in species identification. Despite this
sequence is highly conserved, this sequenced also comprised of polymorphism sequence due to
high number of deletion and insertion (Sarri et al. 2014). Nguyen et al 2008, 2006 has been used
this sequence for identification and speciation of Tor tambroides and Tor douronensis in Borneo
Island of Sarawak (Nguyen et al. 2006). Furthermore, recent study shows that, 16S rRNA gene
sequence was used to identify Tor tambroides species in Peninsular Malaysia (Asaduzzaman et
al. 2018). Even though, 16S rRNA gene sequence has used and classified as universal marker,
current finding show that this sequence are unable to differentiate two species from different
genus (Mohanty et al. 2015).
Cyt b and ATPase 6

Beside COX1 and 16S rRNA gene sequence, Cyt b gene sequence also has been nominated as
reliable and most useful universal marker for fish identification and specification (Arif and Khan
2009). Cyt b gene sequence is encoded for cytochrome b protein which involve in performing
electron transfer reaction. The length of Cyt b gene is about ~1143 bps with low variability.
However, longer sequenced is postulated to increase or improve the efficiency of species
identification and specification (Patwardhan et al. 2014). Another mitochondrial gene sequence
used for universal marker is ATPase 6 gene sequence. ATPase 6 gene sequence is encoded for
ATP synthase membrane subunit 6. This gene sequence is characterized by the high rate of
mutation which suitable to distinguish the species with high haplotype diversity (Divya et al.
2015). In term of identification of Tor species, Nguyen et al 2008 has used both Cyt b and
ATPase 6 together with 16S rRNA gene sequence to improvise the efficiency for identification
and specification of Tor tambroides and Tor douronensis in Borneo Island, Sarawak Malaysia
(Nguyen et al. 2008).

Microsatellite DNA marker

Microsatellites is a repetitive sequence of DNA with certain motifs up to 50 times and occurs at
more than thousands of location and widely distributed within the DNA sequence. This
microsatellites also known as simple sequence repeats (SSR) or simple sequence length
polymorphisms (SSLP). This sequence highly susceptible to the DNA mutation as compared to
other region of DNA which result in high genetic diversity. Furthermore, microsatellites is also
known as polymorphic DNA loci that comprised repeated sequence which useful for universal
marker for fish identification and specification (Vieira et al. 2016). This marker has been used by
Nguyen et al. 2007, 2008 for identification and specification of Tor tambroides and Tor
douronensis in Island Borneo, Sarawak Malaysia (Nguyen et al. 2007; Nguyen 2008).
Furthermore, similar technique with using microsatellites marker from other cyprinid also been
used for identification and specification of Tor tambroides and Tor douronensis in Borneo
Island, Sarawak, Malaysia (Esa et al. 2011).
Future fate of ASEAN’s Tor species

From this review we have notified that the number of publication related to Tor species in
ASEAN countries is decreasing and remained limited. Some of the ASEAN countries including
Thailand, Laos PDR, Cambodia and also Borneo Island have no recent study and previous
publication related to Tor species back to 5 years ago. Furthermore, some of the recent studies
show that the disappearance of Tor species at a river during ichthyology study showing a high
possibilities of decreased population number of Tor. Research finance or resources in developing
countries would be a major challenges and limitation for the researchers to conduct population
study of Tor species. Therefore, forming a group research throughout ASEAN countries,
focusing on Tor research in ASEAN and sharing the resources would help the developing
countries to update the current population number and its availability in the riverine system.

Another obstacle in Tor species study is unstable taxonomy which result in difficulties
for the identification and specification. Previous studies have shown that various universal
marker have been used by every different research group for identification and specification of
ASEAN’s Tor species. These independent and truncated data results from various approach
further ameliorate the instability of Tor taxonomy. Hence, requires more researches and
resources to fill up the data and knowledge gap. Consequently, this review would provide new
direction and platform to improve the identification and specification of ASEAN’ Tor species by
using similar and standardize approach.

Out of nine ASEAN’s Tor species, eight has been listed as red list species by IUCN Red
List. Three of them are classified as endangered species and the rest are deficient in data.
Currently, with the unknown status of ASEAN Tor species, conservation of Tor species would
be the priority and responsible for each ASEAN countries. Due to the rapid development in
developing countries, lot of river basin disappear, increased polluted rivers, dam development
further contribute to the decreasing population number of ASEAN’s Tor species. Previously,
conservation of Tor species are relies on captive breeding in the farm. Several place known to
conduct in house conservation program that include Freshwater Fisheries Research Centre,
Glami Lemi, Titi, Jelebu, Negeri Sembilan and Inland Fisheries Research and Production Centre,
Tarat, Sarawk in Malaysia and also Research Institute for Freshwater Aquaculture and Fisheries
Extension, Bogor in Indonesia. However, there are several limitations that would challenge the
aquaculture process. That limitation include asynchronous spawner behavior of Tor species,
limitation of parental breeder, slow in growth and maturation and genetic diversity. These
limitation would requires further study to overcome the limitation and understand further the
characteristic and behavior of ASEAN’s Tor species. From this review, we found the necessity
of understanding the genetic of ASEAN’s which will provide an overview of ASEAN’s Tor
species in term of the growth, maturation, breeding, and feeding process.

Conclusion

Up-to-date, the current population status and taxonomy within ASEAN’s Tor genus remained
elusive and unstable respectively. Furthermore, absence of whole DNA genome sequence further
widen the knowledge gap of ASEAN’s Tor species. With this review, it would help to update our
current understanding about ASEAN’s Tor species and open up the new direction of ASEAN’s
Tor research.

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