First records of Croaking Gourami, Trichopsis vittata (Cuvier, 1831) (Teleostei: Osphronemidae), from Myanmar and Bangladesh

Abstract

The Croaking Gourami, Trichopsis vittata (Cuvier, 1831), is native to Southeast Asia and Sundaland, with introductions reported from USA, Philippines and India. The species was found by us in Myanmar (1997 and 2013), and Bangladesh (2014 and 2016). DNA analysis supports the view that T. vittata is a species complex. Specimens from Bangladesh, Myanmar and the European aquarium trade are the same genotype as specimens from Thailand, possibly corresponding to Trichopsis harrisi Fowler, 1934, considered a synonym of T. vittata. Non-native populations are likely due to release from aquarium specimens.

Full text

First records of Croaking Gourami, Trichopsis vittata (Cuvier, 1831)
(Teleostei: Osphronemidae), from Myanmar and Bangladesh
Michael Norén,1 Sven O. Kullander,1 Md. Mizanur Rahman2, Abdur Rob Mollah2
1 Department of Zoology, Swedish Museum of Natural History, P.O. Box 50007, SE-104 05, Sweden. 2 Department of Zoology, University of
Dhaka, Dhaka-1000, Bangladesh.
Corresponding author: Michael Norén, michael.noren@nrm.se
Abstract
The Croaking Gourami, Trichopsis vittata (Cuvier, 1831), is native to Southeast Asia and Sundaland, with introduc-
tions reported from USA, Philippines and India. The species was found by us in Myanmar (1997 and 2013), and
Bangladesh (2014 and 2016). DNA analysis supports the view that T. vittata is a species complex. Specimens from
Bangladesh, Myanmar and the European aquarium trade are the same genotype as specimens from Thailand, possibly
corresponding to Trichopsis harrisi Fowler, 1934, considered a synonym of T. vittata. Non-native populations are
likely due to release from aquarium specimens.
Key words

Academic editor: Zeehan Jaafar | Received 13 January 2017 | Accepted 16 May 2017 | Published 12 July 2017
Citation: Norén M, Kullander SO, Rahman MM, Mollah, AR (2017) First records of Croaking Gourami, Trichopsis vittata (Cuvier, 1831) (Teleostei:
Osphronemidae), from Myanmar and Bangladesh. Check List 13 (4): 81–85. https://doi.org/10.15560/13.4.81
Check List 13(4): 81–85
https://doi.org/10.15560/13.4.81
4
13
81–85
Date 2017
Copyright Norén et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use,
distribution, and reproduction in any medium, provided the original author and source are credited.
Introduction
Trichopsis vittata (Cuvier, 1831), known as Croaking
         
      
Java, Borneo, Sumatra, Peninsular Malaysia, Thailand,
and the Mekong basin in Cambodia, Laos, and Viet-
nam (Kottelat 1985, Rainboth 1996, Baird et al. 1999).
Trichopsis vittata      
stagnant waters with abundant vegetation, such as rice
        
to survive in brackish water up to 20 psu salinity, and
        
2016). It feeds on small planktonic crustaceans and insect
larvae (Rainboth 1996). The male is territorial and builds
a small inconspicuous bubble nest among the vegetation,
in which the eggs are deposited and guarded by the male
until the larvae become free-swimming (Britz et al. 2001,
Liengpornpan et al. 2006). Due to its small size there is
    T. vittata, but it is occasionally
sold in markets as part of mixed catches, and is regularly

The FAO Introduction of Species database lists T.
vittata as introduced only to the USA, where a feral
population has persisted in southern Florida since the
  and Pecora 2013), but
established feral populations of T. vittata have also been
reported from the Philippines (BFAR 2006), without sup-
porting data, and most recently from Chennai in Tamil
Nadu, India (Knight and Balasubramanian 2015). Here
NOTES ON GEOGRAPHIC DISTRIBUTION

82 Check List 13(4): 81–85
we present additional localities from Myanmar and Ban-
gladesh, up to 1000 km west of the natural range of the
species and 1500 km northeast of the previously known
feral population in Chennai, and discuss the origin and
future development of feral T. vittata in South Asia.
Methods
All vouchered localities in Myanmar and Bangladesh are
presented in Table 1 and mapped in Figure 1. All listed
specimens were collected using beach seine or handheld
  
       -
chondrial cytochrome c oxidase subunit I (COI) gene
was sequenced from 5 T. vittata collected in Bangladesh,
6 collected in Myanmar, and 3 aquarium specimens
(GenBank accession numbers: KT250366, KY327366,
KY327367) purchased in Sweden, using the primers and
protocol of Ward et al. (2005).
The obtained DNA sequences were assembled in
Geneious version 9 ( et al. 2012), and combined
       T. vittata, T.
cf. vittata, or T. sp. cf. vittata deposited on GenBank, to
produce a matrix from which to estimate sequence diver-
gence as pairwise p-distance
nucleotide sites between 2 compared DNA sequences
(please note that this is a simple measure of dissimilar-
      
term p-value). A neighbor-joining majority rule bootstrap
tree (Jukes-Cantor model, 500 repetitions, random addi-
tion sequence) rooted with Betta macrostoma (GenBank
Accession number KM485437) was produced to visual-
ize genetic similarity (not shown) The Identication tool
of the Barcode of Life portal (http://boldsystems.org) was
used to obtain the Barcode of Life Barcode Index Num-
bers (BINs) of clusters of similar published sequences,
corresponding to putative species. Voucher specimens are
deposited in the collections of Dhaka University and the
Swedish Museum of Natural History.
Published occurrence data used to construct occur-
rence map (Fig. 2) were obtained from the Academy
of Natural Sciences at Philadelphia, Auburn University
Museum of Natural History, California Academy of
Sciences, Field Museum, Florida Fish and Wildlife Con-
servation Commission, Illinois Natural History Survey,
GBIF-MNHN (Paris), Oregon State University, Royal
Ontario Museum, Florida Museum of Natural History,
University of Michigan Museum of Zoology, United
States National Museum of Natural History, Smithsonian
Institution, Yale University Peabody Museum, and Natu-
ral History Museum of Denmark (Accessed through the
Fishnet2 Portal, , 2015-5-26).
Collecting was made with permission from the Depart-
ment of Environment in Bangladesh to the Department of
Zoology, University of Dhaka, and from the Department
of Fisheries in Myanmar to Swedish teams.
Table 1. Voucher data for the non-native Croaking gourami (Trichopsis vittata) recorded from Myanmar and Bangladesh. NRM = Swedish Museum of Natural History; USNM = National Museum of Natural History,
Smithsonian Institution (database record only).
Voucher numbers nCountry Genbank Acc# River drainage Locality Lat. °N Lon. °E Date
NRM 39950 1 Myanmar Salween Roadside ditch at Zayitchaung village, on the road Kyaikto-Thaton, 5 km before Thaton 17.0131 97.3356 16 Mar. 1997
USNM 378943 6 Myanmar Salween Kawkareik market — — 1 Jan. 2002
NRM 58106 1 Myanmar Salween Thathon market 16.9271 97.3621 17 Mar. 2008
NRM 58339 1 Myanmar Salween God Chaung, 2 miles on the way from Thathon to Pha-An 16.9086 97.3887 17 Mar. 2008
NRM 58410–58584, 67131 101 Myanmar KT250368 Salween Roadside ditch near bridge over Salween River at Tayoke Hla village, 5 miles to Pha-An 16.8331 97.6261 17 Mar. 2008
NRM 65183, 65291, 66324 12 Myanmar KY327369 Ayeyarwaddy Lake in small village, north of Nyaungdon, 68 km NW Yangon 17.1036 95.5895 24 Nov. 2013
NRM 65142, 65145, 65146, 66311–66312,
66395–66398
83 Myanmar KY327368
KY327370
Yangon Irrigation channel on road Yangon - Pathein, 25 km NW Yangon 16.9113 95.9441 24 Nov. 2013
NRM 67073 1 Myanmar KY327372 Ayeyarwaddy Small stream in village, alongside road, 3 km NE Pantanaw, 75 km WNW Yangon 16.9992 95.4884 24 Nov. 2013
NRM 67056-67058 3 Myanmar KY327371 Ayeyarwaddy Small stream in Kha Yaykan village, 9 km N Innma village, 95 km NW Yangon 17.1451 95.3312 25 Nov. 2013
NRM 66672-66674, 66676–66681, 67094–67095 65 Bangladesh KY349109 Old Brahmaputra Turag River at Kamarpare, near Dhaka city 23.8985 90.3844 1 Dec. 2014
NRM 66578–66581, 67096 12 Bangladesh KT250365
KT250367
Old Brahmaputra Fish market in Shonbari, Sreenagar 23.544 90.2964 2 Dec. 2014
NRM 69396, 69461 2 Bangladesh KY327373
KY327374
Old Brahmaputra Dhaleshwari River near Abdullahpur Bridge, Keranigonj 23.3850 90.2046 25 Feb. 2016
NRM 69220, 69434–69435 3 Bangladesh Old Brahmaputra Turag River at Ashulia point near Dhaka city 23.5135 90.2049 5 Mar. 2016
NRM 68345, 69493 3 Bangladesh Meghna Shorail, roadside ditch 8 km north of Brahmanbaria 24.0472 91.1026 19 Mar. 2016
NRM 68562 5 Bangladesh Meghna Titas River in Akhaura 23.884 91.2012 19 Mar. 2016

Norén et al. | First records of Trichopsis vittata from Myanmar and Bangladesh 83
Results
Trichopsis vittata can be separated from the 2 other cur-
rently recognized species in the genus, Trichopsis pumila
(Arnold, 1936) and Trichopsis schalleri Ladiges, 1962, by
the presence of a thin dark line below the eye. Trichopsis
vittata also grows to a larger maximum size, and normally
has 3 longitudinal dark bands on the posterior half of the
body instead of 2 as in T. schalleri and T. pumila (in T.
pumila the upper of the 2 bands is broken into a row of dark
spots surrounded by iridescent scales) (Fig. 2). Rainboth
(1996) states that T. vittata grows up to 70 mm standard
length (SL). Sithtananan (2010) measured 848 individuals
from all over the native distribution and found a maximum
length of 48.8 mm SL, and average length 31 mm SL.
Discussion
There are no published records of T. vittata from Bangla-
desh or Myanmar previous to the present report, not even
from the Myanmar portion of the Mekong River, but the
species has been common in collections from Indochina
and Indonesia for more than a century, suggesting that
the Tenasserim and Sino-Burman ranges separating the
Indochinese watersheds from the Burmese have been
         
from the Swedish Museum of Natural History collected
a single T. vittata from a roadside ditch 5 km north of
the town of Thaton, in Mon State in eastern Myanmar.
Table 1. Voucher data for the non-native Croaking gourami (Trichopsis vittata) recorded from Myanmar and Bangladesh. NRM = Swedish Museum of Natural History; USNM = National Museum of Natural History,
Smithsonian Institution (database record only).
Voucher numbers nCountry Genbank Acc# River drainage Locality Lat. °N Lon. °E Date
NRM 39950 1 Myanmar Salween Roadside ditch at Zayitchaung village, on the road Kyaikto-Thaton, 5 km before Thaton 17.0131 97.3356 16 Mar. 1997
USNM 378943 6 Myanmar Salween Kawkareik market — — 1 Jan. 2002
NRM 58106 1 Myanmar Salween Thathon market 16.9271 97.3621 17 Mar. 2008
NRM 58339 1 Myanmar Salween God Chaung, 2 miles on the way from Thathon to Pha-An 16.9086 97.3887 17 Mar. 2008
NRM 58410–58584, 67131 101 Myanmar KT250368 Salween Roadside ditch near bridge over Salween River at Tayoke Hla village, 5 miles to Pha-An 16.8331 97.6261 17 Mar. 2008
NRM 65183, 65291, 66324 12 Myanmar KY327369 Ayeyarwaddy Lake in small village, north of Nyaungdon, 68 km NW Yangon 17.1036 95.5895 24 Nov. 2013
NRM 65142, 65145, 65146, 66311–66312,
66395–66398
83 Myanmar KY327368
KY327370
Yangon Irrigation channel on road Yangon - Pathein, 25 km NW Yangon 16.9113 95.9441 24 Nov. 2013
NRM 67073 1 Myanmar KY327372 Ayeyarwaddy Small stream in village, alongside road, 3 km NE Pantanaw, 75 km WNW Yangon 16.9992 95.4884 24 Nov. 2013
NRM 67056-67058 3 Myanmar KY327371 Ayeyarwaddy Small stream in Kha Yaykan village, 9 km N Innma village, 95 km NW Yangon 17.1451 95.3312 25 Nov. 2013
NRM 66672-66674, 66676–66681, 67094–67095 65 Bangladesh KY349109 Old Brahmaputra Turag River at Kamarpare, near Dhaka city 23.8985 90.3844 1 Dec. 2014
NRM 66578–66581, 67096 12 Bangladesh KT250365
KT250367
Old Brahmaputra Fish market in Shonbari, Sreenagar 23.544 90.2964 2 Dec. 2014
NRM 69396, 69461 2 Bangladesh KY327373
KY327374
Old Brahmaputra Dhaleshwari River near Abdullahpur Bridge, Keranigonj 23.3850 90.2046 25 Feb. 2016
NRM 69220, 69434–69435 3 Bangladesh Old Brahmaputra Turag River at Ashulia point near Dhaka city 23.5135 90.2049 5 Mar. 2016
NRM 68345, 69493 3 Bangladesh Meghna Shorail, roadside ditch 8 km north of Brahmanbaria 24.0472 91.1026 19 Mar. 2016
NRM 68562 5 Bangladesh Meghna Titas River in Akhaura 23.884 91.2012 19 Mar. 2016
Figure 1. Distribution of Trichopsis vittata in Asia, based on occurrence records in FishNet II (1906–2014), Knight and Balasubramanian
(2015), and University of Dhaka and Swedish Museum of Natural History records (Myanmar 1997–2013; Bangladesh 2014, 2016). BD =
Bangladesh.
Figure 2. Trichopsis vittata. A. Live male specimen photographed
in aquarium (not preserved), B. Preserved specimen (NRM67094c),
32.8 mm SL, collected from Turag River, Bangladesh, 2014. The dark
pigmented line below the eye is a diagnostic character for T. vittata.
The individual was a juvenile, showing that the species
was already reproducing in the region. In 2008, numer-
       
Kayin State in eastern Myanmar, and in 2013 we found
      

84 Check List 13(4): 81–85
Myanmar west of the city of Yangon, and additional
specimens were obtained in the Kawkareik district in
eastern Myanmar by Mr. U Tin Win. The earliest records
of T. vittata from Myanmar are from the eastern part of
the country, but no collections are available from the
same time in western Myanmar, so the dispersal within
Myanmar should not be assumed to have been from east
to west. In 2014 we found the species in abundance in the
-
     
Sreenagar (Dhaka Division) in central Bangladesh (Table
1). The species was recorded again in 2016 from the
Turag River, and also in the nearby Dhaleswari River as
well as 2 localities in the lower Meghna River northeast
of Dhaka (Table 1). No T. vittata were found in southeast-
ern Bangladesh (Chittagong Division) during a survey in
2015, or in northeastern Bangladesh (Sylhet Division),
surveyed in 2016, despite the presence of suitable habitat.
Panijpan et al. (2015) analyzed mitochondrial COI
and nuclear RAG1 DNA sequences of a large number of
Trichopsis from Southeast Asia, and found that T. vittata
is not a single species but a species complex comprising
4 ecologically and morphologically similar but geneti-
cally distinct biological species. However, there were no
DNA sequences available from Java, the type locality
of T. vittata (Roberts 1996), and Panijpan et al. (2015)
were unable to determine which, if any, of these 4 clades
represented the true T. vittata. Dahruddin et al. (2016)
published 6 sequences of T. vittata from Java, which
in our analysis correspond to the clade that Panijpan et
al. (2015) referred to as “Trichopsis cf. vittata 4”, from
southern Thailand (BIN cluster ID BOLD:AAB9368).
As a coarse empirical rule-of-thumb, for the COI
gene, the dissimilarity between any 2 members of the
same species is typically < 1%, while the dissimilarity
 
(Ward 2009). This disjunct distribution of genetic similar-
ity has been termed “the barcode gap” (Meyer and Paulay
   -
ity through a so-called barcode analysis, where a DNA
 
a database of corresponding DNA sequences from well-
    
also of single unknown sequences.
The COI sequences of T. vittata from Myanmar,
       
  p-distance) to each other and to 78 published
COI sequences, corresponding to a clade Panijpan et
al. (2015) referred to as “Trichopsis sp. (cf. vittata) 3”
(BIN cluster ID BOLD:ACS9775), from western and
central Thailand, but dissimilar (> 2.4% p-distance) to all
individuals of T. vittata from other areas. This supports
the conclusion by Panijpan et al. (2015) that the current
T. vittata is a species complex, and indicates that T. vit-
tata  
introduced populations in Bangladesh and Myanmar all
originates from central and western Thailand. According
to Panijpan et al. (2015), “Trichopsis sp. (cf. vittata) 3” is
distributed in Thailand from Chiang Mai in the northwest,
to Trat in the southeast, and Surat Thani in the south.
Fowler (1934) described T. harrisi from Kratt (Trat), but
Kottelat (2001) considered T. harrisi a junior synonym of
T. vittata. A taxonomic revision of Trichopsis is needed,
and may come to the conclusion that the correct name for
“Trichopsis sp. (cf. vittata) 3” is T. harrisi.
The probable source of the feral population of T.
vittata    
also in the other locations the presumed source is release
       
and Balasubramanian 2015). However, while Thailand is
  T. vittata does
         
Pygmy gourami (T. pumila) is the most common spe-
cies of Trichopsis
populations have only been reported from the Philippines
(BFAR 2006), without data. This suggests either that T.
vittata is able to colonize new territory even when propa-
gule size and frequency are low, or that it spreads also
through some mechanism other than release of aquarium
  
mosquitos or as a contaminant among shipments of live

Wongsiri (1982) found that in Thailand the 3 most
  T. vit-
tata, Guppy (Poecilia reticulata Peters, 1859) and Tilapia
(Oreochromis sp.). The ability of T. vittata to survive in
very small volumes of water, and to tolerate polluted or
low-oxygen water, could make it suitable for mosquito
-
tionally released as a mosquito control agent.
Another possible avenue for introduction is as a con-

East Asian Topmouth Gudgeon, Pseudorasbora parva
(Temminck & Schlegel, 1846), was introduced to East-

cyprinids imported for aquaculture trials and has since
spread to many river systems in Europe (Gavriloaie et al.
2008). In Asia, T. vittata could spread as a contaminant in
shipments of for instance live Silver Barb, Barbonymus
gonionotus (Bleeker, 1849), native to Southeast Asia,
sympatric with T. vittata, and an important aquaculture
species in South and Southeast Asia (Gupta and Rab
1994), but both Myanmar and Bangladesh have a range
of cultured species (Siddiqui et al. 2007) which present
similar potential. Nevertheless, the records from India,
Myanmar and Bangladesh are close to major cities, where
aquarium releases are more likely, and in the absence of
evidence that T. vittata spreads as a contaminant, and con-
sidering that the feral populations are genetically similar
to specimens from the aquarium trade, we consider the
release of specimens from the aquarium trade the most
probable mechanism of introduction.
Trichopsis vittata has many of the traits typical of
         
oxygen conditions, is human-associated (through aquaria,


Norén et al. | First records of Trichopsis vittata from Myanmar and Bangladesh 85
has a short generation time, guards its eggs (uniparental
care), and has a prior history of successful invasions. The
preferred habitat of T. vittata (shallow, often temporary,

areas suitable for rice-production, suggesting that it could
potentially colonize most of southern and eastern Asia.
However, T. vittata does not have any mechanism for
long-distance dispersal, as it is non-migratory and does
not have planktonic eggs or larvae, and non-human medi-
ated spread is likely to be slow. In addition, it may locally
be limited by life-history variables or interactions with
the native biota, as seems to be the case with the popula-

   T.
vittata have been reported, but Knight and Balasubra-
manian (2015) speculated that in India it may compete
for niche space with native small osphronemids such as
Dwarf Gourami [Trichogaster lalius (Hamilton, 1822)]
   Pseudosphromenus cupanus
 
aggressive displacement of native species, or that it may
act as a vector for non-native parasites or pathogens, such
as the trematode Euclinostomum heterostomum (Rudol-
phi, 1809) (Purivirojkul and Sumontha 2013).
Acknowledgements
This study is part of the project “Genetic characterization

(Swedish Research Council, contract D0674001 to Sven
Kullander and Abdur Rob Mollah).
Authors’ Contributions
All authors contributed to collection and text. MN and
MR extracted and sequenced DNA. MN made the analy-
ses. SOK made images and table.
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