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Growth and morphological development of laboratory-reared larval and

juvenile snakeskin gourami Trichogaster pectoralis

Article  in  Ichthyological Research · January 2010

DOI: 10.1007/s10228-009-0118-x

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Ichthyol Res (2010) 57:24–31
DOI 10.1007/s10228-009-0118-x
FULL PAPER
Growth and morphological development of laboratory-reared
larval and juvenile snakeskin gourami Trichogaster pectoralis
Shinsuke Morioka Æ Sayaka Ito Æ Shoji Kitamura
Received: 6 November 2008 / Revised: 15 May 2009 / Accepted: 19 May 2009 / Published online: 30 June 2009
Ó The Ichthyological Society of Japan 2009
Abstract Morphological development, including that of
fins, labyrinth organ, body proportions, and pigmentation, in
laboratory-hatched larval and juvenile snakeskin gourami
Trichogaster pectoralis is described. Body lengths (BL;
mean ± SD) of larvae and juveniles were 2.3 ± 0.1 mm
just after hatching (day 0) and 8.2 ± 0.6 mm on day 22,
reaching 14.1 ± 2.3 mm on day 48. Aggregate fin ray
numbers attained their full complements in juveniles
[11.8 mm BL. Preflexion larvae started feeding on day 2
following upper and lower jaw formation, the yolk being
completely absorbed by day 12. Subsequently, oblong con-
ical teeth appeared in postflexion larvae [8.2 mm BL
(day 16). Melanophores on the body increased with growth,
with a large dark spot developing on the lateral midline at the
caudal margin of the body in flexion larvae [6.1 mm BL.
Subsequently, a broad vertical dark band from the eye to the
caudal peduncle developed in postflexion larvae [8.9 mm
BL. Proportions of head and pre-anal lengths became con-
stant in postflexion larvae greater than ca. 9–10 mm BL,
whereas those of maximum body depth, eye diameter, and
snout length failed to stabilize in fish of the size examined in
this study. First soft fin ray of the pelvic fin elongated,
reaching over 40% BL. The labyrinth organ differentiated in
postflexion larvae [7.4 mm BL (day 22). Comparisons of
larval and juvenile morphology with another anabantoid
species Anabas testudineus were also made, revealing sev-
eral distinct differences, particularly in the numbers of
myomeres and fin rays in the dorsal/anal fins, mouth location
and body shape.
S. Morioka (&)
S. Ito
S. Kitamura
Fisheries Division, Japan International Research Center for
Agricultural Sciences, 1-1 Owashi, Tsukuba, Ibaraki 305-8686,
Japan
e-mail: moriokas@affrc.go.jp; morishinlao@yahoo.co.jp
123
Keywords Trichogaster pectoralis
Larvae
Juveniles

Morphology
Introduction
The snakeskin gourami Trichogaster pectoralis, belonging
to Osphronemidae, suborder Anabantoidei, is one of the
most common air-breathing freshwater fishes in the Indo-
china peninsula (Burggren and Haswell 1979; Rainboth
1996; Kottelat 2001), its distribution having been increased
due to introductions to neighboring countries, e.g.,
Malaysia, Indonesia, the Philippines, and Papua New
Guinea, where breeding populations of the species are now
established (West and Glucksman 1976; Welcomme 1988;
Juliano et al. 1989). The species is often found in rice
paddies, ditches, and streams with dense vegetation
(Rainboth 1996; Vidthayanon 2002), and it is an important
aquaculture target in Thailand, Vietnam, and Malaysia
(Vromant et al. 2001).
In Laos, 10 species of Anabantoidei are known to occur
(9 species of Osphronemidae, in the genera Betta, Macro-
podus, Osphronemus, Trichogaster, and Trichopsis, and
1 species of Anabantidae, in Anabas) (Kottelat 2001).
These species are significant targets of commercial and
small-scale fisheries in the region (Iwata et al. 2003;
Welcomme and Vidthayanon 2003), although differences
in marketable values exist according to species and size. At
the same time, human populations have rapidly grown in
the region, and environmental changes, such as urbaniza-
tion, increasing land use for cropping, biological invasion
by alien fishes (e.g., Oreochromis spp., Hypophthalmich-
thys molitrix, Aristichthys nobilis) and hybrid fishes
(crossbred Clarias spp.) (Iwata et al. 2003; Welcomme and
Vidthayanon 2003; Na-Nakorn et al. 2004; Senanan et al.
Morphological development of Trichogaster pectoralis
2004) have occurred. Such impacts, disturbing regional fish
diversity, potentially lead to declines of indigenous fish
stocks, including Anabantoidei. Accordingly, the stock
assessment of these species is immediately required, and
species identification particularly in larval and juvenile
stages is important. Hence, accumulation of early life-his-
tory information is essential, including morphological
descriptions of indigenous species. Furthermore, since
Anabantoidei has long been a systematically contentious
taxonomic group, in view of species’ diversification and
evolutionary ecology (Rüber et al. 2006), descriptions of
early life-stage morphology should provide significant new
information, as only Anabas testudineus has thus been
treated so far (Morioka et al. 2009). The objective of the
present study, therefore, was to facilitate the identification
of early life stages of the snakeskin gourami Trichogaster
pectoralis. The morphology of larvae and juveniles are
described in detail from a series of laboratory-reared
specimens.
Materials and methods
Parental fishes and egg collection. Broodstocks of Tric-
hogaster pectoralis were collected from small water bodies
in the Namxuang area, 40 km north of Vientiane City,
Laos, on 2 July 2007. Broodstocks were reared in the
Living Aquatic Resources Research Center, Vientiane
City, Laos. A LH-RH-analogue hormone (Suprefact;
HOECHST AG, Germany) was injected in combination
with a dopamine inhibitor (Motilium; OLIC, Thailand) at
the rate of 4 mg/100 g FW (fish wet weight) for the former
and 2 mg/100 g FW for the latter, into one male (body
weight 70 g) and one female (90 g) at 1630–1700 hours on
4 July 2007, prior to placing them in a spawning tank
containing 100 l water. Because Trichogaster species are
known to form a bubble nest beneath floating aquatic plants
for spawning (Cole et al. 1999), two taro leaves (ca. 30 cm
diameter) were floated in the spawning tank. Water tem-
perature during the induced breeding period was 28.2–
29.2°C.
Rearing larvae and juveniles. Fertilized eggs were
obtained through the induced breeding process with the
newly hatched larvae being separated into six rectangular
aquaria (30 cm length 9 40 cm width 9 30 cm depth
containing 36 l of water each) and reared under ambient
water temperatures ranging from 26.8 to 30.2°C. From
day 2 after hatching, fish were fed cultured zooplankton
(Brachionus spp.) at a density of 10 individuals/ml in the
rearing tanks three times a day until day 16. From days 5 to
48 inclusive, Artemia nauplii as well as Brachionus spp.
were also added at a density of 2–3 individuals/ml three
times a day.
25
Observations and measurements. Fertilized eggs were
collected ca. 1 h after spawning, and larvae and juveniles
on days 0–3, 5, 7, 9, 11, 13, 16, 22, 29, 38, and 48 after
hatching. All were preserved in 5% formalin immediately
after sampling. Observations and measurements were
made in the Fisheries Division of Japan International
Research Center for Agricultural Sciences, Tsukuba,
Japan. The fertilized eggs and some fish samples were
registered in the Museum, Tokyo University of Marine
Science and Technology (MTUF). Measurements were
made on 7 eggs [MTUF-P(L)-16363] and 7–15 fish on
each day of sampling, for a total of 127 larvae (2.1–
12.0 mm in body length) and 19 juveniles (11.8–17.8 mm
in body length). Of these, 7 were dissected for observa-
tions of the labyrinth organ [MTUF-P(L)-16378–16384],
the remainder (n = 139) being used for observations on
general morphology, number of myomeres, pigmentation,
fin development, and the following morphometric mea-
surements (mm): body length (BL), head length (HL),
pre-anal length (PAL), maximum body depth (BD), eye
diameter (ED), snout length (SnL), length of first soft ray
of pelvic fin (PFRL). The HL, SnL, and PAL were
measured from day 1 after operculum formation and anus
opening, PFRL from its appearance on day 29, and the
remainder from day 0. Body length was taken as noto-
chord length before hypural formation, and standard
length thereafter. Some specimens were stained in alizarin
red S or alcian blue 8GX for observations of fin ray
formation and jaw development. Fourteen specimens were
sketched [MTUF-P(L)-16364–16377]. Egg diameter was
measured after spawning. Measurement methods followed
Leis and Trnski (1989).
Results
Spawning and hatching. Spawning took place approxi-
mately 14 h after hormone injection, at 0700 hours on
5 July 2007, and the spawned eggs were concentrated in the
bubble nest beneath the taro leaves. Approximately 37,500
fertilized eggs were obtained with a fertilization rate of
94.8%. Eggs were isolated, epipelagic and oval in shape,
with the egg diameter ranging from 0.89 to 1.03
(mean ± SD: 0.97 ± 0.06, n = 7) mm in maximum axis
and 0.70 to 0.80 (mean ± SD: 0.74 ± 0.04, n = 7) mm in
minimum axis. Hatching took place approximately 23 h
after spawning at 0600 hours on 6 July 2007.
Larvae and juveniles. General morphology. The body
length (BL, mm) of newly hatched larvae (day 0) ranged
from 2.1 to 2.3 (mean ± SD: 2.2 ± 0.1, n = 7) mm,
reaching 5.1 ± 0.2 mm on day 5 (n = 10), 7.0 ± 0.3 mm
on day 11 (n = 10), 8.2 ± 0.6 mm on day 22 (n = 10),
10.8 ± 1.4 mm on day 38 (n = 10), and 14.1 ± 2.3 mm
123
26
Fig. 1 Changes in body length (mm) of laboratory-reared larval and
juvenile snakeskin gourami Trichogaster pectoralis from days 0 to 48
after hatching. Vertical bars indicate standard deviations
Table 1 Body length (BL) and age at each developmental stage of
Trichogaster pectoralis
Stage BL (mm) Age (days) Number of fish
Preflexion larva 2.1–5.8 0–5 43
Flexion larva 5.8–7.4 5–16 48
Postflexion larva 7.2–12.0 16–48 36
Juvenile [11.8 [38 19
on day 48 (n = 15) (Fig. 1). Body lengths (mm) of larvae
and juveniles at each developmental stage are shown in
Table 1.
Newly hatched larvae (n = 7) presented with a large
oval yolksac (vertical axis 0.96 ± 0.06 mm, horizontal
axis 0.70 ± 0.05 mm); anterior margin of yolksac bor-
dering on ventral aspect of bent head (Fig. 2a); yolk split
into two portions in day 2 preflexion larvae (4.4 ± 0.1 mm
BL, n = 10) (Fig. 2c); yolk moving dorsally to above
abdominal cavity beside upper base of pectoral fin
(Fig. 2c), completely absorbed in flexion larvae by day 12
(7.0 ± 0.2 mm BL, n = 10) (Fig. 2h).
Head initially bent with ventral aspect bordering on
anterior margin of yolksac (Fig. 2a), subsequently stretch-
ing and separating from yolksac from day 1 (3.3 ± 0.1 mm
BL, n = 7) (Fig. 2b); operculum appearing on day 1
(Fig. 2b); lateral line appearing on day 2 (Fig. 2c). Myo-
meres fully observable (numbering 7–8 ? 25–26 = 32–34)
with yolk absorption in flexion larvae [5.8 mm BL (day 7;
Fig. 2f), subsequently becoming invisible in postflexion
larvae [8.5 mm BL (day 22; Fig. 2k). Gas bladder visible
between abdominal cavity and notochord in preflexion
larvae on day 3 (4.3 ± 0.2 mm BL, n = 10; Fig. 2d),
becoming invisible in postflexion larvae [7.2 mm BL on
day 16 (Fig. 2j).
Ventral finfold initially originating on anterior 50–53%
BL (Fig. 2a), divided into anterior and posterior portions
relative to anus position in day 1 larvae (Fig. 2b), anterior
123
S. Morioka et al.
portion disappearing in postflexion larvae [11.6 mm BL
(day 29; Fig. 2l), posterior portion in juveniles (day 38;
Fig. 2m); dorsal finfold initially originating on anterior 36–
40% of body (Fig. 2a), disappearing in postflexion larvae
[9.0 mm BL (day 29; Fig. 2l); caudal finfold initially
round, fan-shaped (Fig. 2a), a notch in upper half of finfold
appearing in day 5 preflexion larvae (5.1 ± 0.2 mm BL,
n = 10; Fig. 2e), caudal finfold disappearing in juveniles
(day 38; Fig. 2m). Pterygiophores of anal fin clearly visible
in flexion larvae [6.0 mm BL (Fig. 2g) and becoming
invisible in juveniles (day 38; Fig. 2m).
Mouth and anus opened in day 1 larvae (Fig. 2b), mouth
reaching a vertical through anterior ca. 40% of eye dia-
meter on day 1 (Fig. 2b), subsequently posterior edge of
mouth moving forward with growth, located in anterior
half of snout in juvenile stage (Fig. 2m, n); upper and
lower jaws formed in day 1 larvae (Fig. 2b) (upper jaw
length 3–4% BL on day 1, constant thereafter); a few
sharp, oblong conical teeth appearing on both jaws in
postflexion larvae [8.2 mm BL (day 16), teeth increasing
in number with growth. Nostril appearing in day 2 larvae
(Fig. 2c), dividing into two in juveniles [14.5 mm BL
(day 48; Fig. 2n).
Head length 17–18% BL in day 1 larvae (Fig. 2b),
proportion subsequently increasing with growth, reaching
ca. 30–35% BL in postflexion larvae[9.7 mm BL (day 29;
Fig. 2l). Pre-anal length 45–50% BL initially in day 1
larvae (Fig. 2b), proportion decreasing to 36–38% BL with
tail extension in preflexion larvae \5.6 mm BL (day 5;
Fig. 2e), subsequently increasing to 46–50% BL in post-
flexion larvae [8.5 mm BL (day 22; Fig. 2k). Maximum
body depth initially 34–38% BL (Fig. 2a), proportion
rapidly decreasing to 16–18% BL with yolk absorption and
body extension in preflexion larvae \5.6 mm BL (day 5;
Fig. 2e), subsequently increasing continuously to over 30%
BL in juveniles [11.8 mm BL (Fig. 2m). Eye diameter
initially 13–14% BL (Fig. 2a), proportion rapidly
decreasing to 8–9% with head and body extension in pre-
flexion larvae \5.6 mm BL (day 5; Fig. 2e), proportion
subsequently increasing to 12–15% BL in postflexion lar-
vae \9.7 mm BL (day 29; Fig. 2l), decreasing slightly
thereafter. Snout length 0.3–0.6% BL on day 1 (Fig. 2b),
increasing to 7–8% BL in postflexion larvae\10.6 mm BL
(day 29; Fig. 2l) and decreasing slightly thereafter. First
soft ray of pelvic fin elongating following initial appear-
ance, proportion attaining over 40% BL of body length on
day 48 (Fig. 2l–n).
Fin development. Dorsal soft rays appearing in flexion
larvae [6.8 mm BL (day 11–13; Figs. 2i, 3a), spines in
postflexion larvae [8.2 mm BL (day 16–22), attaining full
complement (XI–XII, 10) in postflexion larvae [10.6 mm
BL (day 38; Figs. 2l, 3a); soft ray segmentation initiated in
postflexion larvae [9.8 mm BL (day 29), completed (8–9
Morphological development of Trichogaster pectoralis
Fig. 2 Laboratory-reared larval and juvenile snakeskin gourami
Trichogaster pectoralis. a Newly hatched larva [2.2 mm BL,
MTUF-P(L)-16377]; b preflexion larva, day 1 [3.3 mm BL, MTUF-
P(L)-16376]; c preflexion larva, day 2 [4.1 mm BL, MTUF-P(L)-
16375]; d preflexion larva, day 3 [4.7 mm BL, MTUF-P(L)-16374];
e preflexion larva, day 5 [5.3 mm BL, MTUF-P(L)-16373]; f flexion
larva, day 7 [5.9 mm BL, MTUF-P(L)-16372]; g flexion larva, day 9
[6.1 mm BL, MTUF-P(L)-16371]; h flexion larva, day 11 [6.7 mm
segmented rays) in postflexion larvae [10.7 mm BL
(day 38). Anal soft rays appearing in flexion larvae [6.5
mm BL (day 11–13; Figs. 2h, 3b), spines in postflexion
larvae[8.1 mm BL (day 16–22), attaining full complement
(VII–VIII, 38–40) in postflexion larvae [9.8 mm BL
(day 38; Figs. 2m, 3b); soft ray segmentation initiated in
postflexion larvae [8.0 mm BL (day 29), completed
(35–38 segmented rays) in juveniles (day 48). Caudal soft
rays appearing in flexion larvae [5.8 mm BL (day 7;
Figs. 2f, 3c), attaining full complement (9 ? 9 = 18) in
postflexion larvae [8.9 mm BL (day 22; Figs. 2k, 3c); soft
ray segmentation initiated in flexion larvae [6.5 mm BL
(day 11), completed (16 segmented rays) in postflexion
larvae [8.9 mm BL (day 22). Pectoral soft rays appearing
in postflexion larvae [8.2 mm BL (day 16 to 22; Figs. 2k,
3d), attaining full complement (10–11) in postflexion larvae
[9.1 mm BL (day 29; Figs. 2l, 3d); soft ray segmentation
initiated in postflexion larvae [8.7 mm BL (day 29),
27
BL, MTUF-P(L)-16370]; i flexion larva, day 13 [6.9 mm BL, MTUF-
P(L)-16369]; j postflexion larva, day 16 [7.3 mm BL, MTUF-P(L)-
16368]; k postflexion larva, day 22 [8.9 mm BL, MTUF-P(L)-16367];
l postflexion larva, day 29 [11.6 mm BL, MTUF-P(L)-16366];
m juvenile, day 38 [13.2 mm BL, MTUF-P(L)-16365]; n juvenile,
day 48 [17.8 mm BL, MTUF-P(L)-16364]. b0 –d0 Pre-anal dorsal view,
k0 –n0 ventral view of pelvic fin. YS Yolksac, GB gas bladder
completed (8–9 segmented rays) in juveniles [11.8 mm
BL (day 26). Pelvic soft rays appearing in postflexion
larvae [8.9 mm BL (day 29; Figs. 2l, 3e), a spine appear-
ing in postflexion larvae [11.6 mm BL (day 29; Fig. 2l),
attaining full complement (I, 4) in juveniles [11.8 mm
BL (day 38; Figs. 2m, 3e); first soft ray becoming
remarkably elongate following its appearance at 8.9 mm
BL (day 29; Fig. 2l–n); first soft ray segmentation initiating
in postflexion larvae [11.6 mm BL (day 29), while other
soft rays not segmented. Branched rays not present in any
fins.
Pigmentation. Melanophore deposition in eyes observed
in day 1 larvae (Fig. 2b). Many light stellate melanophores
initially present over entire surface of yolksac, decreasing
in number with yolk absorption (Fig. 2a–h). Two small
punctate melanophores present on snout of day 1 larvae
(Fig. 2b), disappearing in day 2 larvae (Fig. 2c), subse-
quently two or three punctate melanophores appearing on
123
28
snout and interorbital section in day 3 larvae (Fig. 2d),
increasing in number with growth; several punctate mela-
nophores appearing on upper jaw in flexion larvae at
5.8 mm BL (day 7; Fig. 2f), on lower jaw in flexion larvae
at 6.1 mm BL (day 9; Fig. 2g), increasing in number with
growth (Fig. 2f–n). Several stellate melanophores initially
scattered on head (Fig. 2a), becoming punctate and
increasing in number in day 1 larvae (Fig. 2b), subse-
quently decreasing in number in preflexion larvae by
5.8 mm BL (day 5; Fig. 2e), increasing thereafter with
growth (Fig. 2f–n).
Several punctate melanophores present on throat and
ventral region of abdomen in day 2 larvae (Fig. 2c),
decreasing and completely lost in postflexion larvae at
11.6 mm BL (day 29; Fig. 2l). Two or three stellate mela-
nophores appearing on upper region of operculum in day 2
larvae (Fig. 2c), increasing and becoming punctate with fish
growth (day 13); several stellate and punctate melano-
phores appearing on circumorbital section just behind eye in
postflexion larvae at 8.9 mm BL (day 22; Fig. 2k),
increasing and extending to the melanophore assembly on
operculum in juveniles at 17.8 mm BL (day 48; Fig. 2n).
Dozens of small punctate melanophores present on dor-
sal body along dorsal finfold except posterior tip in day 1
larvae, covering entire dorsal region of body in flexion
larvae [6.9 mm BL from day 13 (Fig. 2i–n). Small punc-
tate melanophores present on ventral body along ventral
finfold except posterior tip in day 1 larvae (Fig. 2b), sub-
sequently pterygiophores of anal fin with melanophores
developing distally in flexion larvae [6.0 mm BL (day 9;
Fig. 2g); melanophores posterior to anus constant until
early juveniles [13.3 mm BL along base of pterygio-
phores (Fig. 2m), subsequently increasing and covering
posterior ca. 70–80% of ventral margin along anal fin
(Fig. 2n).
Melanophores on lateral body initially few (Fig. 2a, b),
subsequently over a dozen punctate melanophores
appearing on lateral line since its appearance in day 2
larvae (Fig. 2c), increasing with growth, but disappearing
in postflexion larvae at 11.6 mm BL (day 29; Fig. 2l);
several stellate melanophores sparsely distributed on lateral
body in flexion larvae at 6.1 mm BL (day 9; Fig. 2g) in
addition to those on lateral line, subsequently becoming
punctate and covering most of the upper lateral body in
postflexion larvae at 8.9 mm BL (day 22; Fig. 2k); many
punctate and stellate melanophores appearing beneath lat-
eral line in postflexion larvae at 8.9 mm BL (day 22;
Fig. 2k), extending to those on operculum with growth,
thereby forming a vertical broad dark band in juvenile
stage (Fig. 2m), thereafter relatively less dense melano-
phores form five light horizontal bands on the posterolat-
eral half of the body in juveniles at 17.8 mm BL (day 48;
Fig. 2n); dozens of punctate melanophores appearing
123
S. Morioka et al.
around caudal margin of body in flexion larvae at 6.1 mm
BL (day 9; Fig. 2g), becoming more dense and forming a
large dark spot with growth (Fig. 2g–n).
Small dense melanophores present on the dorsal surface
of gas bladder from its initial appearance in day 3 larvae
(Fig. 2d), disappearing as gas bladder becomes invisible in
postflexion larvae at 7.3 mm BL (day 16; Fig. 2j); mela-
nophores appearing internally on ventral side of notochord
(Fig. 2g), reaching caudal peduncle in flexion larvae at
7.3 mm BL (day 16; Fig. 2j), becoming invisible in post-
flexion larvae at 8.9 mm BL (day 22; Fig. 2k).
Several punctate melanophores appearing on caudal fin
in flexion larvae at 5.9 mm BL (day 7; Fig. 2f), increasing
thereafter; several punctate melanophores appearing on
dorsal and anal fin in flexion larvae at 6.9 mm BL (day 13;
Fig. 2i), those on lower half of anal fin becoming more
dense in postflexion larvae at 11.6 mm BL (day 29;
Fig. 2m), subsequently increasing to densely cover entire
anal fin in larger juveniles (Fig. 2n).
Labyrinth organ development. Labyrinth organ appear-
ing in postflexion larvae [7.4 mm BL (day 22) as pri-
mordial hypertrophy of upper portion of first gill arch
(Fig. 4b), hypertrophy subsequently developing (Fig. 4c–e)
and forming a spherical structure on that portion of the gill
arch in juveniles at 12.6 mm BL (day 38; Fig. 4f) and a
fan-shaped membranous structure in juveniles at 16.4 mm
BL (day 48; Fig. 4g).
Discussion
The developmental stages of Trichogaster pectoralis were
determined in this study, with the species considered to
have attained the juvenile stage by 11.8 mm BL (based on
completion of aggregate fin ray numbers; Table 1;
Figs. 2m, 3). However, body proportions did not become
constant within the size range of fish used in the study
(max. 17.8 mm BL).
The present results on egg diameters and duration from
spawning to hatching more or less coincided with ones
reported by Amornsakun et al. (2004), although the num-
bers of spawned eggs and the yolksac period were both
greater in the present study, despite both studies utilizing
more or less corresponding broodstock sizes under similar
water temperatures. The difference in egg number, i.e.,
over 400,000 eggs/kg female in the present study and ca.
280,000 eggs/kg female in Amornsakun et al. (2004), may
be explainable by differences in maturation levels of the
broodstocks. However, the causative factor for different
yolksac periods, i.e., 11 days in the former and 4.5 days in
the latter, remains unclear, although small, well-absorbed
yolks in the final phase of the yolksac period (Fig. 2g, h)
may have been overlooked by Amornsakun et al. (2004).
Morphological development of Trichogaster pectoralis 29

The 11-day yolksac period observed in the present study

(Fig. 3) indicated that Trichogaster pectoralis has a ca.
10-day preparatory period from days 2 (start of feeding) to
11 (just before complete yolk absorption) inclusive, for the
shift from endogenous to exogenous energy-dependent
periods (under the experimental temperature regime during
this study). The shift from endogenous to exogenous
energy-dependent periods has been examined in several
species of marine fishes, covering aspects of their devel-
opment and survival during the larval stages (Kohno 1998;
Moteki et al. 2001). However, similar investigations of
freshwater fishes, including T. pectoralis, are still scarce,
except for the climbing perch Anabas testudineus (Morioka
et al. 2009) and the North African catfish Clarias gariepinus
(Matsumoto et al. 2001), despite being necessary for con-
siderations of evolutionary ecology, including survival
strategies during early life stages, as well as for achieving
any improvements in seed productivity. By comparison,
A. testudineus (Anabantidae), also included in suborder
Anabantoidei, showed a ca. 5-day preparatory period from
days 2 to 6 inclusive (Morioka et al. 2009) under similar
water temperatures, a period much shorter than that of
T. pectoralis. In addition, the duration from spawning to
hatching in A. testudineus (ca. 11 h, Morioka et al. 2009)
was also much shorter than in T. pectoralis (ca. 23 h)
observed in the present study. These differences may be
attributable to the differences in reproductive strategies of
the two species, i.e., dispersal spawning without nesting and
parental care in A. testudineus, compared with spawning in
a bubble nest with parental care in T. pectoralis (Rüber et al.
2006) as well as its congener T. trichopterus (Kramer 1973),
although confirmation of this is required.
Although the function of the labyrinth organ and asso-
ciated ecological aspects (Burggren and Haswell 1979)
have been reported for adult T. pectoralis and its congener
T. trichopterus (Burggren 1979), ontogeny and develop-
mental morphology of the former in larval and juvenile
stages are poorly known. In the air-breathing species
Macropodus opercularis of Osphronemidae (Matayoshi and
Shokita 1982) and A. testudineus of Anabatidae (Morioka
et al. 2009), descriptions of the developmental morphology
of the labyrinth organs showed that the occurrence of air-
breathing followed the onset of primordial hypertrophy of
the first gill arch and continued with subsequent enlarge-
ment of the organ. These observations suggest that air-
breathing in T. pectoralis also starts in the postflexion larval
stage at the onset of primordial hypertrophy of the first gill Fig. 3 Relationships between body length (mm) and fin ray numbers
arch ([7.4 mm BL; Fig. 4). Since T. pectoralis is an obli- in laboratory-reared larval and juvenile snakeskin gourami Tricho-
gaster pectoralis. a Dorsal fin, b anal fin, c caudal fin, d pectoral fin,
gate air-breather (Burggren and Haswell 1979), it is highly
e pelvic fin
resistant to low levels of dissolved oxygen in the sur-
rounding water, suggesting a considerable aptitude of the of T. pectoralis was observed to possess only a single-lay-
species for seed production and aquaculture under high ered membrane in the specimen of 16.4 mm BL (Fig. 4),
stocking densities. In the present study, the labyrinth organ whereas it was more developed at 14.9 mm BL having

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Fig. 4 Development of labyrinth organ in laboratory-reared larval
and juvenile snakeskin gourami Trichogaster pectoralis. a Flexion
larva, day 16 [7.1 mm BL, MTUF-P(L)-16384]; b postflexion larva,
day 22 [7.6 mm BL, MTUF-P(L)-16383]; c postflexion larva, day 29
[8.4 mm BL, MTUF-P(L)-16382]; d postflexion larva, day 29
three-layered membranes in A. testudineus (Morioka et al.
2009). This difference as well as the faster morphological
development in A. testudineus (Morioka et al. 2009) may
reveal their specific survival strategies during early life
stages, although more information is necessary.
Although nine species of Osphronemidae, including
T. pectoralis, are known to be distributed in Laos (Kottelat
2001), their early life-stage morphology has not been
described to date. However, the morphology of larval and
juvenile Anabas testudineus was reported in detail by
Morioka et al. (2009). Although newly hatched larvae of
both species have similarities in body size, head shape
(bent and bordering the anterior yolksac margin), and body
shape (large yolksac with stellate melanophores), T. pec-
toralis had a greater number of melanophores on the
yolksac surface. On the other hand, a number of small
punctate melanophores were present on the dorsal and
ventral aspects of the body in newly hatched larvae of
A. testudineus, being scarce in T. pectoralis. During the
preflexion and flexion larval stages, the major similarities
between the two species are the divided yolksac forming
two portions located behind the upper base of the pectoral
fin and, the differences were found in the number of
myomeres, particularly those posterior to the anus (25–26
in T. pectoralis and 19–21 in A. testudineus), and the
smaller PAL as a percentage of BL in T. pectoralis. In
addition, during those stages, the posterior end of the
mouth was beneath the eye at mouth opening, subsequently
moving forward to level with the middle of the snout in
T. pectoralis, and to the anterior 30% of the eye in
A. testudineus. Following the postflexion larval stage, the
identification of the two species is made easier by the
development of several distinct differences, as follows:
(1) more slender body shape with smaller head in T. pec-
toralis than A. testudineus, (2) dorsal and anal fin ray
numbers of D XI–XII, 10, A VII–VIII, 38–40 in the former
123
S. Morioka et al.
[9.6 mm BL, MTUF-P(L)-16381]; e postflexion larva, day 38
[10.7 mm BL, MTUF-P(L)-16380]; f juvenile, day 38 [12.6 mm
BL, MTUF-P(L)-16379]; g juvenile, day 48 [16.4 mm BL, MTUF-
P(L)-16378]. LO Labyrinth organ, MS membranous structure(s), GR
gill rakers. Horizontal bar 1 mm
and D XVI–XVII, 10–11, A IX–X, 10–11 in the latter,
(3) melanophore distributions on the lateral body surface
forming a broad vertical band in the former and several
horizontal bands in the latter, and (4) first soft ray of pelvic
fin elongated in the former, but not in the latter. In addition,
T. pectoralis is zooplanktivorous by the early juvenile stage
and subsequently omnivorous (Boonsom 1984), whereas
A. testudineus is highly piscivorous, even in the larval
stages smaller than 8 mm BL (Morioka et al. 2009).
Acknowledgments We express our sincere gratitude to K. Sano,
Laboratory of Global Fisheries Science, University of Tokyo, and
staff of the Aquaculture Improvement and Extension Project of Japan
International Cooperation Agency for their kind cooperation on
broodstock collection. We are also grateful to L. Khamsivilay and
B. Vongvichith, Living Aquatic Resources Research Center, Laos, for
their logistical support. Our thanks also go to G. Hardy for his polite
and constructive English revision.
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