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Determining the level of parental care relating fanning behavior of five species of clownfishes in captivity

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  • Ramakrishna Mission Vivekananda Educational Research Institute

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Clownfishes Amphiprion sebae, A. clarkii, A. percula, A. ocellaris and Premnas biaculeatus were reared in captive condition and their parental care activities were studied. Role of both male and female in parental care and terminating in successful hatching of eggs studied is described. Site selection, clutch formation, spawning, guarding, mouthing and fanning of the eggs were quantified. A. sebae exhibits more effective parental care for longer durations. The time spent on fanning was longer in A. sebae and shorter in A. clarkii. Fanning frequency was higher in A. percula but lower in A. clarkii. In the five species the fanning duration and frequency were positively correlated to hatching success. Gradation of dissolved oxygen levels adjacent to the clutch area due to the fanning behavior is discussed. Relation between body weight and the fecundity, fanning duration and frequency with hatching success are documented.
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Indian Journal of Geo-Marine Sciences
Vol. 41 (5), October 2012, pp. 430-441
Determining the level of parental care relating fanning behavior of five species of
clownfishes in captivity
Swagat Ghosh, T. T. Ajith Kumar* & T. Balasubramanian
Centre of Advanced Study in Marine Biology, Faculty of Marine Sciences, Annamalai University,
Parangipettai - 608 502, Tamilnadu, India
*E-mail: tt_ajith@yahoo.co.in
Received 25 March 2011; revised 4 August 2011
Clownfishes Amphiprion sebae, A. clarkii, A. percula, A. ocellaris and Premnas biaculeatus were reared in captive
condition and their parental care activities were studied. Role of both male and female in parental care and terminating in
successful hatching of eggs studied is described. Site selection, clutch formation, spawning, guarding, mouthing and fanning
of the eggs were quantified. A. sebae exhibits more effective parental care for longer durations. The time spent on fanning
was longer in A. sebae and shorter in A. clarkii. Fanning frequency was higher in A. percula but lower in A. clarkii. In the
five species the fanning duration and frequency were positively correlated to hatching success. Gradation of dissolved
oxygen levels adjacent to the clutch area due to the fanning behavior is discussed. Relation between body weight and the
fecundity, fanning duration and frequency with hatching success are documented.
Key wards: Clownfish, Parental care, Spawning, Fanning, Hatching success
Introduction
With smaller burly size and brilliant colors,
the ornamental clownfishes are in great demand.
Hence breeding of marine clownfishes in captivity
may to ensure their conservation in nature and fetch
additional income to coastal folk. In the aquarium
trade, the clownfish are predominant and have
precious value. Clownfish exhibit protoandrous
hermaphrodism, which is denotes first male and the
most common mode of their reproduction is egg
deposition on a secure substrate, usually in close
proximity to the anemone, with a distinct type of
clutching behavior. Eggs are termed as demersal and
result in relatively larger fry. According to many
studies, the effort a male expends on parental care is
positively correlated with the certainty of paternity1-3,
but this is not always the case4,5. In clownfish males
most often provide care for the eggs, when
care at all is provided6. In the clownfish the male
cares for the eggs by fanning and guarding them
until they hatch, which takes about 6 to 10 days.
Instinct parental care to protect their eggs and
young ones during the vulnerable periods is of
great value. Aim of the present study is to document
the parental care exhibited by five different species
of clownfish to understand their behavior during the
incubation period.
Materials and Methods
Description of the clownfishes
Among the different marine ornamental fishes,
clownfishes are ranked as one of the most popular
attractions all over the world. Their small size,
hardness, pretty colour features, high adaptability
to live in captivity, acceptability of artificial diet,
their fascinating display behavior and symbiotic
relationship with sea anemones are the important
features. The clownfishes belong to the subfamily,
Amphiprioninae and the family, Pomacentridae;
30 species are recorded, of which one of the selected
clownfish belong to the genus Premnas and others
to namely belong to Amphiprion sebae, A. clarkii,
A. percula, A. ocellaris.
Sebae clownfish [Amphiprion sebae (Bleeker, 1853)]
Size of the fish, Male, 7.5 and Female, 11.5 cm
(Fig. 1a)
Dark brown to blackish with two white bars, the
mid body bar starting slight backwards and extending
on to rear part of dorsal fin. Snout, breast and belly
often yellow orange, tail yellow.
Clark’s clownfish [Amphiprion clarkii (Benneff, 1830)]
Size of the fish, Male, 7.0 cm and Female, 9.0 cm
(Fig. 1b)
GHOSH et al.: DETERMINING THE LEVEL OF PARENTAL CARE RELATING FANNING BEHAVIOR OF FIVE SPECIES
431
Usually black with variable amount of orange on
head, ventral parts and fins, three milky white bars on
head, body, base of caudal fin, transition between
darker body and bar across caudal fin base usually
abrupt, caudal fin whitish yellow edges but sometime
yellow.
Percula clownfish [Amphiprion percula (Lacepede,
1802)]
Size of the fish, Male, 6.4 cm and Female, 7.5 cm
(Fig. 1c)
Bright orange with three white bars, the middle
with forward projecting bulge, bars often outlines
with black that varies in width. Although A. percula’s
vibrant colours are eye catching, it is easily confused
with A. ocellaris (false clownfish). However, one can
distinguish the two by counting the number of dorsal-
fin spines. The A. percula usually has 10 dorsal-fin
spines, while A. ocellaris usually has 11 and also, the
latter never has thick black margins outlining the fins.
False clownfish [Amphiprion ocellaris (Cuvier, 1830)]
Size of the fish, Male, 6.5 cm and Female, 8.0 cm
(Fig.1 d)
Normally bright in orange colour with three white
bars, the middle one with forward projecting bulge,
bars have narrow black margins or disappeared. It is
really identical, but never a thick black margin around
the white bars.
Maroon clownfish [Premnas biaculeatus (Bloch, 1790)]
Size of the fish, Male, 7.5 cm and Female, 10.5 cm
(Fig. 1e)
Bright red to brownish-red in colour with three
relatively narrow white or grey bars, all fins same
colour as body, usually with a long spine in both
opercula. Male is brighter red than female with
brilliant white stripes and the female bars generally
grey, but can be switched rapidly to white if fish is
motivated.
Species acclimatization in captive environment
The study was carried from June 2008 to February
2009 in Faculty of Marine Sciences marine
ornamental hatchery. Six numbers in each of the
selected species of the clownfishes and three numbers
in each species of the anemones (Stichodactyla
mertensii and Heteractics gigantica) were procured
from the fish suppliers from Kolathur, Chennai.
They were transported to the hatchery and
accommodated in separate conditioning tanks.
After the pair formation, the potential couple of
each species was shifted to separate spawning
tanks (1000 l capacity) along with the host anemone.
The tank was provided with live rock, dead corals
and shells to mimic those of the natural habitat.
The fishes were fed daily at 9:00-9:30 a.m., 1:30-2:00
and 6:00-6:30 p.m. with boiled oyster and prawn
meat and anemones fed with prawn meat daily ones.
After two months, fishes were started spawning.
Behavioral observation
Behavior of the fishes to clean the substratum, lay
eggs, guard the eggs was recorded and hatch out was
documented. Every fanning bout was taken to start,
when the male’s pectoral fin began to ventilate the
egg clutch, and when the male stopped fanning and
began to swimming around. The fanning duration was
estimated since laying till hatching of fry. Fanning
duration (minutes), frequency (times/hour) and
hatching success (%) of each clutch of each species
were obtained by using stop watch and the data were
recorded from continues observation during the
morning, afternoon, evening and dusk periods in a
day before feeding. The means of fanning duration,
frequency and hatching success were calculated for
each species. Hatching success was estimated by
dividing the number of hatchling from the initial
number of eggs in a clutch. Effect of dissolve oxygen
and its changes in the vicinity area of the eggs was
estimated in each species by using D.O. meter
(Singapore ECOSAN).
Results
Fanning time
Before making the clutch, the parents of A. sebae
cleared the so called clutch area by removing algae on
the substratum. Generally, the shape of the clutch area
was round or oval. Eggs were deposited in a clutch
and attached to the substratum with sticky fluid.
Clutch diameter ranged from 4.5 cm to 8.5 cm and the
number which increased with size of the female.
Fecundity ranged between 600 and 1500 eggs and the
male care of the eggs by ventilating continuously by
pectoral fins; the female also took the responsibility
but occasionally.
The longest and shortest duration of fanning by the
male were 1.43 minutes and 0.43 minutes (one way
ANOVA F 8,39= 3.31, p<0.05) (Figure 2a). In the case
of female, maximum and minimum fanning duration
were 0.92 minutes and 0.03 minutes respectively
(one way ANOVA F 8,39= 13.81, p= NS) (Figure 2b).
INDIAN J. MAR. SCI., VOL. 41, NO. 5, OCTOBER 2012
432
In the case of A. clarkii, the clutch diameter
varied between 3.5 - 4.5 cm and the fecundity ranged
500-800 eggs in each spawning. Its fecundity was
lesser than A. sebae but more than that of A. percula
and A. ocellaris. The maximum and minimum fanning
durations by the male were 1.12 minutes and 0.39
minutes respectively (one way ANOVA F 9, 28= 23.43,
p= NS) (Figure 3a) and in the case of the female,
it was 0.7 minutes and it was also noticed that in
few occasion there were no fanning for an hour
(one way ANOVA F 9,28= 3.94, p<0.05) (Figure 3b).
In A. percula, the parents made a clutch which was
oval in shape and the eggs were deposited and
attached to the substratum with the sticky end. The
clutch area diameter varied from 3.5-5.0 cm and the
fecundity ranged between 300-500 eggs. The male
was fanning the eggs continuously but the female did
it occasionally. The maximum and minimum fanning
duration by the male were 1.62 minutes and 0.37
minutes respectively (one way ANOVA F 8,40= 8.69,
p=NS) (Figure 4a) and in the case of female, the
maximum and minimum fanning duration in total
Fig. 1—Different species of clownfish with deposited eggs a. Amphiprion sebae, b. A. clarkii, A. percula, A. ocellaris and
e. Premnas biaculeatus.
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433
Fig. 2—Fanning duration (Sep. - Feb.) in A. sebae (2a) male and (2b) female (Error bar - SE, - SD and - mean).
Fig. 3—Fanning duration (Sep. - Feb.) in A. clarkii (3a) male and (3b) female (Error bar - SE, - SD and - mean).
fanning period were 0.88 minutes and 0.06 minutes
respectively (one way ANOVA F 8,40= 26.20, p= NS)
(Figure 4b).
In A. ocellaris, the parents made the clutch
diameter varied from 3.5-4.5 cm. Fecundity of
this fish ranged between 200-500 eggs. Here,
both the male and female parents fanned vigorously.
Male was involved in mouthing the clutch
and removing the unfertilized and dead eggs. The
maximum and minimum fanning done by the
male were 1.67 minutes and 0.37 minutes (one way
ANOVA F 8,39= 3.47, p<0.05) (Figure 5a) and that
of the female were 0.92 minutes and 0.05 minutes
respectively (one way ANOVA F 8,39= 6.00, p<0.05)
(Figure 5b).
In P. biaculeatus, the clutch area diameter was
4-5 cm. Fecundity of this fish ranged from
300-600 eggs. Mostly, the male was caring the
eggs and the female spent maximum time with the
sea anemone. The maximum and minimum fanning
made by the male were 1.67 minutes and 0.39 minutes
respectively (one way ANOVA F 9,35= 1.88, p= NS)
(Figure 6a). In the case of the female, maximum
and minimum fanning duration were 0.92 minutes
and 0.01 minutes respectively (one way ANOVA
F 9,35= 37.16, p= NS) (Figure 6b).
Fishes of the genera, Amphiprion and Premnas
have obligate associations with anemones and lay
their eggs on the hardy substratum under the shelter of
the anemone’s stinging tentacles (Wilkerson, 1998).
INDIAN J. MAR. SCI., VOL. 41, NO. 5, OCTOBER 2012
434
Fig. 4—Fanning duration (Sep. - Feb.) in A. percula (4a) male and (4b) female (Error bar - SE, - SD and - mean).
Fig. 5—Fanning duration (Sep. - Feb.) in A. ocellaris (5a) male and (5b) female (Error bar - SE, - SD and - mean).
Fig. 6—Fanning duration (Sep. - Feb.) in Premnas biaculeatus (6a) male and (6b) female (Error bar - SE, - SD and - mean).
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435
Fig. 7—Average fanning duration monthly variation (Sep. - Feb.) in five clown fishes (7a) male and (7b) female (Error bar - SE, - SD
and - mean).
Fig. 8—Comparing fanning duration with incubation period (stages days wise) in five clown fishes (8a) male and (8b) female.
Similar observation was recorded in all our five
species. The average fanning duration of male
anemone fishes viz., A. sebae, A. clarkii,
A. percula, A. ocellaris and P. biaculeatus have
mean S.D.) viz. 1.32 (±0.13), 0.65 (±0.34),
1.10 (±0.31), 1.20 (±0.28), 1.19 (±0.32) minutes
(one way ANOVA F 4,29= 4.70, p<0.05) (Figure 7a)
and in the case of female were the mean (± S.D.)
0.80 (±0.22), 0.30 (±0.29), 0.65 (±0.19), 0.62
(±0.28) and 0.56 (±0.31) minutes respectively
(one way ANOVA F 4,29= 2.93, p<0.05) (Figure 7b).
In the present study, fanning duration also illustrated
with the incubation period, which is refers to their
everyday fanning ability on the clutch. In case of
male, fanning on first three were more compare
to fourth to sixth days and again increased fanning
on the days of seventh to up to hatching days
(Figure 8a). In case of female, it has linear relations
and gradually increased from the first day to till
before hatching (Figure 8b).
Trends in the fanning mechanism have revealed
that the parents are adjusting their fanning timing in
response to oxygen availability. In the present study,
it was observed that A. sebae, A. percula and
A. ocellaris bred throughout the six months period
at regular intervals and fanned their eggs normally.
In the case of A. clarkii, it did not spawn for
two months period and P. biaculeatus did not breed
INDIAN J. MAR. SCI., VOL. 41, NO. 5, OCTOBER 2012
436
for one month. The average fanning duration of
the male fishes during the study period (two way
ANOVA F 5,29=1.21, p= NS and F 4,29=3.68, p<0.05)
were mean (±S.D.) 1.32 (±0.13) minutes in A. sebae,
and in A. clarkii 0.54 (±0.49) minutes.
Generally A. clarkii spent maximum time spent
along with sea anemone and showing less
parental care and hatching duration also more
compare to A. sebae. In A. percula, A. ocellaris and
P. biaculeatus mean S.D.) fanning times were
1.10 (±0.31), 1.20 (±0.28), 1.07 (±0.57) minutes
respectively. In the female fish of A. sebae, A. clarkii,
A. percula, A. ocellaris and P. biaculeatus, the
mean (±S.D.) fanning times were 0.80 (±0.22), 0.30
(±0.29), 0.65 (±0.19), 0.62 (±0.28) and 0.56 (±0.31)
minutes respectively (two way ANOVA F 5, 29 =1.75,
p= NS and F 4,29=3.37, p<0.05).
Fanning frequency
In the present study, the fanning frequency
indicating cares taking capacity of parents and
the development of eggs for successful hatching. The
average fanning frequency of male clownfishes viz.
A. sebae, A. clarkii, A. percula, A. ocellaris and
P. biaculeatus was mean (±S.D.) 40.18 (±5.45),
42.68 (±3.48), 35.75 (±6.33), 42.00 (±4.37), 39.02
(±4.47) / hrs (one way ANOVA F 4,29= 2.70, p<0.05)
(Figure 9a) and in the case of females, the average
fanning frequency was mean (±S.D.) 5.33 (±1.57),
6.85 (±0.55), 5.23 (±4.09), 5.5 (±0.72) and 5.68
(±2.84) / hrs respectively (one way ANOVA F 4,29=
0.47, p<0.05) (Figure 9b). Present finding also
revealed that fanning frequency with the incubation
period, which is refers to their everyday number of
fanning time on the clutch/hr. In case of male,
frequency are first three days more then gradually
decreased fourth to sixth days and again increased
fanning on the days of seventh to up to hatching days
(Figure 10a). In case of female, fanning frequency
gradually increased from the first day to before
hatching (Figure 10b). Among the five species,
A. percula and A. ocellaris took more care of their
eggs and A. clarkii showed lesser care than others.
In the present study, fanning duration, fanning
frequency and hatching success varied with varying
species (Table 1 and 2).
Hatching success
The fanning time and frequency correlated well
with the hatching success of individuals, revealing the
fact that the hatching of eggs depends on the fanning
and mouthing of the parent clownfish. The average
hatching success during the study period varied with
varying species viz. A. sebae, A. clarkii, A. percula,
A. ocellaris and P. biaculeatus were mean S.D.)
87.18 (±6.83), 46.64 (±36.45), 84.34 (±5.35), 87.47
(±11.34), 87.47 31.92) % (one way ANOVA F 4,29=
3.84, p<0.01) respectively (Figure 11).
Effect of Dissolved oxygen
Present finding also sustaining that the differences
between pre-fanning dissolved oxygen and post-
fanning dissolved oxygen variation were measureed,
which signifying cares captivating ability of parents
and the development of eggs for successful hatching.
The average increased dissolved oxygen lavel in
Fig. 9—Comparing fanning frequency in five clown fishes (9a) male and (9b) female (Error bar - SE, - SD and - mean).
GHOSH et al.: DETERMINING THE LEVEL OF PARENTAL CARE RELATING FANNING BEHAVIOR OF FIVE SPECIES
437
Fig. 10—Comparing fanning frequency with incubation period (stages days wise) in five clown fishes (10a) male and (10b) female.
Table 1—Variable measured in male and female with this five
anemone fishes fanning paternity
Degree of paternity (fanning)
Sl.No.
Species Duration
(min.) Frequency
(time/h) Hatching
success (%)
1. A.sebae
Male 1.3 (± 0.13)
40.2 (± 5.45) 87.2 (± 6.83)
Female 0.8 (± 0.22)
5.3 (± 1.57)
2. A.clarkii
Male 0.5 (± 0.34)
35.8 (± 6.33) 46.6 (± 36.45)
Female 0.3 (±0.29) 5.2 (± 4.09)
3. A.percula
Male 1.1 (± 0.31)
42.7 (± 3.48) 84.3 (± 5.35)
Female 0.7 (± 0.19)
6.9 (± 0.55)
4. A.ocellaris
Male 1.2 (± 0.28)
42.0 (± 4.37) 87.5 (± 11.34)
Female 0.6 (± 0.28)
5.5 (± 0.72)
5. P.biaculeatus
Male 1.1 (± 0.32)
39.0 (± 4.47) 87.5 (± 31.92)
Female 0.6 (± 0.31)
5.7 (± 2.84)
All values given are means (± SD) from each pair in all five
cultured anemone fishes.
clownfishes tanks viz. A. sebae, A. clarkii, A. percula,
A. ocellaris and P. biaculeatus were mean (±S.D.)
1.71 (±0.05), 0.46 (±0.06), 1.04 (±6.33), 0.96 (±0.07),
0.68 (±0.38) mg/lit (one way ANOVA F 4,29= 7.02,
p<0.05) (Figure 12). The dissolved oxygen depends
on days wise developmental stages involved fanning
in selective male species of male clownfishes viz.
A. sebae, A. clarkii, A. percula, A. ocellaris and
P. biaculeatus were mean (±S.D.) 1.06 (±0.18), 0.74
Table 2—Variable measured in male and female with this five
anemone fishes fanning observation
Sl.No.Species Ob/D Ob/I Ob/H TOB
1. A.sebae
Male 9 5 11 99
Female 9 5
2. A.clarkii
Male 9 5 5 45
Female 9 5
3. A.percula
Male 9 5 12 108
Female 9 5
4. A.ocellaris
Male 9 5 13 117
Female 9 5
5. P.biaculeatus
Male 9 5 7 63
Female 9 5
Ob/D- observation/day, Ob/I- observation/ incubation, Ob/ H-
observation / no of hatching, TOB- total number of observation
(±0.24), 1.09 (±0.30), 1.19 (±0.29), 1.13 (±0.31) mg/lit
(one way ANOVAF 4,39= 3.46, p<0.01) (Figure 13).
The average increased dissolved oxygen depends on
days wise developmental stages involved fanning in
selective male species of female clownfishes viz.
A. sebae, A. clarkii, A. percula, A. ocellaris and
P. biaculeatus were mean (±S.D.) 0.40 (±0.28), 0.43
(±0.16), 0.48 (±6.31), 0.39 (±0.28), 0.36 (±0.21)
mg/lit (one way ANOVA F 4,39= 3.36, p<0.01)
(Figure 14).
INDIAN J. MAR. SCI., VOL. 41, NO. 5, OCTOBER 2012
438
Fig. 11—Hatching success in five clown fishes (Error bar - SE,
- SD and - mean).
Fig. 12—Increased Dissolved oxygen (D.O.) after fanning in
selective species (Error bar - SE, - SD and - mean).
Total body weight
The relation between number of eggs and total
body weight involved fanning in selective female
species are also studied in the same experiment.
This different weight of fishes and their relation
with the fecundity were variables (one way
ANOVA F 4,64= 1.21, p<0.01) (Figure 15). In A. sebae
maximum weight was observed, compare to
P. biaculeatus, A. clarkii, A. ocellaris and A. percula,
where average body weight was 15±2.89, 14±3.11,
13±3.34, 12±2.5 and 12±2.0 gm, respectively.
The average egg laying capability of the fishes in
Fig. 13—Increased Dissolved oxygen (D.O.) depends on days wise
developmental stages involved fanning in selective male species.
Fig. 14—Increased Dissolved oxygen (D.O.) depends on days wise
developmental stages involved fanning in selective female species.
the present study are 1200, 400, 350, 400 and 800
number per spawning in A. sebae, A. clarkii,
A. percula, A. ocellaris and P. biaculeatus.
The total body weight of the female and male
shows the relation with the fanning time (one way
ANOVA F 4,64= 3.36, p<0.01) (Figure 16 & 17) and
fanning frequency (one way ANOVA F 4,64= 3.36,
p<0.01) (Figure 18 & 19).
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439
Fig. 15—Relation between number of eggs and total body weight
(gm) involved fanning in selective female species.
Fig. 16—Relation between female total body weight (gm) and
Fanning frequency (time/h).
Discussion
Parental care consists of fanning and mouthing
the eggs, in addition to oxygenation and removal of
metabolic wastes7. Mouthing would remove dead
eggs and clean the live ones7. Further timing of
fanning has been negatively correlated with oxygen
concentration within fish clutches throughout
development8-13.
Parental care is maintaining the health of a brooder
in many organisms such as amphipods14, crabs8 and
fishes15. Rombough16 found that in the clownfish,
A. melanopus, such behavior was necessary because
of the semicryptic areas with poor water circulation
where the demersal eggs were laid and the boundary
Fig. 17—Relation between female total body weight (gm) and
Fanning time (min) involved fanning in selective female species.
Fig. 18—Relation between male total body weight (gm) and
Fanning frequency (time/h).
Fig. 19—Relation between male total body weight (gm) and
Fanning time (min) involved fanning in selective male species.
INDIAN J. MAR. SCI., VOL. 41, NO. 5, OCTOBER 2012
440
layers formed rapidly around the eggs, reducing the
transfer of oxygen from the surrounding water to the
developing embryos. Males normally spend more
time than females in active egg fanning, which is
typical of the genus Amphiprion17-19 and in the present
study it was well noticed.
Results have revealed that the eggs in the clutch
area developed rapidly when the male fanned
properly (Table 3 and 4). This suggests that the
male would be able to adjust its paternal care
according to the amount of energy and time he
puts into the egg clutch. Other studies have also
shown that the male is more alert in defending
more valuable broods20. Thus, with regard to fanning
and mouthing, the male with full paternity invests
more effort in the clutches than the female.
Many studies have indicated that the females
of several fish species prefer to spawn in nests
with many eggs21-27. In the three-spined stickleback,
Gasterosteus aculeatus, the females have been
shown to prefer nests with many eggs and this has
offered the classical explanation as to why the
males engage in egg stealing in this species22,28-29.
In the present study also the female always prefer
nests for spawning and the fecundity is ranged
between 200-1500 numbers depends the species
and their gonadal maturity. It was also observed
that once the spawning was completed, the female
used to take rest on the anemone and the male only
guarded the egg with more care.
Mass loss of eggs is a consequence of reduced
food intake by the male during the paternal care,
as caring shortens the time available for
food searching20,24,30-32. The paternal care in the
clownfishes exceeds just fanning and mouthing
and there may be some time loss their eggs suddenly
by consuming them. This eggivorous characteristic
is generally called egg eating behavior which could
be due to adverse environmental conditions, physical
disturbances or any biological hazard. This egg
eating behavior was noticed in the present study and
it was more in A. clarkii due to physical disturbance
and was less in A. ocellaris. Another type of
disturbance noticed in the present study was the slow
movement of the sea anemone along with the parent
fish from the vicinity of clutch area, truly causing a
distance between the clutch and parent. Generally,
this type of movement is very rare in captivity.
In this study, A. sebae with S. mertensii moved
occasionally.
Table 3—Variable measured in male and female, five anemone
fishes fanning observation
Sl.
No. Species Clutch
area(cm)
Fecundity (
nos.
of eggs/clutch)
Fanning behaviour
1. A.sebae 4.5-8.5 600-1500 Both fanned by
pectoral fins
2. A.clarkii 3.5-4.5 500-800 Female very rare.
Male using fanning
3. A.percula 3.5-5.0 300-500 Both are actively
fanning
4. A.ocellaris 3.5-4.5 200-500 Male and female
very active in
fanning behaviour
5. P.biaculeatus 4.0-5.0 200-500 Male actively
fanned, female
occationally
Table 4—Measured in male and female, five anemone fishes in
clutch area, fecundity and fanning duration
Sl.No.
Species CL F FD
1. A.sebae
Male 4.5-8.5 600-1500 1.3 (± 0.13)
Female 0.8 (± 0.22)
2. A.clarkii
Male 3.5-4.5 500-800 0.5 (± 0.34)
Female 0.3 (± 0.29)
3. A.percula
Male 3.5-5.0 300-500 1.1 (± 0.31)
Female 0.7 (± 0.19)
4. A.ocellaris
Male 3.5-4.5 200-500 1.2 (± 0.28)
Female 0.6 (± 0.28)
5. P.biaculeatus
Male 4.0-5.0 300-600 1.1(± 0.3)
Female 0.6 (± 0.31)
CL- clutch area (cm), F- fecundity (Nos. of eggs/
clutch),
FD- fanning duration (Min./ventilation)
Conclusion
In conclusion, the male increase their paternal
caring effort as the significance of the hatching
successes. Increased fanning time not only increases
the hatching success of clownfishes but fanning
frequency also play same role in clownfish parental
care. Female showed less preference for eggs
caring behavior than the male in all the five
species. It also revealed that the fanning was an
imperative biology of their life cycle and it’s a
crucial mechanism to develop their eggs and
receiving hatching success.
GHOSH et al.: DETERMINING THE LEVEL OF PARENTAL CARE RELATING FANNING BEHAVIOR OF FIVE SPECIES
441
Acknowledgement
Authors are grateful to the authorities of
Annamalai University for providing facilities and the
Centre for Marine Living Resources and Ecology,
Ministry of Earth Sciences, Govt. of India for
financial support.
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Publisher Summary This chapter describes respiratory gas exchange, aerobic metabolism, and the effects of hypoxia during early life of fish. Analytical models provide a useful framework for the study of respiratory gas exchange. The cascade model, in particular, has been widely used to describe various aspects of vertebrate respiratory function. The laminar boundary layer is a semistagnant region of water adjacent to the egg surface where oxygen is depleted and metabolic wastes accumulate. The boundary layer actually has no outer limit, but for practical purposes are usually defined as the distance from the egg surface where local conditions are equivalent to 99% of free-stream conditions. Larvae experience respiratory distress during the transition from cutaneous to branchial gas exchange. The metabolic response of larvae to declining oxygen levels appears to be more variable than in embryos. The influence of salinity on the rate of oxygen consumption of embryos and larvae has been investigated in only a few species, but results to date suggest that net ionoregulatory costs are negligible once acclimation has occurred.
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Male three-spined sticklebacks, Gasterosteus aculeatuis, are violently territorial when breeding. Even so, their stomachs commonly contain nonspecific eggs. Curiously, this conflict between the inviolability of territories and the consumption of nonspecific eggs is not resolved by the regular observations of male sticklebacks raiding each other's nests for eggs. This is so because stolen eggs are not eaten. Instead, they are invariably transferred to the thief's nest. Sticklebacks apparently do not recognize their own eggs, so stolen eggs must not serve primarily as food. My evidence that males do not recognize their own eggs is based on the success of van lersel's (1953) nest-switching experiments and on van den Assem's (1967) demonstration that males will hatch eggs that they did not fertilize. Both were laboratory studies, so the possibility that the strange eggs were recognized but not consumed because of an abundance of other food cannot be ruled out. So the plot has thickened. There is an illicit trafficking of eggs between nests, but this trafficking apparently does not explain the observed egg cannibalism. Furthermore, the egg raids themselves are complex in that they are often presaged by a behavioral sequence termed "sneaking" (van den Assem 1967). Here the prospective raider, usually a neighbor, swims to his victim's territorial boundary and then sinks to the bottom and shortly loses his vivid nuptial colors. He then stealthily approaches his victim's nest, creeping millimeter by millimeter along the bottom (Morris 1952; van den Assem 1967; Wootton 1971). Sneaking is often timed to coincide with courtship, and, when the female enters the nest to spawn, the waiting invader suddenly rushes to cover the remaining distance. The sudden "appearance" of the invader in this rush often enables him to pass through the nest, presumably spawning over the eggs, before the resident male does so. This seems to happen because the resident becomes preoccupied with attacks upon the invader (Morris 1952; van den Assem 1967, p. 104). Sneaking may appear to facilitate cuckoldry, and Barlow (1967) has so interpreted such "pseudofemale" behavior in a South American leaf fish, Polycentrus schomburgkii. But the plot is thicker. Additional observations suggest that stolen spawnings may be a subsidiary adaptation in sticklebacks and that sneaking primarily facilitates egg raiding. First, invaders that succeed
Article
Attempts by males to steal fertilizations from other males are common in many species. In some sticklebacks, males also are known to steal eggs from the nests of rivals and to carry them back to their own nests. However, the genetic consequences of these nest-raiding behaviors seldom have been investigated. Here we assess genetically the prevalence of sneaked fertilizations and egg stealing, and we describe the mating system in a natural population of the fifteenspine stickleback. Six microsatellite markers were developed and employed to assay a total of 1307 embryos from 28 nests. Guardian males and all nest-holding males in the local area also were genotyped for two to six loci. Analysis of male genotypes and those of embryos revealed that five of the 28 nests (18%) contained progeny from sneaked fertilizations, and that four of the 24 nests (17%) with resident males contained stolen egg clutches. Comparisons of the composite DNA genotypes of nest-holding males against those of inferred sneakers implicated one nest holder as the sneaker of a nest seven meters from his own. Also, the genetic data demonstrated that nests of males frequently contain eggs from multiple females. The multilocus genotypes of inferred mothers indicated that females mate with multiple males, sometimes over distances greater than one kilometer.
Article
There is evidence that female three-spined sticklebacks, Gasterosteus aculeatus L., prefer to mate with males whose nests contain eggs rather than with males with empty nests. While there is consensus on this point, a dispute exists about whether this preference should be attributed to a direct effect of the eggs on the female's entering the nest or, alternatively, to a positive impact of the eggs on the courtship behaviour and breeding coloration of the male. In the field experiment reported here females strongly preferred nests with eggs over empty nests. Additionally, females were less likely to enter risky nests with eggs: nests that contained fewer eggs than one average clutch or more eggs than the average nest content of parental males in this population. However, in the field possible differences in male attractiveness were not controlled for. In supplementary laboratory experiments the effect on female choice of possible changes in male attractiveness (intensified courtship and coloration) as a result of the presence of eggs in the nest was tested. Other differences in male attractiveness as a result of differences in male quality (body size, breeding coloration before the test, territory quality and size) were controlled for. When females had no access to the nests, they showed no preference for males with eggs in their nests in simultaneous choice tests. These results, together with the earlier published data, make it likely that the preference of females for nests with eggs is partly a direct consequence of the eggs themselves. So female sticklebacks are influenced by the mate choice behaviour of other females, but remain selective as to the actual nest content.