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MILKFISH, Chanos chanos (FORSSKAL, 1775), FRY SEASONALITY IN VANUATU: THEIR AVAILABILITY AND ABUNDANCE IN THE COASTAL SHORELINE OF EFATE ISLAND
By
Ronnick Spenly Shedrack
A Thesis submitted in fulfillment of the requirements for the Degree of Master of Science in Marine Science
Copyright © 2017 by Ronick Spenly Shedrack
School of Marine Studies
Faculty of Science, Technology and Environment
The University of the South Pacific
November, 2017
Statement by Autho
I, Ronick Spenly She
best of my knowledge
overlapping with mat
institution, except wh
Signature:
Name: Ronick Spenl
Statement by Superv
The research in this th
knowledge is the sole
Signature:
Name: Dr. Marta Fe
Declaration of Originality
or
drack, declare that this thesis is my own work
e, it contains no material previously publishe
terial submitted for the award of any other de
here due acknowledgement is made in the text
Date: 10 November
ly Shedrack Student ID No: S110
visor
hesis was performed under my supervision an
e work of Mr. Ronick Spenly Shedrack.
Date: 20/11/2
erriera Designation: Princip
k and that, to the
d, or substantially
gree at any
t.
2017
62153
nd to my
2017
pal Supervi
iii
Dedication
This thesis is dedicated to my dear parents Belinda Toa Spenly and Spenly Shedrack
Salemomo.
iv
Acknowledgements
I express my sincere thanks and appreciation to the Australian Centre of Agricultural
Research (ACIAR) for funding me for the Master of Science degree under the
ACIAR-USP scholarship Scheme and to USP for facilitating the sponsorship and
funding another 6-month for the write up until completion of the program.
The completion of my thesis was made possible through the invaluable support from
my principal supervisor, Dr. Marta Ferreira and co-supervisor, Dr. Susanna Piovano
who provided constructive criticism, suggestion, comments and encouragement
during the course of the study.
Dr. Marta is also immeasurably valued for providing me with support to Otolith
analysis technique that is necessary to complete the thesis. I also acknowledge Dr.
Susanna for various constructive criticisms polishing this study.
I express my appreciation to Mr. Sompert Gereva for his invaluable support, advice
and assistance throughout the project with the field data collection, sampling
materials and providing me space at the Vanuatu fisheries department to complete
the thesis write up. Thanks are also extended to Dr. Timonthy Pickering for his
various advices on the field data collection techniques.
I also express my gratitude to the Vanuatu Meteorological and Geo-hazard
Department staff particular, a special thanks to Melinda Natapei, David Gibson and
Philip Mansale for supporting with the data collection until the completion of the
fieldwork.
I would like to thank Martinez and his colleagues (2006) for allowing me to
reproduce their figure of milkfish life stages in Figure 1.2.
My heartfelt gratitude is extended to my family members, in particular Christian
Shedrack, Semion Shedrack, Ajay Shedrack, Susila Salemomo and Welpi Tane for
assisting me throughout the field data collection.
Lastly, I am thankful to my then girlfriend and now my wife, Rose Charley for her love, support and assistance with lab work, food preparation and for everything, she did for me.
v
Abstract
This study describes the seasonality and abundance of milkfish Chanos chanos fry in
the coastal shorelines of Efate Island in order to assess the feasibility of milkfish
aquaculture in Vanuatu. The study will enable the knowledge of fry collection so that
this species can be cultured for food. Four sampling sites have been assessed in
February 2016 for the presence of milkfish fry namely, Erakor, Kawenu, Mele and
Teouma. The site with the highest abundance was selected for further seasonality
sampling over a one-year period from February 2016 to February 2017.
Milkfish fry were collected from February to May and October to December 2016
and, again from January to February 2017. Milkfish fry was absent in other months
of the year. Two peaks of fry occurrence were observed, the 1st peak was in April and
the 2nd peak was in November. The fry abundance in Teouma was high (relative to
other locations such as Erakor, Kawenu and Mele) over the period of seasonality
sampling. The abundance between Moon phases showed that more fry were caught
in New Moon and 3rd quarter Moon whereby a non-significant difference (p > 0.05)
was observed. Milkfish fry collected on Teouma coast measured between 8 mm to 15
mm total length (TL) and the weight range was between 3 mg to 9 mg. The fry were
aged 13 to 25 days old, which mean that they spend 2 to 4 weeks in the surf zone
before disappearing into other habitats. Environmental variables assessed showed
cloud cover is positively correlated with fry abundance, similarly, wind speed and
current speed with length, and rainfall and turbidity with weight. Other variable such
as temperature in situ do not significantly correlated with fry abundance, weight and
length.
The prolonged fry seasonality in Vanuatu is advantageous for milkfish fry
aquaculture, however the very low abundance documented in Teouma are not.
Before a final decision on the viability of milkfish culture in Vanuatu is made, an
assessment of multiple sites is recommended. To be cost-effective such a study could
be informed by this research with sampling limited to the months when fry
abundances peaks. If milkfish culture does proof to be viable, then the landowners of
fry collection ground may need to manage activities along the coast to reduce
impacts on fry recruitment habitat. Furthermore, fishers may want to consider
vi
managing fishing of milkfish during spawning periods in order to protect spawning
stocks.
vii
Abbreviation and Acronyms
ACIAR Australian Center for International Agricultural research
ANOVA Analysis of variance
BOD Biological demand for oxygen
CT Concentration test oC Degree Celsius
DO Dissolve oxygen
FAO Food and Agricultural Organization
GSI Gonadosomatic Index
IPCC Intergovernmental Panel on Climate Change
VFD Vanuatu Fisheries Department
PIC Pacific Island Country
ppt Parts per thousand
VMGD Vanuatu Meteorology and Geo-hazard Department
COSPPac Climate and Ocean Support Program in the Pacific
SST Sea surface temperature
SPC Pacific Community
USP The University of the South Pacific
TL Total length
viii
Table of Contents Acknowledgements ..................................................................................................... iv
Abstract ........................................................................................................................ v
Abbreviation and Acronyms ...................................................................................... vii
List of figures ............................................................................................................... x
List of tables ............................................................................................................... xii
Thesis organisation.................................................................................................... xiii
Introduction .................................................................................................................. 1
1.1 Milkfish aquaculture ...................................................................................... 1
1.2 Reviews on milkfish (Chanos chanos) .......................................................... 1
1.2.1 Systematics .................................................................................................. 2
1.2.2 Morphology ................................................................................................. 3
1.3 Habitat and life history .................................................................................. 3
1.3.1 Life history ................................................................................................... 4
1.4 Food, Growth and Feeding Habit .................................................................. 7
1.5 Milkfish distribution ...................................................................................... 7
1.6. Milkfish aging, spawning and fry seasonality ................................................... 8
1.6.1 Milkfish spawning ....................................................................................... 9
1.6.2 Seasonality of fry ....................................................................................... 10
1.7 Migration and movement of fry ....................................................................... 11
1.8 Predation on milkfish fry .................................................................................. 12
1.9 Overview of main methods of fry collection .................................................... 12
1.10 Food demand in the Pacific island countries .................................................. 14
1.11 Conclusion ...................................................................................................... 16
1.12 Research objectives ........................................................................................ 16
Chapter 2: Methodology ............................................................................................ 18
2.0 Site selection and description ........................................................................... 18
2.1 Sampling materials and methods ...................................................................... 20
2.1.1 Sampling materials .................................................................................... 20
2.1.2 Bulldozer net construction ......................................................................... 21
2.1.3 Sampling method ....................................................................................... 23
2.2 Fry identification .............................................................................................. 24
2.3 Fry length and weight measurement ................................................................ 24
ix
2.4 Otolith analysis ................................................................................................. 25
2.5 Data analysis ..................................................................................................... 26
Chapter 3: Results ...................................................................................................... 27
3.1 Site assessment and selection ........................................................................... 27
3.2 Milkfish fry occurrence in Teouma .................................................................. 29
3.2.1 Monthly occurrence and abundance .......................................................... 29
3.2.2 Milkfish fry length and weight .................................................................. 30
3.3 Effect of Moon phases ...................................................................................... 32
3.3.1 Abundance by Moon phase........................................................................ 32
3.3.2 Total length and weight in Moon phase over one year .............................. 33
3.4 Comparison of fry abundance with other indices ............................................. 34
3.4.1 Environmental variables correlation with fry abundance .......................... 34
3.4.2 Environmental variables correlation with length and weight of milkfish fry ............................................................................................................................ 35
3.5 Otolith analysis and age of the fry per month and spawning estimates ........... 37
3.6 The age distribution .......................................................................................... 38
3.7 Spawning time estimates for Teouma over one year period ............................ 40
3.8 Comparison of fry abundance in the Melanesian PICs .................................... 41
Chapter 4: Discussion ................................................................................................ 42
Chapter 5: Conclusion ................................................................................................ 50
References .................................................................................................................. 52
Appendix .................................................................................................................... 66
Appendix A. Table of p-values for the ANOVA test ............................................ 66
Appendix B. The daily SST from VMDG and monthly mean, maximum, minimum, median and standard deviation beginning from February 17th 2016 to February 17, 2017. ............................................................................................. 69
Appendix C. Milkfish fry mean length and weight plus the descriptive statistic for Teouma over one year period ....................................................................... 70
x
List of figures
Figure 1.1 Picture of adult milkfish……………………………………………...…...2
Figure 1.2 Milkfish life cycles, 1-7 indicates the seven development stages in the
milkfish life cycle (obtained with permission from Martinez, Tseng et al. (2006)…..4
Figure 1.3 A graphical representation of peak and lean season of milkfish fry occurrence in several countries with data retrieved from Kuo et al. (1979),Villaluz (1986) and Vuto et al. (2014)…..………………………………………...………….19
Figure 2.1 Map of Vanuatu where the study is based. Retrieved from
http://www.nationsonline.org/oneworld/map/vanuatu-map .…….…………….……...19
Figure 2.2 Map of the four sites Erakor, Kawenu, Mele and Teouma where milkfish
fry was assessed during the preliminary study, and the Vanuatu capital (marked with
a star)…………………………………..………………………………………….…20
Figure 2.3 The bulldozer net built and used for this study. Picture shot at Teouma on
March 2016. ……………..……………………………………….…………………22
Figure 2.4 The bamboo frame design for the bulldozer net (Pickering, Tanaka et al.
2012)…………………………………………………….………………………..…22
Figure 2.5 Net size and design modified from Pickering, Tanaka et al. (2012)..…...23
Figure 2.6 Milkfish fry collected in a white plastic basin for visual identification. The
black dot from left to right showed position of grey spot on caudal fin, dark spot on
middle of body (air bladder) and two dark spot (eyes) on side each side of head…..24
Figure 3.1 Milkfish fry total abundance (number of fry captured during the
exploratory trial done on Teouma, Erakor, Mele and Kawenu). The different letters
indicate significant difference in abundance (p < 0.05) while the same letter indicate
no significant difference (p > 0.05).…………………………………………….…...29
Figure 3.2 Milkfish fry abundance and occurrence with SST by month for Teouma.
The different letters indicate significant difference in fry abundance (p < 0.05)..….30
Figure 3.3 The total length frequency of milkfish fry (N = 369) in the Teouma……31
xi
Figure 3.4 The frequency distribution of milkfish fry (N = 369) weight in
Teouma……………………………………………………………………………...31
Figure 3.5 Milkfish fry mean total length (A) and weight (B) by month of
occurrence. The error bars represent the standard deviation of the mean. …….…...32
Figure 3.6 Milkfish fry cumulative abundance in all Moon phases for Teouma
throughout the year. The different letters indicate significant difference is abundance
between Moon phases (p < 0.05). ……………………………….........…………….33
Figure 3.7 Milkfish fry mean total length (A) and weight (B) in four Moon phases.
The different letters show significant difference in mean total length or weight (p <
0.05) and the error bars represent the standard deviation of the mean. .……..……..34
Figure 3.8 The environmental variable correlation with milkfish fry abundance,
rainfall, cloud cover, wind speed, temperature, turbidity (transparency) and current
speed...……………………………………………………………………………....35
Figure 3.9 Environmental variable, temperature, cloud cover, rainfall, turbidity
(transparency), wind speed and current speed correlation with milkfish fry total
length ..…………………………………………………………..…………………..36
Figure 3.10 Environmental variables, temperature, cloud cover, rainfall, turbidity
(transparency), wind speed and current speed, correlation to milkfish fry weight
...…………………………………………………………………………..…………37
Figure 3.11 Photographs a, sagittae otolith from 14 days old larvae, 11 mm total
length (TL). (x400), b an otolith of a 15 day larva of 11 mm TL and, c a otolith of a
22 day larva of 14 mm TL. In photograph a, letters. A, anterior; P, posterior; D,
dorsal; V, ventral. The dot (●) are the increment dark zone, the star ( ) indicate the
core and the arrow (↓) indicate the discontinuity zone. …………...………………..38
Figure 3. 12 The age frequency of milkfish fry in Teouma coastal shoreline ….…..39
Figure 3.13 The mean age of fry per month of occurrence in Teouma. The error bars
represent the standard deviation of the mean. ..………...…...………………………39
Figure 3.14 Correlation between the age and length of milkfish fry ………….……40
xii
Figure 3.15 The spawning season of milkfish fry (N =63) based on different days over a one year period with mean Monthly SST…………………...……………….41
List of tables
Table 3.1 The monthly environmental parameters (turbidity (transparency), current
speed, salinity and SST) assessed and the landscape features for Mele, Kawenu,
Erakor and Teouma. The monthly values are presented as mean ± standard
deviation…………………………………………………………………………..…28
Table 3.2 the fry abundance between Fiji, Solomon Islands and Vanuatu……….....41
xiii
Thesis organisation
There are six chapters in this thesis.
Chapter 1 presents a literature review about what is known and where the gaps of
knowledge which require further research on milkfish (Chanos chanos) larval
recruitment. Other parameters affecting milkfish larval development and fry
presence along coastal areas were also reviewed, and seasonality of spawning
and culture of milkfish in the Pacific Island Countries (PICs) were outlined. This
chapter also presents the purpose and scope of the study, and the hypotheses and
statement of problems to be addressed. The reason for a milkfish fry seasonality
study in Vanuatu is also outlined and the advantages of having an additional fish
species for the local aquaculture industry are discussed.
Chapter 2 describes the materials and methods used to undertake this study. The study
sites and the methodologies applied to collect the larvae are shown and the data
and statistical analyses highlighted.
Chapter 3 presents the results that include seasonality of fry occurrence and abundance,
abundance in Moon phases, environmental variables affecting abundance and
size of fry, and finally the spawning period estimation.
Chapter 4 contains the discussion of research findings and reflection within the literature
context. The assessment for each single site was discussed with respect to the
abundance of milkfish fry and the parameters assessed. The length, weight and
age frequency of milkfish fry were also discussed with respect to seasonality and
Moon phases.
Finally, chapter 5 contains the conclusion and recommendations for further research to
address limitations and constraints in this study and to further engage researchers
to develop milkfish farming in Vanuatu.
1
Introduction
1.1 Milkfish aquaculture
Milkfish Chanos chanos is one of the many species cultivated to address food
shortage and meet the food protein demand in the Asian-Pacific region (Agbayani et
al. 1989; FAO 2014; Martinez et al. 2006). The knowledge about biology, spawning
and culture of the species has aided the development of milkfish culture industry in
many countries, in particular Asian nations (Bagarinao 1991; Martinez et al. 2006;
Sulu et al. 2016). Milkfish aquaculture dated back about 4 to 6 centuries ago in India,
Philippines, Indonesia and Taiwan (Agbayani et al. 1989; Nelson 2007). The prolong
season of fry appearance aids aquaculture development as fry can be collected and
stocked in water ponds in many months of the year (Bagarinao et al. 1987). In recent
decades, many Asian countries have used technological advances to move into
milkfish mass aquaculture. Pacific Island Countries and Territories (PICTs), milkfish
aquaculture is new, while in Nauru milkfish culture is part of their tradition for
household food consumption (Spennemann 2002). Vanuatu is one PIC with no
quantitative records of milkfish fry appearances or history and culture, however
there is history of local people collecting juveniles on some islands for food
(Pickering et al. 2012).
The knowledge of milkfish fry seasonality is lacking in some PICs although milkfish
fry are available in their coastal areas (Dela Cruz 1997). This knowledge is important
for PICs to adopt milkfish culture, as access to advanced hatchery technology is poor
and culture may depend on availability of wild fry for stocking ponds. Hence, the
purpose of this study is to determine milkfish fry seasonality in Vanuatu and evaluate
the potential of milkfish aquaculture.
1.2 Reviews on milkfish (Chanos chanos)
Milkfish Chanos chanos (Forsskål, 1775) is the only fish species of the family
Chanidae in the order Gonorynchiformes that comprises four families, seven genera
and 27 species (Bagarinao 1994b; Lim et al. 2002; Nelson 2006). The specific
2
epithet chanos (Greek for a fish of remarkable voracity) was attributed to milkfish in
1775 by the Swedish naturalist Peter Forsskål, who made a clear description of the
fish based on a specimen from the Red Sea that was preserved at the Zoological
Museum of the Copenhagen, University in Denmark (Klausewitz 1965). In 1803, the
French naturalist Bernard Germain de Lacépède elevated the specific epithet to the
generic level, naming the species Chanos arabicus, and further works from other
authors later described the milkfish under different names and synonyms (Bagarinao
1994b; Bagarinao 1991; Brian 2015). The review by Bagarinao (1994b) revealed that
nine different names were used to describe milkfish by Cuvier and Valenciennes
while other authors described it under eighteen other synonyms. The name Chanos
chanos was validated and used first by Klunzinger (1870) and following authors did
the same (Fricke 2008) .
A picture of a typical adult milkfish is shown in figure 1.1 whereby it has a single
dorsal fin, large fork tail and silver or milk colour. The milkfish swim over the sandy
bottom and around islands where coral reefs are present (Figure 1.1).
Study species: Chanos chanos
Kingdom: Chordata
Phylum: Vertebrata
Class: Osteichthyes
Order: Gonorynchiformes
Family: Chanidae
Genus: Chanos (Lacépède, 1803)
Species: Chanos chanos (Forsskål, 1775)
Figure 1.1 Picture of adult milkfish (Source: www.aqtinfo.com)
1.2.1 Systematics
Milkfish is considered a more ancestral ostariophysian which belongs to a Monotype
gonorynchiform family and is related mostly to the freshwater Ostariophysi
(Bagarinao 1994b). Although milkfish morphological characteristics have
significantly contributed to a much better understanding of the species, molecular
3
studies determined the number of chromosomes in milkfish as diploids, 2n = 32
consisting of 7 pairs of metacentric, 2 pair of submetacentric and 7 pair of
acrocentric pairs of chromosomes (Arai et al. 1976). The number of chromosomes is
low when compared to other primitive teleosts (Brian 2015). The genetic
identification has helped to distinguish milkfish populations, whereby 9 populations
were described in the Indo-Pacific of which 4 populations were determined as the
Indian, Thailand, Philippine-Taiwan-Indonesia and Tahiti (Bagarinao 1994b;
Bagarinao 1991; Kumagai 1990; Senta et al. 1977). Samples of larvae and juvenile
specimens obtained from Kiribati, Tonga, Hawaii and Panama have revealed four
other populations and a suggestion was made that there is a possibility for one
population in the coast of Africa (Bagarinao 1994b).
1.2.2 Morphology
Adult milkfish has been described as having a silvery colour, muscular, streamlined
body and powerful forked tail (Bagarinao 1994b). Recently Brian (2015) stated that,
“The mouth is small and lacks teeth. There is a notch on the upper jaw in the mid-
line into which a lower jaw protuberance fits. The large eyes have an adipose eyelid.
The intestine is very long with many folds. The lower part of the esophagus has a
"gizzard”, an area with longitudinal folds. Lateral line scales 70-92, with 3-11 on
the tail fin”.
There are also variant forms of milkfish reported, the “goldfish” form from
Philippine and Indonesia and the “dwarf or hunchback –shade type” found in
Hawaii, Indonesia and Australia, but the knowledge about these other forms of
milkfish is limited since they rarely occur (Bagarinao 1994b).
1.3 Habitat and life history
Milkfish are usually found in offshore marine waters and shallow coastal
embayment, but also frequently enter estuaries and occasionally penetrate freshwater
streams (Bagarinao 1994b). Adults occur in small to large schools near the coasts or
around the islands w
and larvae are pelagic
in coastal wetlands
(Bagarinao et al. 198
sexually and spawn i
phytoplankton and z
benthic invertebrates
Milkfish at the differ
the open ocean in adu
spawn (Figure 1.2).
Figure 1.2 Milkfish limilkfish life cycle (ob
Milkfish fry enters a
juvenile habitat. Acco
in estuaries and near
of water flow (Lin et
swamps and lagoons
1.3.1 Life history
The life history and
many researchers, a
4
where reefs are well developed (Bagarinao 19
c for up to 2-3 weeks and older larvae migrat
(mangroves, estuaries) occasionally enterin
87). Juveniles and sub-adults return to the sea
in fully saline marine water (Nelson 2007). M
zooplankton; juveniles and adults feed on cy
s, and even pelagic fish eggs and larvae
rent life stages frequent various habitats befor
ult stage where they spend the rest of their li
ife cycles, 1-7 indicates the seven developmenbtained with permission from Martinez, Tseng
a coastal lagoon and swamp before they m
ording to Martinez et al. (2006), milkfish fry
river mouth. The fry tend to migrate into riv
t al. 2003; Pillay et al. 2005) and other coas
to enter into juvenile stage.
habitat of milkfish in the wild was docume
although many errors have been pointed o
991). Milkfish eggs
te inshore and settle
ng freshwater lakes
a where they mature
Milkfish fry feed on
yanobacteria, small
(Bagarinao 1991).
re they migrate into
ves and continue to
nt stages in the g et al. (2006)
migrate further into
y are more abundant
vers in the direction
stal habitats such as
ented in the past by
out in more recent
5
research, like the review by Bagarinao (1994b). In the last few decades milkfish have
been demonstrated to mature and spawn under various condition of captivity, and
hatcheries have produced larvae to supply ponds (Bagarinao 1991). Nowadays the
aquaculture industry has employed facilities and techniques that facilitate breeding in
captivity and the transition of fry to juvenile in captivity, including mass propagation
with minimum mortality (Bagarinao 1991; Nelson 2007).
Milkfish fertilized eggs are spherical and pelagic with a diameter ranging from 1.1 to
1.2 mm, and the development of the embryo takes 20 - 35 hours at a temperature
range from 26 ºC to 32 ºC, and at a salinity between 29 to 34 parts per thousand (ppt)
(Bagarinao 1991). Delsman (1926) was the first scientist who took note of milkfish
eggs when he examined the oocytes and later collected 15 eggs from the Java sea;
later his identification was tested and proven to be correct after induced spawning
was done in captivity (Bagarinao 1994b; Liao et al. 1979) . The spawning grounds of
milkfish are in clean and clear saline water with warm temperature (25-30 ºC) and
shallow water (< 200 m) over coral reefs and sandy beaches at a distance of about 6
km offshore (Brian 2015). The spawning location is also based on the need to
minimize predation (Brian 2015), and close to shore to enhance larvae migration to
coastal habitat (Johannes 1978).
Milkfish larvae are categorized into 5 developing stages: i) yolk-sac larvae (3.3-5.4
mm total length (TL), lasts for 3 days); ii) pre-flexion larvae (3.4-6.2 mm TL, lasts
for 5 days); iii) flexion larvae (4.4-9.9 mm TL, lasts for 6 days); iv) post flexion
larvae (9.5-14.9 mm TL, lasts for 7 days); and v) transformation larvae (9.5 -16.5
mm TL also named as fry are free swimming larvae, last from 2 to 4 weeks), after a
detailed description was made by various authors (Bagarinao 1991). Milkfish larvae
are pelagic, while at hatching, the yolk sac larvae are about 3.5 mm TL; the lava
begin to feed when their eyes are fully pigmented (3 days old) and the mouth will
open although some yolk is still attached (Bagarinao 1994a). It takes 36 hours for the
larvae to grow into 5 mm total length and to consume about 90 % of the yolk until
day 5 when the yolk is completely exhausted. The size of the egg, larvae, amount of
yolk and mouth is far greater in milkfish than any other tropical marine fish
(Bagarinao et al. 1986). Younger larvae occur both onshore and offshore while older
larvae are found near shore only (Bagarinao et al. 1987). A study on larvae
6
movement in the Great Barrier Reef by Leis et al. (1991) revealed that the larvae
move to the juvenile habitat through passive advection and active migration. Active
migration is employed by larvae that gain a certain degree of morphological
characteristics, probably at 10 mm TL and age of 2 weeks old (Bagarinao 1994b).
The juvenile milkfish has a minimum length of 20 mm and the shape and structures
of adult (Bagarinao 1994b; Bagarinao 1991). The juveniles, length between 2 to 10
cm TL, are called “fingerlings”, particularly in the aquaculture industry (Bagarinao
1994b). Milkfish larger than 30 cm are found in a variety of habitats such as
estuaries, coral lagoons, tidal creeks, mud flats and tide pools that are protected areas
and characterized by rich food supply (Buri 1980; Kumagai et al. 1985; Kumagai et
al. 1981). The juveniles feed mainly on food from the bottom such as cyanobacteria,
diatoms, detritus, filamentous green algae and invertebrate such as worms and
crustacean (Bagarinao 1994b; Bagarinao 1991). The wild juveniles feed during the
day as determined by gut content analysis, whereby morning guts were empty while
those during the day were loaded with food (Kumagai et al. 1985).
An adult milkfish has a length of 50-150 cm TL and weight range of 4-14 kg with
age from 3-15 years (Bagarinao 1991). More recently, Brian (2015) has noted in a
review of the milkfish of Iran that this species has the size of up to 1.85 m and 18.6
kg. Adult milkfish are powerful swimmers and are seen in large schools near islands
and the coast where reefs are well developed, and also in coastal lagoons where they
often swim with their dorsal fins above water like sharks (Bagarinao 1994a; Brian
2015). Adults are seen and caught near shore usually during breeding season
(Kumagai 1990; Tampi 1957). Milkfish reproduction in the wild is not well
understood, although milkfish has been successfully bred in captivity in the
Philippines, Taiwan, Hawaii and in Indonesia, and in Kiribati (Bagarinao 1994b). In
nature milkfish may reach sexual maturity at age 3-5 years and in captivity at age 8-
10 years (Bagarinao 1994b; Brian 2015). The females may spawn more than twice
per year, both in captivity and in the wild (Marte et al. 1986b; Schuster 1960; Silas et
al. 1982; Tampi 1957), and 3-13 kg female may produce 0.5 to 6 million eggs.
Spawning usually happens at night and is triggered by lunar and seasonal periodicity
(Kumagai 1990; Marte et al. 1986b).
7
1.4 Food, Growth and Feeding Habit
Milkfish larvae normally feed on phytoplankton and zooplankton (Bagarinao 1991;
Brian 2015), and juveniles in a variety of food, including cyanobacteria, detritus,
diatoms, filamentous green algae, fish eggs and larvae, and some invertebrates such
as crustaceans and worms (Bagarinao et al. 1986). Milkfish larvae are particulate
visual feeders. They feed by snapping prey such as rotifer, water flea, copepod and
brine shrimp (Bagarinao 1991). However, Carreon et al. (1984), evaluated the
feeding response of larvae on diets of plankton and detritus (containing bacteria,
various species of cyanobacteria, diatoms, rotifers and protozoans) in race way tanks
whereby growth, health and survival was better in larvae supplied with natural
planktons in comparison to those reared on detritus.
An adult milkfish feeds on plankton, benthic plants and animals (Bagarinao 1994b),
they feed on plankton by swimming through plankton masses and also larval fish
schools. They also graze on the rock's surface and floating algae therefore, adult
milkfish is an opportunistic generalist (Bagarinao 1994b). Although juveniles and
adult milkfish feeds on both day and night they are more active during the day
(Kawamura et al. 1981).
1.5 Milkfish distribution
Milkfish are widely distributed in tropical, subtropical seas and around Oceanic
islands, where seawater temperatures are greater than 20 °C, ranging from Red Sea
and African coast to East Pacific, South Pacific coast of the U.S.A, Central America,
Central and Western Pacific, Northern Pacific, South Pacific to New Zealand, the
Indian Ocean and around Australia (Figure 1.3). Milkfish is said to be an Indo-West
Pacific fish species, although it ranges across, to the eastern Pacific (Brian 2015).
The adults can swim at the speed of 2 km/h although they were never caught in the
high seas (Brian 2015). It is thought that milkfish recolonized the eastern Pacific by
dispersal from the Indo-West-Pacific via the equatorial counter current (Briggs
1961), even though there is still debate regarding these theory (Bagarinao 1994a;
8
Bagarinao 1991; Brian 2015; Leis 1984). Milkfish are not found in tropical waters
that are affected by cold ocean currents (Bagarinao 1991).
1.6. Milkfish aging, spawning and fry seasonality
Otoliths are fish ear bones, small calcified structures found inside the inner ear
region to help with hearing and orientation (Begg et al. 2005; Popper et al. 2005).
The shapes are different for each three type of otolith such as the lapilli, sagittae and
asterisci (Panfili et al. 2002). The sound wave is detected from the movement of
otolith and the fish movement that send mechanical signals which assist in hearing
and orientation (Popper et al. 2005).
Otoliths have been used widely to determine age, growth rate, development and life
history of fish and are helpful in understanding fish systematics and evolution (Fey et
al. 2005; Popper et al. 2005). Otoliths are formed at the hatching of the eggs and
grow daily rings made of calcium ion from the endolymph precipitate (Warner et al.
2005). The rings continue to grow around the core and form a layer called
microstructures, increment or growth rings. The daily growth rings or increments
become narrower as the fish age increase, which also pose a difficulty in determining
the life history of older fish (Baumann et al. 2005).
The growth of milkfish larvae follows a sigmoid growth curve (Liao et al. 1979).
Kumagai et al. (1981) stated that fry in shore waters grow at a rate of 0.5 mm per
day. Tzeng et al. (1988) have determined that the first otolith increment in reared
milkfish larvae is formed 2 days from hatching and continues on a daily basis. A
subsequent study by Tzeng et al. (1989) on wild caught larvae using the
oxytetracycline method has validated that increments form at the rate of 1 per day
regardless of growth rate. Kumagai (1990) has determine the size distribution
frequency of 10,000 milkfish fry collected in shore waters where he concluded that
shore caught fry grow at the rate of 0.5 mm/day and larvae remains at 5 mm for
about 4 days from hatching. Therefore the relationship between size and age was
derived from the following equation (Equation 1) from Bagarinao (1991).
Equation 1: TL = 5.0 + 0.5 (D-4)
9
Where, TL=total length and D=day from hatching
Furthermore, the growth of shore caught milkfish fry is temperature dependent and
the transformation stage begins in 5 days of rearing (Villaluz et al. 1983a). The
growth of fry also varies with factors such as diet and feeding (Bagarinao 1991).
1.6.1 Milkfish spawning
The spawning age of milkfish was determined to be 3 to 10 years in a temperature
range of 25 to 30 °C and salinity of 29 to 34 ppt (Bagarinao 1991). Milkfish females
can produce large amount of eggs with ovaries weighing 10 % to 25 % of the body
weight when mature (Bagarinao 1991). The frequency of spawning as mentioned
previously occurs probably more than once per year, based on indications from a
female caught in the Philippines that contained 3-4 batches of oocytes with different
sizes (Bagarinao 1991).
In captivity, female milkfish at the Southeast Asian Fisheries Development
Centre/Aquaculture Department (SEAFDEC/AQD) can spawn to a maximum of
three times per year naturally (Marte et al. 1986a). Spawning occurs around midnight
and this was confirmed based on the egg development stages that were collected in
the Panay island in the Philippines (Bagarinao 1991), although spawning at day time
can occur less frequently. Spawning behavior of milkfish was observed in floating
cages at the SEAFDEC/AQD in Philippines whereby an increase in swimming
activity, chasing, leaping and slapping of water is seen from afternoon to early
evening.
Milkfish spawning was assessed in the Philippines in Moon phases whereby the
spawning occurred around midnight of every 1st and 3rd quarter Moon phase
(Bagarinao 1991). It was also suggested by Delsman in 1929 that, spawning during
neap tide minimize flushing of the eggs and facilitate larvae to remain in near shore
waters (Bagarinao 1991), and the same was also suggested by Johannes (1978) for
marine species in the tropical region with pelagic eggs. The young milkfish larvae
are more abundant in quarter Moon phases and older ones on the New and Full
Moon phases while fry were more abundant during New Moon and Full Moon of
10
which the same occurrence happened in the Philippines and other localities
(Bagarinao 1991; Kumagai 1984).
1.6.2 Seasonality of fry
Many authors have determined seasonality and the occurrence of milkfish fry in
different countries. Villaluz (1986) stated that milkfish fry seasonal occurrence may
be predictable but the abundance will vary from year to year. The season is long near
the equator and becomes shorter at the higher latitudes and in regions affected by
trade winds or monsoon. The peak season may coincide with one or both of the
yearly wind shifts (Villaluz 1986).The seasonality of milkfish fry has been
determined for Fiji with a seasonal peak from December to February (Villaluz 1990)
and for Kiribati with two (2) seasonal peaks, one from late December to early
February and the other from May to September (Wainwright 1982). The
identification of peak seasons is a preliminary step necessary to develop milkfish fry
collections for aquaculture and has been done in Fiji and Kiribati (Bagarinao 1994a;
Billings et al. 2010). In Hawaii, a study conducted on the gonadosomatic index (GSI)
of milkfish over 36 months has indicated that breeding season is between June and
August with a synchronized spawning behaviour observed (Kuo et al. 1979). Among
the thirteen countries for which the seasonality of milkfish fry was established
(Figure 1.4), Fiji is the only one that lies along the same latitude of Vanuatu and thus
may help to predict the fry occurrence.
Figure 1.3 A graphicoccurrence in several(1986) and Vuto et al
1.7 Migration and m
The fry swimming ve
Bagarinao (1991) h
however, they becom
and distribute evenly
11
cal representation of peak and lean season of l countries with data retrieved from Kuo et al. (2014).
movement of fry
elocity is within the range of 9-11 cm/s (Kaw
has stated that milkfish fry near shore wat
me loosely aggregated. The fry arrive in batche
y along extensive sections of beach. There is
f milkfish fry al. (1979),Villaluz
wamura et al. 1984).
ters do not school;
es at different times
s a need for further
12
studies to determine mechanism of larval transport to understand the natural
recruitment of fry along inshore habitat (Bagarinao 1991). Some parameters that
influence the abundance of fry along the coast were discussed by Kumagai (1984)
such as currents, tides, bottom profile and proximity to inland waters whereby
currents and tides were the factors that influence most the abundance along the
shoreline. Fry are more abundant at flood tide due to the effects of the strong current
and collected in large numbers at the surf zone (Bagarinao 1991). The time of day
and wind speed does not have any influence on the fry abundance (Bagarinao 1991).
The arrival of fry at the shoreline is in the form of small batches and this movement
was suggested to be passive, driven by physical factors as such as longshore current
(Kumagai 1984; Villaluz 1986). Although Buri & Kawamura (1983) claimed that fry
in inshore waters were actively migrating into coastal habitat, Kawamura (1984)
suggested that migration was also passive towards nursery habitat. More recently,
Bagarinao (1994a) stated that, milkfish fry move inshore by both passive advection
and active migration.
1.8 Predation on milkfish fry
The most important predators of milkfish larvae and fry are the Hawaiian tenpounder
Elops hawaiensis and the Indo-Pacific tarpon, Megalops cyprinoides (Buri 1980;
Pickering et al. 2012). Other common predators of milkfish are Ambassis sp.,
Terapon sp., Epinephelus sp., Lutjanus sp., Sphyraena sp., Chaetodon sp.,
Meiacanthus grammistes, Oxyurichthys microlepis, and Scatophagus argus (Buri
1980). Bagarinao (1991) suggested that milkfish fry may be eaten also by mugilids
and siganids. The fry of tarpon and tenpounder are very similar to milkfish but they
are predators of milkfish fry and fingerlings. These two species can be distinguished
from milkfish as they have longer and wider bodies and much larger mouth, different
swimming movement and light amber colour (Villaluz 1986).
1.9 Overview of main methods of fry collection
13
Milkfish fry are well-captured using gears operated by filtering water, such as
plankton nets. Milkfish fry have well developed vision so they may respond
negatively (i.e. avoidance) if they recognize the gear or positively (driven) with the
gear if they see the net as some form of shelter (Bagarinao 1991). Black nets were
observed to be more effective in catching fry although white nets are preferred due to
easy identification of fry during capture (Kawamura et al. 1980). Experiments in
tanks on the behavior of the fry response to gear revealed that, milkfish fry gear with
herding design will function well rather than those with filtering (Kawamura et al.
1980). The good condition of the fry is identified through their ability to swim in one
direction facing the current (Villaluz 1984).
The selection of a more appropriate and effective method to collect milkfish larvae
or fry from the wild depends also on the topography of the site, currents, wind
direction and tidal fluctuation (Villaluz 1986). However, the effectiveness of the
methods can also be affected by small population size as shown in small lagoons
with small population (Buri 1980).The fry are caught better with driving gears than
filtering, and the gears may perform well in shallow part of the fry ground. It has
been reported that milkfish fry used to congregate close to the fish shelter located
offshore where they can be caught in sizable quantities (Villaluz 1986). The fry
always stay at the surface, even when they are scared so the depth of the catching
gear will not have to be increased to catch more fry (Villaluz 1986; Villaluz et al.
1983b). The gears used in fry collection are fine nylon nets with a mesh size ranging
from 0.3 to 1.6 mm. The bulldozer net method is a very useful fry collection gear,
which is normally operated in shallow lagoon. The bulldozer net gear was use first in
the Philippines and it was reported to be effective in the night (Villaluz 1986).
However, more recently, Vuto et al. (2014) showed that there is no significant
difference in density of fry between the day and night.
Milkfish fry swim at the low speed of 9-11 cm/s (Kawamura 1984). They are caught
in large numbers with juveniles and also the larvae of other predatory species
(Bagarinao 1991). Sampling methods used in different countries are selected based
on the topography of the sites, such as skimming net, hand scoop net, seine net
collector, beach drag seine net, push net or bulldozer, barrier net and plankton net
(Garcia 1990; Pickering et al. 2012). A skimming net is used in mud flats and
14
mangrove areas, and the hand scoop net is used mainly in mudflats, pools and canals.
The seine net collector is used mainly in mud flats. A beach drag seine net is used
mainly in deep waters of the shoreline. The barrier net is used mainly in estuaries
with bridge as deploy platform for high and low water influx (Villaluz 1984).
Plankton nets are used in deep waters and also on the bridge as a barrier for trapping
fry during high tide (Kawamura 1984). The best design for fry collection in sandy
beaches is the bulldozer net whereby one person will push the dozer net while the
other person will check the back of the net for fry using a plastic basin. The fine
mesh size net of 30 μm is used for collecting the fry constructed onto a bamboo
frame. The separation of the fry will be made in a white basin whereby milkfish fry
will be spotted as having two spot on the eyes, a fork tail, and a black spot on the
middle and the characteristic is of a darting swimming behavior into the current
when the water swirled (Pickering et al. 2012).
1.10 Food demand in the Pacific island countries
In the PICs and Territories, fish consumption is common and a major source of
protein food as many of the villages are located along the coasts. The average fish
consumption per capita in PICs and Territories is 16 -18 kg per person per year (Bell
et al. 2009). The decrease of protein food supply from the oceans is an ongoing
problem all around the world due to increased demand from the growing human
population, and aquaculture is deemed to be an efficient approach to address food
demand in the coming future (World Bank 2013). Capture fisheries have so far
contributed to food production, but unfortunately they are unable to supply the
growing demand as more people request fish and fish products (Tidwell et al. 2001).
Aquaculture alone has accounted for 47 % of fish supply as capture fisheries leveled
off in the last twenty years (FAO 2016) . Overfishing plays a major role in the
decline of coastal ecosystems (Myers et al. 2003) and climate change may play a
significant part as well (Williams et al. 2010). Research into aquaculture is
increasing globally to identify new approaches that may support food security and
sustain community livelihood through sustainable production. Different marine
species are studied which increases chances for potential later application in
15
aquaculture. Milkfish is one of the marine species farmed previously as traditional
industry in Asian countries where annual restocking of fingerlings was reared from
wild caught fry (Nelson 2007). The culture of milkfish was reported to spread to
some PICs and Territories, and the potential is recognized to address food demand
(Pickering et al. 2011). In Vanuatu, milkfish forms an extremely small component of
substance diet when compared to its multispecies fisheries. There has been a history
of collecting seasonal shoals of milkfish fry and juveniles for food (Pickering et al.
2012). In the Solomon Islands, milkfish aquaculture is based on fry collection from
the wild, and the economic analysis done by Sulu 2016 has confirmed that
subsistence farming is possible if labor cost is zero (Sulu et al. 2016). As such,
milkfish subsistence farming may be predictable also to contribute to subsistence diet
in Vanuatu. Moreover, reef fish is becoming very expensive in Efate, which is the
most populous island in Vanuatu and where the capital of Vanuatu is located,
therefore, if lower income households were able to culture milkfish this could
improve both their food security and culture options.
The Vanuatu government has recognized milkfish as a priority species for culture in
the Aquaculture Action Plan (SPC 2008), but no attempt has been made yet on
culturing the milkfish Chanos chanos species. In some PICTs such as Fiji, Kiribati
and Solomon Island attempted were made by collecting the fry from the wild for
culture based on the knowledge of the seasonality (Billings et al. 2010; Vuto et al.
2014; Wainwright 1982). The Australian Centre for International Agricultural
Research (ACIAR) funded project FIS/2012/076 – Improving community based-
aquaculture in Fiji, Kiribati, Samoa and Vanuatu, is now supporting the government
action plan. In this master’s thesis the seasonality of milkfish fry will be determined
as part of the plan for Vanuatu. Upon completion of this, the Vanuatu government
will have at disposal useful information for further work, such as trial culturing and
assessing the economic viability for milkfish culture. The knowledge of milkfish fry
seasonality is the primary basis of the aquaculture process where there is a lack of
advanced breeding technology, such as in the PICs and Territories. There has been a
history of collecting seasonal shoals of milkfish fry and juveniles for food (Pickering
et al. 2012) but to date there has been no attempt to quantify milkfish fry seasonality
in Vanuatu. Hence, the knowledge of milkfish fry seasonality and abundance is
important for further engagement in milkfish farming in Vanuatu.
16
1.11 Conclusion
The literature review has re-affirmed importance of quantifying the seasonality of
milkfish fry in Vanuatu in order to establish baseline information for fry collection,
which is a prerequisite for milkfish aquaculture development. Environmental
variables can also affect the fry seasonality and spawning time and density.
Temperature is known to be a variable that influences milkfish spawning. Milkfish
fry seasonality is latitude dependant with two peaks close to the equator and one
peak at higher latitudes. This chapter provided some knowledge of milkfish studies
in the past that will help to establish the seasonality study in Vanuatu. The methods
of fry collection, time of spawning, and other variables that influence fry appearance
and abundance were highlighted in the review. The methods for fry collection were
reviewed and the best method based on the topography of the sites will be adopted in
this study.
1.12 Research objectives
The main aim of this research is to determine the availability of milkfish fry and their
seasonality and abundance in a coastal area of Efate Island, Vanuatu.
The following is a list of questions that this study is addressing:
1. Is milkfish present in the selected study areas on Efate Island, Vanuatu?
2. What is the seasonality and abundance of milkfish fry in the selected study
area on the coast of Efate Island, Vanuatu?
3. What is the age of milkfish fry at the coast of Efate, Vanuatu?
There are three hypotheses to be tested:
1. Fry abundance in Vanuatu shows a lunar and annual seasonality
2. Environmental factors such as temperature, cloud cover, rainfall, wind speed,
current speed, and turbidity are correlated with fry abundance, length and
weight.
17
3. Fry abundance in the New Moon, 1st quarter Moon, Full Moon and 3rd quarter
Moon are significantly different.
The following is a list of objectives to answer the questions raised above.
1. To determine the presence, seasonality and abundance of milkfish fry at different
sites related to the Moon phase in the coastal zones of Efate Island.
2. To estimate the age at which fry appear along the coast of Efate Island.
3. To determine the impact of environmental variables on fry abundance, length and
weight.
18
Chapter 2: Methodology 2.0 Site selection and description
Efate Island is Vanuatu's third largest island and it is where the capital, Port Vila is
situated (Figure 2.1). Efate Island has the highest population due to migration of
people from the outer island in Vanuatu to Port Vila town for employment and
education (Amos 2007). A site inspection was conducted around Efate Island to
check for the eligible environmental characteristic such as presence of rivers,
lagoons, sandy shoreline, coral reefs, mangrove swamps and brackish shoreline for
fry occurrence. The Coast of Efate Island has bays and lagoons with mangrove
swamps and rivers. The fringing reefs are narrow and sandy beach is present in the
shoreline, as observed during site inspection. The river inputs into the lagoons create
turbid in shore water, which is preferred for milkfish fry recruitments. The four
closest sites to Port Vila were selected.
The four sites selected for the study were Erakor, Kawenu, Mele and Teouma
(Figure 2.2). Their coordinates according to Google Earth are as follows: Erakor:
17°46'54.04"S, 168°19'30.52"E, Kawenu: 17°43'49.30"S, 168°18'18.50"E, Mele:
17°41'31.02"S, 168°15'58.24"E, Teouma: 17°47'27.14"S 168°22'49.73"E. All the
four sites have input of freshwater from rivers and lakes into the coastal shorelines,
with Teouma having two rivers flushing into the lagoon, one from the East and the
other from the West. The similarities between sites were the coastal sandy beaches,
brackish water and coral reef within the lagoon. The differences were the size of the
lagoon, the source of fresh water (river and swamp), the presence and size of
mangrove canopy and the distance of the sites relative to Port Vila town (Figure 2.2).
19
Figure 2.1 Map of Vanuatu where the study is based. Retrieved from http://www.nationsonline.org/oneworld/map/vanuatu-map
20
Figure 2.2 Map of the four sites Erakor, Kawenu, Mele and Teouma where milkfish fry was assessed during the preliminary study, and the Vanuatu capital (marked with a star).
2.1 Sampling materials and methods
2.1.1 Sampling materials
A Secchi disk to test water turbidity was made from timber, and the rope on which
the disk was mounted was marked up to 100 cm with ±0.5 cm precision. A Kedida
concentration tester (CT) -3080 pen type salt and temperature meter were used to
measure salt concentration in mg/l with precision of ± 1 mg/l and temperature in
degree Celsius (°C) with ± 0.1 °C precision. A thermco water bath thermometer of ±
0.5 °C precision was used later to measure temperature during continuous sampling.
21
A plastic bottle, stop watch and a nylon rope of 5 m were used to measure current
speed along the surf zone of the coastline by deploying a plastic bottle and letting it
drift along a 5 m distance while taking the time with a stop watch. Additional
environmental parameters such as rainfall, cloud cover and wind speed were
obtained from the Vanuatu meteorology and Geo-hazard department (VMDG).
2.1.2 Bulldozer net construction
The bulldozer net was built from bamboo, and tire rubber was used to tie the bamboo
sticks together (Figure 2.3). The net was seamed with a 100 μm fishing line onto the
nylon rope to keep the edges from wearing and also to retain the shape. A wire was
used to attach the net at each corner of the frame and a fishing line was used to tire
the net into position. The pouch was a fine mesh size of 150 μm while the wings of
the net were made of a larger nylon mesh size of 250 μm.
The diagram of the frame with the measurements is shown in figure 2.4. The length
of the bamboo frame was 2.6 m and the back of the frame was narrow to allow for
easy pushing and collection of the fry. The front was wide which was used for the
wing controlling the fry.
The diagram of the net and measurement are shown in figure 2.5. The pouch was
short and covered about 1/3 of the total length. The back of the net was deep to
enable fetching when hauled to check for milkfish fry (Figure 2.5).
22
Figure 2.3 The bulldozer net built and used for this study. Picture shot at Teouma on March 2016.
Figure 2.4 The bamboo frame design for the bulldozer net (Pickering et al. 2012)
23
Figure 2.5 Net size and design modified from Pickering et al. (2012)
2.1.3 Sampling method
The net was first tested for its effectiveness and efficiency at the Kawenu lagoon
on the side towards Port Vila. The first deployment showed the catching of a
juvenile grouper after three trials pushing over 25 m along the surf zone.
Sampling trips were made once in every Moon phase for one-year period.
Overall, 69 sampling trips were made during the period of this study from
February 17th 2016 to February 15th , 2017 of which 18 trips were made for the
preliminary site assessment. A total of 7 to 8 net deployments were made per
sampling trip whereby each deployment covers a push over 25 m distance at a
time of 5-7 minutes. Each deployment is referred to as effort whereby the net was
checked for the fry presence for each deployment.
Fry abundance was checked in the surf zone. Net deployment was done only
during daylight hours between 6 am to 6 pm during the 1st and 2nd hour of high
tide. The abundance of larvae is expressed as number collected per effort
(Morioka et al. 1993) where the effort is the number of bulldozer net
deployments.
24
2.2 Fry identification
Milkfish fry identification was made visually with the help of a white plastic
basin on the field. Milkfish fry (Figure 2.6) is characterized by lack of
pigmentation, two dark spots (the eyes) on each side of the head, a dark spot (the
air bladder) on the middle of the body and also a light grey spot on the caudal
fin(Pickering et al. 2012). The fry uses a darting swimming motion with the
tendency to swim against the current.
Figure 2.6 Milkfish fry collected in a white plastic basin for visual identification. The black dot from left to right showed position of the grey spot on the caudal fin, dark spot in middle of body (air bladder) and two dark spot (eyes) on each side of the head.
2.3 Fry length and weight measurement
After identification, the larvae were dipped into 95% ethanol for 10 min to
enhance hardness for measuring and weighing. After 10 min, the TL of the larvae
was measured with a ruler and expressed in millimeters and weighed in grams
with an electronic balance (FX-1200 SN1010061) with a scale of 3 decimal
places and precision of 0.1 g. After mass and length measurements were done,
25
each larva was transferred into small containers of 20 ml in 95 % ethanol until
further use (Miller et al. 1994).
2.4 Otolith analysis
Before otolith analysis, a maximum of 5 fry from each sample collected in each
month was measured and TL measures were converted to equivalent fresh-state
using the equation from Morioka et al. (1993)on milkfish length conversion
applying the isopropanol equation. The equation for isopropanol was used as
there was none for ethanol available. The isopropanol equation employed was,
(Equation 2):
Equation 2:
Where TL is the total length of fresh specimen and Li is the length of the
specimen in isopropanol (Morioka, Ohno et al. 1993).
Examination of otolith was made on the sagittae on increment of Teouma wild
caught fry excluding the ones from Erakor, Kawenu and Mele, as the seasonality
study only focused on one site. A total of 91 fry was randomly selected from the
samples collected in Teouma for the examination.
The sagittae were removed from both sides of the milkfish fry cranium by teasing
them with an insulin syringe. Each sagitta was then mounted onto microscope
slides and immersion oil was used for otolith observation. Count of the total
number of increments was done under an Olympus dissecting light microscope
SZ-ST type of magnification ranging from 100 to 500x. The photos were taken
with an Infinity 1 canon camera (A-1) connected to the microscope and counts
were done twice, three days apart, by using the same photographs. A third count
was done under the microscope. The three counts were validated using analysis
of variances (ANOVA) single factor in excel and only the counts from the
microscope were used in the age estimate.
The age of milkfish was determined from the reading of otoliths. In milkfish the
first increment is formed two days after hatching during yolk sac re-absorption
period and the increments continue to grow daily regardless of the growth rate
26
(Bagarinao 1991). The relationship between the number of increments and the
age in days is the following (Equation 3):
Equation 3: D = N + 1
Where N = number of increments and D = Days from hatching
The equation, developed by Bagarinao (1991) was used to backdate the time of
hatching and thus the date of spawning was calculated from the counting done
under the microscope.
2.5 Data analysis
The comparison of milkfish fry abundance in the different sites, months and
Moon phases were determined using ANOVA in R version 3.4.0 (R Core Team
2007) with a significance level of 0.05. The difference in weight and length
between Moon phases was also determined by the same test. The fry seasonality
data was presented from August 2016 to January 2017 and from February 2016
to July 2016 to better depict the seasonal occurrence of milkfish fry.
The environmental data collected during sampling and those collected on
sampling day by VMDG were analyzed to evaluate their effect on the milkfish
fry abundance, length and weight. Regression analysis with multiple predictors,
also known as multiple regressions, was run and model selection was made by
comparing the most complex model with simpler ones in R for all variable data
such as sea surface temperature measured in situ, cloud cover, rainfall, turbidity
and wind speed. The Pearson correlation test for statistical software version 10
(Statistica Inc 2011) was conducted to verify the correlation coefficient and
significance level at 95% significance level and then used as final. Lastly the
correlation between age, length and weight of fry was done in Microsoft excel
version 15 (Microsoft Officer 2013).
27
Chapter 3: Results
A total of 47 fry were captured in the four sites altogether for the site assessment,
with 369 fry collected in the Teouma bay during a one year period and were used for
the following analysis except for aging, which was done on a subsample of 91 fry.
3.1 Site assessment and selection
Results for the four environmental parameters (i.e. turbidity, current speed, salinity,
and temperature) assessed over a month in Erakor, Kawenu, Mele and Teouma are
summarized in table 3.1. Erakor, Kawenu, Mele and Teouma, had a mean turbidity
of 80 ± 12.2 cm, 75.8 ± 10.3 cm, 89.3 ± 4.0 cm and 69.5 ± 11.2 cm respectively.
During the first 1-2 hour of high tide, Teouma had a mean current speed of 0.01 ±
0.0 m/s and Mele, Kawenu and Erakor of 0.01 ± 0.1 m/s. In ascending order
Teouma, Kawenu, Erakor, and Mele had mean salinity of 8.4 ± 2.5 parts per
thousand (ppt), 14.1 ± 3.0 ppt, 14.5 ± 3.5 ppt and 15.3 ± 3.5 ppt respectively. The
mean monthly sea surface temperature (SST) measured showed Mele had the lowest
(30.4 ± 1.7 C), while Teouma, Kawenu and Erakor had a higher temperature (30.7 ±
0.5 C, 31.1 ± 3.5 C and 31.3 ± 2.0 C, respectively).
Table 3.1 The monthly environmental parameters (turbidity (transparency), current speed, salinity and SST) assessed and the landscape features for Mele, Kawenu, Erakor and Teouma. The monthly values are presented as mean ± standard deviation.
Sites
Landscape
features
Turbidity
(transparency)
(cm)
Current
speed
(m/sec)
Salinity
(ppt)
SST
(ºC)
Mele Sandy beach,
coral reef, two
rivers
89.3 ± 4.0 0.1 ± 0.1 15.3 ± 3.5
30.4 ± 1.7
Kawenu Sandy beach,
coral reef, spring
75.8 ± 10.3 0.1 ± 0.1 14.1 ± 3.0 31.1 ± 3.5
28
water, mangrove
habitat
Erakor Sandy beach,
coral reef, lagoon,
swamps
80.0 ± 12.2 0.1 ± 0.1
14.5 ± 3.5
31.3 ± 2.0
Teouma Sandy beach,
coral reefs, two
rivers
69.5 ± 11.2 0.1 ± 0.0 8.4 ± 2.5 30.7 ± 0.5
Milkfish fry appeared in the four sites with the least abundance recorded in Erakor,
Kawenu and Mele (1, 2, and 1 fry, respectively) and was significantly different (p =
0.01) from those in Teouma with the highest (43 fry) (Figure 3.1). The lowest
number of fry at Erakor, Mele and Kawenu were not significantly (p > 0.99) different
from each other.
The study proceeded only in Teouma, based on low numbers of abundance in the
other sites. Thus, the following results refer only to Teouma.
29
Figure 3.1 Milkfish fry total abundance (number of fry captured during the exploratory trial done on Erakor, Kawenu, Mele and Teouma). The different letters indicate significant difference in abundance (p < 0.05) while the same letter indicates no significant difference (p > 0.05).
3.2 Milkfish fry occurrence in Teouma 3.2.1 Monthly occurrence and abundance
Milkfish fry occurrence in Teouma started in October and ended in early May
(Figure 3.2). The month of November and April showed the two highest peaks in
abundance (120 and 92 fry respectively) with no significant difference where p =
0.67 (Appendix 1). In the months of October, December, January, February, March
and May the abundance was low (19, 24, 3, 55, 41 and 15 fry, respectively) and
significantly different from those November (92) and April (120) where p < 0.05
(Appendix 1). The months of June, July, August and September showed no
occurrence of fry, but were not statistically different (p > 0.05) from the months with
low abundance October, December, January, February, March and May except for
July and February which were significantly different to each other where p = 0.03
(Appendix 1). The sea surface temperature (SST) collected from both Meteorology
department and in situ shows milkfish fry occurrence at monthly mean SST of 26℃
and above while no occurrence is observed at mean SST below 26℃.
30
Figure 3.2 Milkfish fry abundance and occurrence with SST by month for Teouma. The different letters indicate significant difference in fry abundance (p < 0.05).
3.2.2 Milkfish fry length and weight
The mean total length (TL) of the collected fry was 12.8 ± 1.2 mm (Appendix 3).
The Moreover, 59 % of the fry had length from 13 to 14 mm and 33 % had the length
between 11 and 12 mm (Figure 3.3). The minimum length recorded was 8 mm and
the maximum was 15 mm.
The weight of the milkfish fry ranged from 3 mg to 9 mg, with an average weight of
5.8 ± 1.6 mg (Figure 3.4, Appendix 3). A total of 65 % of fry in Teouma had a
weight range between 5 mg and 7 mg while 17 % of the fry caught had a weight of 8
mg and 9 mg.
The fry TL by months shows that May had the highest mean TL (Figure 3.5, A) and
is significantly different in April and October (p = 0.04 and p = 0.04 respectively)
with the lowest mean
sampling shows no st
Figure 3.3The total le
Figure 3.4 The frequ
31
n TL. The comparison of weight by month
tatistical significance (p > 0.05) with the weig
ength frequency of milkfish fry (N = 369) in th
ency distribution of milkfish fry (N = 369) we
h (Figure 3.5, B) of
ght of fry collected.
the Teouma
eight in Teouma
Figure 3.5 Milkfish frThe different letters inrepresent the standar
3.3 Effect of Moon p
3.3.1 Abundance by
The distribution of cu
showed that the highe
followed by 3rd quarte
the two (Figure 3.6).
Moon (N = 57 and 34
abundance in New M
in the 1st quarter and
0.12).
32
fry mean total length (A) and weight (B) per sndicate significant difference (p<0.05). The e
rd deviation of the mean.
phases
Moon phase
umulative fry abundance at the four consecuti
est fry abundance (N = 160) was observed du
er (N = 120), with no significant difference (p
Milkfish fry abundance was lower in the 1st
4 fry, respectively), and were significantly dif
Moon (p = 0.001 and p = 0.001 respectively). T
3rd quarter Moon phase were not significantl
sampling month. error bars
ive Moon phases
uring New Moon,
p = 0.44) between
quarter and Full
fferent from the
The fry abundance
ly different (p =
33
Figure 3.6 Milkfish fry cumulative abundance in all Moon phases for Teouma
throughout the year. The different letters indicate significant difference is the
abundance between Moon phases (p < 0.05).
3.3.2 Total length and weight in Moon phase over one year
The mean total length of the fry captured at 1st quarter, Full Moon and 3rd quarter
(13.1 ± 0.9 mm, 13.5 ± 1.0 mm and 13.1 ± 1.2 mm, respectively) were not
significantly different (p > 0.05) from each other (Figure 3.7.A, Appendix 1). The fry
in 1st, 3rd and Full Moon had a mean total length higher than the fry captured at New
Moon (12.2 ± 1.2 mm) and were significantly different (p < 0.05) from each other
(Figure 3.7).
The fry captured in the 1st quarter Moon phase had the lowest mean weight (4.7 ± 1.5
mg) followed by the Full Moon (4.9 ± 1.8 mg) and not significantly different (p =
0.61) from each other (Figure 3.7, B). In the 3rd quarter and New Moon the fry had
the highest mean total weight (5.3 ± 1.7 mg and 5.4 ± 1.6 mg, respectively) and these
were not significantly different (p = 0.10) from each other. The fry sampled at the
Full Moon had a mean weight not significantly different (p > 0.05) from the other
Moon phases (Appendix 1).
Figure 3.7 Milkfish frThe different letters s0.05) and the error ba
3.4 Comparison of f 3.4.1 Environmental
The environmental va
and current speed (ha
the coastal shoreline a
However, cloud cove
0.05) whereby fry abu
34
fry mean total length (A) and weight (B) in foushow significant difference in mean total lengars represent the standard deviation of the m
fry abundance with other indices
l variables correlation with fry abundance
ariables assessed (temperature, rainfall, turbid
ad no significant influence on the milkfish fry
as shown by the low correlation values (p > 0
er had a significant influence on the milkfish f
undance increased with cloud cover.
ur Moon phases. gth or weight (p < mean.
e
dity, wind speed
y abundance along
0.05) (Figure 3.8).
fry abundance (p <
Figure 3.8 The envirotemperature, turbidityabundance.
3.4.2 Environmental Milkfish fry length sh
cover, rainfall and tur
speed showed a signi
respectively) with fry
35
onmental variables, rainfall, cloud cover, winy (transparency) and current speed, correlate
l variables correlation with length and wei
howed non-significant correlations with temp
rbidity (p > 0.05) (Figure 3.9). The wind spee
ficant (p < 0.05) positive correlation (r = 0.24
y length.
nd speed, ed with milkfish fry
ight of milkfish fry
perature, cloud
ed and current
4 and r = 0.21,
Figure 3.9 The enviro(transparency), wind length
Milkfish fry weight s
cloud cover, wind spe
= 0.02) positive corre
transparency increase
36
onmental variables, temperature, cloud coverspeed and current speed correlation with mi
howed non-significant (p > 0.05) correlations
eed and current speed (Figure 3.10). There w
elation in milkfish fry weight with rainfall and
e.
r, rainfall, turbidity lkfish fry total
s with temperature,
as a significant (p
d turbidity as
37
Figure 3.10 The environmental variables, temperature, cloud cover, rainfall, turbidity (transparency), wind speed and current speed, correlation to milkfish fry weight
3.5 Otolith analysis and age of the fry per month and spawning estimates
The age of the fry was assessed by measuring the number of increments in the
otoliths. In figure, 3.11 are shown examples of this assessment. In more detail, the
increment of a young larvae of 14 days old (Figure 3.11, a) has wider rings indicated
with a series of black dots while older larvae of 22 days (Figure 3.11, c) have narrow
increments. The star at the center of otolith indicates the core which was formed
before the first increment and the discontinuity zone shown by the light zones
indicated with the bar (Figure 3.11, a). There are four dimensions on the otolith
sagittae (Figure 3.11, a) of a milkfish fry, the posterior, ventral, dorsal and the
anterior. The 3 photos (Figure 3.11, a, b, c) indicate a typical 14 days, 15 days and a
22 day old milkfish fry otolith increments.
38
Figure 3.11 Photographs a, sagittae otolith from 14 days old larvae, 11 mm total length (TL). (x400), b is an otolith of a 15 day larva of 11 mm TL and, c is an otolith of a 22 day larva of 14 mm TL. In photograph a, letters. A, anterior; P, posterior; D, dorsal; V, ventral. The dots (●) are the increment dark zone, the star ( ) indicates the core and the arrow (↓) indicate the discontinuity zone.
3.6 The age distribution
The milkfish fry at Teouma coast were of age that ranged from 13 days to 25 days
old (Figure 3.12). The fry age that ranged from16 days to 18 days accounted for 57
% of the total fry caught while 29 % of the fry were aged between 19 to 22 days.
39
Figure 3. 12 The age frequency of milkfish fry in Teouma coastal shoreline
The age distribution between months revealed by otolith analysis two distinct age
groups, the first from October to December and the second was from February to
May (no otolith analysis was performed with samples from January) (Figure 3.13).
The oldest fry appears in March, April and May.
Figure 3.13 The mean age of fry per month of occurrence in Teouma. The error bars represent the standard deviation of the mean.
40
The relationship between total length and age shown in figure 3.14 indicates that the
length was directly proportional to the age where r = 0.46. Thus length also increases
with age at occurrence of fry and were significantly correlated (p < 0.05).
Figure 3.14 Correlation between the age and length of milkfish fry 3.7 Spawning time estimates for Teouma over one year period The backdating of the otolith data for each sampling trip revealed that milkfish-
spawning season starts in September and ends in April (Figure 3.15). Moreover,
there were two spawning groups or seasons, one from September to November and
the other from January to April. Milkfish spawning does not occur in December,
May, June, July and August. It is observed that milkfish spawning activity in
Teouma is link to the mean Monthly SST of above 25 ℃, and is the predicted
spawning threshold (Figure 3.15).
41
Figure 3.15 The spawning season of milkfish fry (N =63) based on different days over a one year period with mean Monthly SST
3.8 Comparison of fry abundance in the Melanesian PICs
The fry abundance in Teouma for one day over 1 hour of sampling using
bulldozer net is very low (0-79 fry) compared to the fry abundance reported in
Fiji and the Solomon island with 200-100 and 0-1000 respectively over one day
of sampling (Table 3.2).
Table 3.2 The fry abundance between Fiji, Solomon Islands and Vanuatu
42
Chapter 4: Discussion
The main goals of this study were to identify milkfish fry seasonality, abundance,
and spawning season, and to identify the environmental parameters such as Moon
phases, cloud cover, current speed, rainfall, turbidity, temperature and wind speed for
their effects on milkfish appearance and abundance in Vanuatu. The knowledge of
fry seasonality is crucial for Vanuatu, as hatcheries are too expensive to operate due
to the high cost of induced spawning technology, bloodstock hatchery, and
management. Similarly, according to FitzGerald (2004), the high cost of capital
investment along with investment on technically trained staff will make the
development of hatchery not advisable in the initial stage of milkfish aquaculture.
Therefore, the study can aid policy implementation based on this information for
milkfish culture purposes in Vanuatu.
Initially, four sites close to Port Vila on Efate Island, namely Erakor, Kawenu, Mele,
and Teouma were selected for a preliminary assessment. The sites were selected due
to the presence of rivers, freshwater sources, and landscape features such as a lagoon,
sandy beach, coral reef and coastal mangrove forest. The presence of rivers and
lagoon creates favorable environmental conditions known to be important for
milkfish fry appearance (Bagarinao et al. 1987; Bagarinao 1991; Brian 2015). All the
selected sites have similar topographic features of a gentle slope on the surf zone
whereby a bulldozer net method of fry collection was selected as it was used for the
first time in the Philippines in 1983 and was deemed effective along surf zones of
sandy coastal shorelines (Villaluz 1986). The pushes are made parallel to the
coastline facing the direction of the wind. The bulldozer net design used in this study
was a modification from the one used by Pickering et al. (2012). In Vanuatu, the
bulldozer net was never previously used for any purpose and that includes milkfish
fry collection, nevertheless, sampling done with the bulldozer net was proven
efficient in this study and thus its further use is highly recommended in locations
with similar characteristics.
After the completion of one-month assessment in the four sites, it was observed that
milkfish fry was present in all four sites. However, Erakor, Kawenu, and Mele, the
43
sites closest to Port Vila, had the lowest abundance in milkfish fry. Teouma was the
site furthest from Port Vila where the council of chiefs has control of the coastal
resources. Recently they imposed regular seasonal banning of fishing from
November to April as they believe it is the season when much of the spawning
occurs. In Pakistan, a study on the breeding season of Schizothorax plagiostomous
by Jan et al. (2017) using GSI techniques resulted that spawning occurs twice a year,
once in March-April and October-November. Similarly, an evaluation of closed area
for fish stock conservation in Europe by Horwood et al. (1998) revealed that closure
areas were likely to have benefited many commercial and unregulated fisheries
species. In addition, a study of the sustainability of two coral reef fish (Amphiprion
percular and Chaetodon vagabundus) larvae recruitment in a Marine Protected Area
(MPA) in Papua New Guinea revealed that even small MPAs may be self-sustaining
in larvae recruitment from the adult spawning in the same location (Berumen et al.
2012). This may be the same case for the seasonal closure in Teouma where adult
milkfish around the coast will only sustain fry recruitment along the Teouma
shoreline. However, this needs to be verified with a more comprehensive research. In
addition, the presence of two rivers within the lagoon in Teouma also makes a good
fry collection ground. This is similar to what was found by Sulu et al. (2016) in the
Solomon Island that better fry collection grounds are usually located close to river
mouths and swamp outlets. Furthermore, Bagarinao (1991) has stated that the most
important component of the water chemistry is the muddy smell, composed of
organic compounds called geosmin which is formed from the bacterial breakdown of
detritus and benthic zooplanktons that triggers most of the fry appearance to the
coastline to feed where conditions are favorable.
The physical parameters of turbidity, current speed, salinity, and temperature, were
measured at sampling during the site assessment and some differences between the
sites were recorded, that may aid to explain the differences in milkfish fry
abundance. It was evident in Teouma that physical parameters such as temperature,
current speed, turbidity, and salinity do not fluctuate when compared to the other
three sites Erakor, Kawenu, and Mele. The stability of the environmental conditions
is crucial for milkfish larvae survival, similarly, the high survival rate of milkfish
44
juvenile was observed by Jaspe et al. (2012) in the pond when there is enough food
supply and water quality is stable.
Milkfish fry appearance in Teouma started in October and continued to occur until
early May which is similar to their occurrence in Fiji from September to May
(Pickering et al. 2012) and has two peaks which are similar as in the Solomon island
(Sulu et al. 2016). A study on the gray mullet Mugil cephalus fry seasonality in
South Africa by Bok (1979) revealed that recruitment along the coastline occurs
from May to November and the peak period of occurrence took place from July to
October. The seasonality of Mugil cephalus in South Africa is different to milkfish
fry occurrence in Vanuatu. In India, Sarojini (1951) has determined that the fry of
Mugil tade and Mugil parsia usually occur from November to February (Oren 1981).
On the Teouma coast of Efate, milkfish fry recruitment had two peaks of occurrence,
one in November and the other in April (120 and 94 fry respectively) which is
similar to the number of peaks recorded in Kiribati although the peaks were shorter
compared to Kiribati with peaks from May to September and December to April
(Wainwright 1982). However, fry seasonality in Kiribati is similar to those of
Solomon Islands (Sulu et al. 2016), as it occurs throughout the year, which is
different to Fiji and Vanuatu with 8 and 9 months respectively. In October,
December, January, February, March, and May a low number of milkfish fry was
observed in Teouma. The months of October and January were the beginning of the
first and second period of fry seasonal appearance, which later leads to the peak
occurrence in November and April. The reduction in fry abundance in March may be
related to the effects of storm surge due to cyclone Winston which started early in
February in North West of Port Vila in Vanuatu. Similarly, a visual census carried
out in 8 coral patch reef in Lizard Island of great barrier reef during larval seasonal
peak occurrence 12 days before and 2 days after a cyclone resulted in fewer juveniles
recruited into the coral reef due to storm surge (Lassig 1983). The study has
concluded that cyclone has marked effects on the fish assemblage as a whole. In
Teouma, the sampling done 2 days after the week of storm resulted in no fry and
they become available only on the sampling day, which is 10 days after the storm
surge of the cyclone. Therefore, the larvae could have suffered additional mortality,
45
which resulted in a decline of fry abundance in March. No fry was captured in the
months of June, July, August, and September. The close range in length of fry
occurrence in Fiji and Vanuatu may be related to the closeness in latitudinal position
between the two countries as it was stated that countries with similar latitude would
have similar periods of occurrence (Bagarinao 1994b). Milkfish fry in the coastal
shoreline of Teouma occurs mainly in the summer months, which may be the same
for their occurrence reported in Fiji. However, the low abundance observed in this
study could also be related to sampling chance, as there is no significant difference in
abundance in the month of low fry occurrences and those of no fry occurrence. A
different method of sampling is required to avoid errors in sampling chance as in the
Solomon Islands and Fiji more fry was caught using other methods (hand dip net,
scoop net) in narrow, shallow lagoon compared to bulldozer net (Sulu et al. 2016).
By comparing the fry abundance in Teouma to Fiji and Solomon Island’s data, fry
abundance in Teouma (0-79 fry per hour) may not be so different using the bulldozer
if sampling is done for more than one hour per day. In Fiji and Solomon islands, fry
were collected at 200-1000 and 0-1000 fry in a sampling day respectively (Napulan
2012; Sulu et al. 2016).
Milkfish fry TL for Teouma coast ranged between 8 mm and 15 mm. This is similar
to those appeared in Japan and Philippine which ranged from 10 mm to 17 mm
(Kumagai 1984; Senta et al. 1981), however, they are more similar to those occurred
in Solomon island with a length range of 10 mm to 15 mm (Sulu et al. 2016). In the
Philippines, milkfish larvae of less than 9-10 mm TL are present in offshore surface
waters but are driven onshore by wind-driven current and tidal current (Kumagai
1984). The weights of the fry in Teouma ranged from 3 mg to 9 mg and were similar
to those reported in Taiwan by Liao et al. (1977) with the weight range from 3.2 mg
to 11.2 mg. In regards to seasonal abundance, milkfish fry was abundant in summer
compared to winter. The two peaks in fry seasonality were in line with previous
findings (Kumagai 1984), where it was determined that fry seasonality is long near
the equator with two peaks of which April-May is higher and progressively shorter at
high latitudes. The fry seasonality in Teouma occurs at monthly mean sea surface
temperature (SST) above 25℃ (figure 3.2) and most spawning activity also occurred
46
at temperatures above 25℃, similarly, milkfish fry occurrence in Fiji, Solomon
Islands, and Kiribati occurs at the same temperature (Bagarinao 1991; Sulu et al.
2016). Therefore, the SST threshold value for prediction of milkfish fry occurrence is
25℃ and above and is noted in the Philippines (Bagarinao 1991).
The milkfish fry occurrence in the different Moon phases showed a high frequency
in New Moon and 3rd quarter Moon phase (120 and 94 fry respectively). This
pattern observed in Teouma was different to the pattern of fry occurrence determined
in the Philippines whereby spawning occurs in quarter Moon period and fry are most
abundant in Full and New Moon period (Bagarinao 1991; Kumagai 1984). In
addition, in the Solomon Islands, the catch during the New Moon period is higher
than the Full Moon, which is similar to those observed in Teouma but a non-
significant difference was identified (Sulu et al. 2016). It was anticipated by Garcia
(1990) that milkfish fry is more abundant between full Moon and New Moon periods
after observing the seasonality of fry in the Philippines. However, this study carried
out in Vanuatu supports that fry is more abundant in New Moon period than in the
Full moon period. This is different to what is observed in Solomon Island and in the
Philippines. A study conducted by Horký et al. (2006), on the behavior of fish in
Moon phase using pikeperch, in the behavior revealed that New Moon has increased
diurnal activity compared to other Moon phases, similarly as the sampling was done
during the day, more fry was caught in New Moon. The results of this study also
show that smallest fry are more abundant in quarter moon period and otolith analysis
also proved that younger fry occurs in the similar period. Concerning the weight of
the fry in Moon phases, those fry appeared in New Moon have more weight.
According to Horký et al. (2006), fry activities increase in the New moon period.
This study supports that as fry activity increased in New Moon period, feeding could
also increase which result in more weight of fry in New Moon.
The environmental variables assessed (temperature in situ, rainfall, turbidity, wind
speed and current speed) had no significant influence on milkfish fry abundance
along the coastal shoreline of Teouma. The monthly mean SST (figure 3.2, 3.15)
indicates a possible link with fry seasonality although there was no correlation, and
this may need further assessment to validate. Fry were caught only on the months
when monthly mean SST was above 25 ℃ and this could be the threshold for
47
prediction fry occurrence. Similarly, countries like Fiji, Solomon Island and Kiribati
have records of fry occurrence at monthly mean SST above 25℃ (Bagarinao 1991).
However, cloud cover had a significant influence on milkfish fry abundance whereby
fry abundance increased with cloud cover that may reduce fry capability of seeing
and escaping the net. It was also noted by Kawamura (1984) after testing fry
visibility with black and white net mesh that fry escape white net mesh while they
retain more in the black mesh net. Nevertheless, a white color net mesh was used in
this study to ease milkfish fry identification, and as a result, fry tends to be caught in
higher numbers during high cloud cover possibly because net visibility is reduced. In
future studies, a black net should also be tested in Teouma to address the low capture
numbers and evaluate if these are related to the color of the mesh used.
The correlation of environmental variables studied with the length of fry over one
year period revealed that milkfish fry length was not significantly correlated with
temperature, rainfall, turbidity and cloud cover. However, wind speed and current
speed showed a significant correlation whereby larger fry were caught with
increasing wind speed and current speed. The wind speed drives fry into the net by
blowing onto the surface current causing larger fry difficulty to escape the net as the
net was deployed facing the wind direction. It was also reported in the Philippines
that milkfish fry were more abundant in favorable wind direction, particularly when
the wind blows towards the shoreline (Senta et al. 1981; Villaluz et al. 1983b).
The environmental variables, temperature, cloud cover, wind speed and current
speed, showed no significant correlations to milkfish fry weight. However, there was
a significant positive correlation of the fry weight to rainfall and increased turbidity.
Rainfall is important to fry as it increases river flow, causing suspension of detritus
in the coastline where more zooplanktons feed on them. Similarly, Edwards (2001)
has determined that zooplankton feeds on detritus and as more zooplanktons are
available, the fry feeding activity increases and thus the weight increases. In
addition, rainfall increases saturated oxygen concentration (Li et al. 2015), whereby
it was determined by Mallya (2007) using a case study on Atlantic halibut cultured
under various saturated oxygen concentration whereby growth and feed conversion
increase with saturated oxygen.
48
The age of the milkfish fry were assessed by measuring the number of increments
from 92 milkfish larvae. The fry (8-15 mm TL) captured on Teouma coast were of
age ranging from 13 days to 25 days old which is similar to the one determined in the
coast of Japan documented by Kawamura et al. (1984) where milkfish fry within 10 -
17 mm TL were aged between 14 to 29 days. Similarly, the fry result from Teouma
showed that more fry were aged between 16 and 18 days old which is not different to
those determined by Kawamura et al. (1984) whereby the fry length 12.5 -14.0 mm
TL were aged 18 to 21 days old. The result of the study revealed that milkfish fry
spend 2 to 4 weeks before disappearing for the nursery which correspond to the size
at transition stage reported by Bagarinao (1991). The fry length was positively
correlated to age, similarly, (Sackett et al. 2013) have stated that fish length increases
with age.
The spawning season of milkfish fry was determined by backdating of the otolith
data and revealed that milkfish spawning starts in September and ends in April, this
links to the period of fry occurrence mentioned previously in the first section,
whereby fry occurs in October following the spawning activity in late September.
Furthermore, there were two spawning groups or seasons, one from September to
November, and the other from January to April, which relates to the two peaks of fry
occurrence. The peak period of fry occurrence is short in the two periods of
spawning season. Helfman et al. (2009) found that typically pelagic fishes spawn
over a 4 month period with a short period of maximal activity, after determining the
spawning period of Cod in the North Sea of the Southeast coast of England where
spawning occurs between January and May where 70% of eggs produced during 6
weeks period. Milkfish spawning in Vanuatu does not occur in December, May,
June, July, and August. According to the results of the backdating, there is a period
of 44 days lapse between the first phase of spawning and the second phase of
spawning activity. The spawning season of mullet in Australia was studied by Horký
et al. (2006) who determined that Mugil chepalus spawning occurs between March
and September which is different to that of milkfish Chanos chanos in Teouma coast
of Vanuatu. The Mugil cephalus of the Mediterranean shore of Israel spawn from
July to December (El Meseda et al. 2006). It is seen that different species of mullet
spawn at different times of the year based on salinity and geographical location as
determined for Mugil cephalus (El Meseda et al. 2006). Similarly, Martinez et al.
49
(2006), has reported that different localities have different milkfish spawning seasons
whereby spawning can occur more than once during the annual spawning season. In
addition, an oocytes analysis of a hormone-induced trial on milkfish by Lee et al,
(1986) indicated that milkfish can spawn several times during the annual breeding
season (Garcia 1990). The otolith analysis resulted that some months fry appear
onshore were then to be older. The cause of the difference in age maybe related to
many factors.
Otolith analysis revealed that fry caught in November and February had the lowest
mean age. Older fry were caught in two periods, the first was in December and the
second period was in March and April. Further research is needed to determine
factors correlated to differences in fry age between the months of occurrence.
However, by synthesizing data from both seasonality of fry occurrence, age and
spawning period, it may be possible that this data can be used for management
purposes to predict fry seasonal occurrence. The months when occurrence peaks can
be used to determine abundance in other islands and possible fry collection ground
may need to be banned from catching of milkfish during the breeding season.
Therefore, milkfish aquaculture development in Vanuatu can be enhanced with this
knowledge of fry seasonal occurrence.
50
Chapter 5: Conclusion
This study has concluded that milkfish fry is available along the coast of Efate Island
in Vanuatu and it was present in all four sites assessed, though the abundance
recorded was low and thus unlikely to be sufficient to support milkfish aquaculture.
The sites close to Port Vila, Erakor, Kawenu and Mele showed low abundance in
milkfish fry compared to Teouma, which is the furthest site from Port Vila and was
selected for the seasonality study. The season of milkfish fry occurrence on the
Teouma coast of Efate started in October and ended in May, with two peaks of
occurrence identified, one in November and the other in April. The fry occurrence
was correlated with the spawning, which probably occurred from September to
November and from January to April, as inferred from aging analysis. Since the
study was done in the central province of Vanuatu, it is recommended that further
assessment of milkfish fry should be carried out also in other islands where
conditions are favorable for their occurrence.
The abundance of wild fry collected over the period of this study may not be
sufficient to support a subsistence culture of milkfish, as the numbers were quite low.
A comprehensive assessment of the abundance and seasonality is needed across the
Vanuatu groups of islands for the presence of milkfish fry and their occurrence in
order to close the seasonality of occurrence in Vanuatu and other methods of fry
collection may also needed to be tested on the abundance. A thorough study on fry
abundance will help determine whether milkfish culture in Vanuatu can rely on the
collection of fry from the wild. Additionally, the absence of fry in some months
could also be related to sampling chance, as there is no significant difference in fry
abundance between the month of low occurrence and absent. Most of the
environmental variables did not correlate with abundance, length, and weight of
milkfish fry, which may require further research to prove. Concerning Moon phases,
fry were more abundant in the 3rd quarter and New Moon compared to 1st quarter
Moon phase and Full Moon.
The period of fry occurrence is 8 months long, which is similar to that of Fiji.
However, the fry abundance was too low so the aquaculture of milkfish is still a
question that needs further research although the seasonality can be predictable. A
51
nationwide assessment can be done using this study as a pilot for spawning and fry
occurrence. The island of Santo, Malekula and Tanna would be the other sites that
look promising for the assessment, as there is plenty of bay and rivers connected to
the sea. However, it is recommended that after a complete assessment the seasonality
study should be done repeatedly over years to be able to build a complete cycle of
milkfish fry occurrence in Vanuatu.
52
References
Agbayani, R.F., Baliao, D.D., Franco, N.M., Ticar, R.B., and Guanzon, N.G. (1989)
An economic analysis of the modular pond system of milkfish production in
the Philippines. Aquaculture 83(3-4), 249-259.
Amos, M. (2007) Vanuatu fishery resource profiles. Apia, Samoa: Secretariat of the
Pacific Regional Environment Programme.
Arai, R., Nagaiwa, K., and Sawada, Y. (1976) Chromosomes of Chanos chanos
(gonorynchiformes, Chanidae). Japanese Journal of Ichthyology 22(4), 241-
242.
Bagarinao, T. (1994a) The natural life history of milkfish. SEAFDEC Asian
Aquaculture 16(3), 3-6.
Bagarinao, T. (1994b) Systematics, distribution, genetics and life history of milkfish,
Chanos chanos. Environmental biology of fishes 39(1), 23-41.
Bagarinao, T., and Kumagai, S. (1987) Occurrence and distribution of milkfish
larvae, Chanos chanos off the western coast of Panay Island, Philippines.
Environmental biology of fishes 19(2), 155-160.
Bagarinao, T., and Thayaparan, K. (1986) The length-weight relationship, food
habits and condition factor of wild juvenile milkfish in Sri Lanka.
Aquaculture 55(3), 241-246.
53
Bagarinao, T.U. (1991) 'Biology of milkfish (Chanos chanos Forsskal).' (SEAFDEC
Aquaculture Department)
Baumann, H., Peck, M.A., and Herrmann, J.-P. (2005) Short-term decoupling of
otolith and somatic growth induced by food level changes in postlarval Baltic
sprat, Sprattus sprattus. Marine and Freshwater Research 56(5), 539-547.
Begg, G.A., Campana, S.E., Fowler, A.J., and Suthers, I.M. (2005) Otolith research
and application: current directions in innovation and implementation. Marine
and Freshwater Research 56(5), 477-483.
Bell, J.D., Kronen, M., Vunisea, A., Nash, W.J., Keeble, G., Demmke, A., Pontifex,
S., and Andréfouët, S. (2009) Planning the use of fish for food security in the
Pacific. Marine Policy 33(1), 64-76.
Berumen, M.L., Almany, G.R., Planes, S., Jones, G.P., Saenz-Agudelo, P., and
Thorrold, S.R. (2012) Persistence of self-recruitment and patterns of larval
connectivity in a marine protected area network. Ecology and Evolution 2(2),
444-452.
Billings, G., and Pickering, T. (2010) Fiji launches milkfish aquaculture project for
food security. Secretariat of the PaciÀc Community Fisheries Newsletter, No.
132, Noumea.
Bok, A.H. (1979) The distribution and ecology of two mullet species in some fresh
water rivers in the Eastern Cape, South Africa. Journal of the Limnological
Society of Southern Africa 5(2), 97-102.
54
Brian, W.C. (2015) Review of the milkfishes of Iran (Family Chanidae). Iranian
Journal of Ichthyology 2(2), 65-70.
Briggs, J.C. (1961) The east Pacific barrier and the distribution of marine shore
fishes. Evolution 15(4), 545-554.
Buri, P. (1980) Ecology on the feeding of milkfish fry and juveniles, Chanos chanos
(Forsskal) in the Philippines. Mem. Kagoshima Univ. Res. Center S. Pac 1,
25-42.
Carreon, J.A., Laureta, L.V., Estocapio, F.A., and Abalos, T.U. (1984) Milkfish
seedling survival in raceways of freshwater recirculating systems.
Aquaculture 36(3), 257-272.
Dela Cruz, E. (1997) Potential of milkfish farming development in Fiji. Vol. 2017.
(FAO: Rome)
Edwards, A.M. (2001) Adding detritus to a nutrient–phytoplankton–zooplankton
model: a dynamical-systems approach. Journal of Plankton Research 23(4),
389-413.
El Meseda, M., Gharabawy, S., and Assem, S. (2006) Spawning induction in the
Mediterranean grey mullet Mugil cephalus and larval developmental stages.
African Journal of Biotechnology 5(19), 1836.
FAO (2014) The State of World Fisheries and Aquaculture. 2014 edn. pp. 22. (FAO:
Rome)
55
FAO (2016) The State of World Fisheries and Aquaculture. 2016 edn. pp. 200.
(Rome)
Fey, D.P., Bath Martin, G.E., Morris, J.A., and Hare, J.A. (2005) Effect of type of
otolith and preparation technique on age estimation of larval and juvenile
spot (Leiostomus xanthurus). Fishery Bulletin 103(3), 544-552.
FitzGerald, W. (2004) Milkfish aquaculture in the pacific: potential for the tuna
longline fishery bait market. Noumea, New Caledonia: Secretariat of the
Pacific Community.
Fricke, R. (2008) Authorship, availability and validity of fish names described by
Peter (Pehr) Simon Forsskål and Johann Christian Fabricius in the
‘Descriptiones animalium’by Carsten Niebuhr in 1775 (Pisces).
Garcia, L.M.B. Fisheries biology of milkfish (Chanos chanos Forskal). In 'In:
Tanaka, H., Uwate, KR, Juario, JV, Lee, CS, Foscarini, R.(eds.). Proceedings
of the Regional Workshop on Milkfish Culture Development in the South
Pacific, 21-25 November 1988, Tarawa, Kiribati. Suva, Fiji: Food and
Agriculture Organization of the United Nations, South Pacific Aquaculture
Development Project. pp. 66-76', 1990, pp. 66-76
Helfman, G., Collette, B.B., Facey, D.E., and Bowen, B.W. (2009) 'The Diversity of
Fishes: Biology, Evolution, and Ecology.' (Wiley)
56
Horký, P., Slavík, O., Bartos, L., Kolárová, J., and Randák, T. (2006) The effect of
the moon phase and seasonality on the behaviour of pikeperch in the Elbe
River. Folia Zoologica 55(4), 411.
Horwood, J., Nichols, J., and Milligan, S. (1998) Evaluation of closed areas for fish
stock conservation. Journal of Applied Ecology 35(6), 893-903.
Jan, A., Rab, A., Ullah, R., and Ullah, I. (2017) On the Breeding Season of
Schizothorax plagiostomus. Pakistan Journal of Zoology 49(2).
Jaspe, C.J., Golez, M.S.M., Coloso, R.M., and Caipang, C.M.A. (2012) Production
of hatchery-bred early juvenile milkfish (Chanos chanos) in nursery ponds
through supplemental feeding. Animal Biology and Animal Husbandry 4(2),
32-37.
Johannes, R.E. (1978) Reproductive strategies of coastal marine fishes in the tropics.
Environmental Biology of Fishes 3(1), 65-84.
Kawamura, G. The sense organs and behaviors of milkfish fry in relation to
collection techniques. In 'Advances in milkfish biology and culture:
proceedings of the Second International Milkfish Aquaculture Conference, 4-
8 October 1983, Iloilo City, Philippines', 1984, pp. 69-84
Kawamura, G., and Castillo Jr, A.R. (1981) A new device for recording the feeding
activity of milkfish. Bulletin of the Japanese Society of Scientific Fisheries
47(1), 141.
57
Kawamura, G., Hara, S., and Bagarinao, T. (1980) A fundamental study on the
behavior of milkfish fry for improving the efficiency of traditional fry
collecting gear in the Philippines. Mem. Kagoshima Univ. Res. Center S. Pac
1, 65-74.
Kawamura, G., and Washiyama, N. (1984) Age determination of wild-captured
milkfish larvae as indicated by daily growth increments of otolith. Bulletin of
the Japanese Society of Scientific Fisheries.
Klausewitz, W. (1965) 'On Forsskål's collection of fishes in the Zoological
Museum of Copenhagen.' (København : [I Kommission Hos E. Munksgaard],
1965.)
Klunzinger, K.B. (1870) 'Synopsis der fische des Rothen Meeres.' In (Wien :C.
Ueberreuter'she Buchdruckerei: New York) Available at
https://doi.org/10.5962/bhl.title.1148 [Verified July 2017]
Kumagai, S. The ecological aspects of milkfish fry occurrence, particularly in the
Philippines. In 'Advances in milkfish biology and culture: proceedings of the
Second International Milkfish Aquaculture Conference, 4-8 October 1983,
Iloilo City, Philippines', 1984, pp. 53-68
Kumagai, S. (1990) Reproduction and early life history of milkfish Chanos chanos in
the waters around Panay Island, Philippines. Ph. D. Dissertation, Kyushu
University,
58
Kumagai, S., Bagarinao, T., and Unggui, A. (1985) Growth of juvenile milkfish
Chanos chanos in a natural habitat. Marine ecology progress series.
Oldendorf 22(1), 1-6.
Kumagai, S., and Bagarinao, T.U. (1981) Studies on the habitat and food of juvenile
milkfish in the wild. Fisheries Research Journal of the Philippines 6(1), 1-10.
Kuo, C.-M., and Nash, C.E. (1979) Annual reproductive cycle of milkfish, Chanos
chanos Forskal, in Hawaiian waters. Aquaculture 16(3), 247-251.
Lassig, B.R. (1983) The effects of a cyclonic storm on coral reef fish assemblages.
Environmental Biology of Fishes 9(1), 55-63.
Leis, J. (1984) Larval fish dispersal and the East Pacific Barrier. Océanographie
tropicale 19(2), 181-192.
Leis, J.M., and Reader, S.E. (1991) Distributional ecology of milkfish, Chanos
chanos, larvae in the Great Barrier Reef and Coral Sea near Lizard Island,
Australia. Environmental biology of fishes 30(4), 395-405.
Li, X., Huang, T., Ma, W., Sun, X., and Zhang, H. (2015) Effects of rainfall patterns
on water quality in a stratified reservoir subject to eutrophication:
Implications for management. Science of the Total Environment 521, 27-36.
Liao, I.-C., Juario, J.V., Kumagai, S., Nakajima, H., Natividad, M., and Buri, P.
(1979) On the induced spawning and larval rearing of milkfish, Chanos
chanos (Forskal). Aquaculture 18(2), 75-93.
59
Liao, I.C., Yan, H.Y., and Su, M.S. (1977) Studies on milkfish fry. 1. On
morphology and its related problems of milkfish fry from the coast of
Tungkang. Journal of the Fisheries Society of Taiwan 6(1), 73-83.
Lim, C., Borlongan, I., and Pascual, F. (2002) Milkfish, Chanos chanos. Nutrient
requirements and feeding of finfish for aquaculture, 172-183.
Lin, Y., Chen, C., and Lee, T. (2003) The expression of gill Na, K-ATPase in
milkfish, Chanos chanos, acclimated to seawater, brackish water and fresh
water. Comparative Biochemistry and Physiology Part A: Molecular &
Integrative Physiology 135(3), 489-497.
Mallya, Y.J. (2007) The effects of dissolved oxygen on fish growth in aquaculture.
The United Nations University fisheries training programmer, Final project,
pp30.
Marte, C.L., and Lacanilao, F. (1986a) Spontaneous maturation and spawning of
milkfish in floating net cages. Aquaculture 53(2), 115-132.
Marte, C.L., Toledo, J.D., Quinitio, G.F., and Castillo Jr, A.R. Collection of
naturally-spawned milkfish eggs in floating cages. In 'The First Asian
Fisheries Forum. Proceedings of the First Asian Fisheries Forum, 26-31 May
1986, Manila, Philippines', 1986b, pp. 671-674
Martinez, F.S., Tseng, M., and Yeh, S. (2006) Milkfish (Chanos chanos) culture:
situations and trends. Journal-Fisheries Society of Taiwan 33(3), 229.
60
Microsoft Officer (2013) Home and Student - Mcrosoft excel. 2013 edn. (Microsoft)
Miller, J.A., Simenstad, C.A., and Team, W.E. (1994) 'Otolith Microstructure
Preparation, Analysis, and Interpretation: Procedures for a Potential Habitat
Assessment Methodology.' (Fisheries Research Institute, School of Fisheries,
University of Washington)
Morioka, S., Ohno, A., Kohno, H., and Taki, Y. (1993) Recruitment and survival of
milkfish Chanos chanos larvae in the surf zone. Japanese Journal of
Ichthyology 40(2), 247-260.
Myers, R.A., and Worm, B. (2003) Rapid worldwide depletion of predatory fish
communities. Nature 423(6937), 280-283.
Napulan, M.R. (2012) Keys to Successful Milkfish Farming. pp. 1-84. (Fiji)
Nelson, A.L.M. (2007) Cultured Aquatic Species Information Programme: Chanos
chanos (Forsskal, 1775). 2007 edn. (Food and Agriculture Organization of
the United Nations: Rome)
Nelson, J.S. (2006) 'Fishes of the World.' (John Wiley & Sons)
Oren, O.H. (1981) 'Aquaculture of Grey Mullets.' (Cambridge University Press)
Panfili, J., De Pontual, H., Troadec, H., and Wrigh, P.J. (2002) 'Manual of fish
sclerochronology.' In Manuel de sclérochronologie des poissons Available at
http://archimer.ifremer.fr/doc/00017/12801/
61
Pickering, T., Tanaka, H., and Senikau, A. (2012) Capture-based Aquaculture of
Milkfish Chanos chanos in the Pacific Islands. Secretariate of the Pacific
Community, No. 978-982-00-0615-7, Fiji.
Pickering, T.D., Ponia, B., Hair, C.A., Southgate, P.C., Poloczanska, E.S., Della
Patrona, L., Teitelbaum, A., Mohan, C.V., Phillips, M.J., and Bell, J.D.
(2011) Vulnerability of aquaculture in the tropical Pacific to climate change.
Vulnerability of tropical Pacific fisheries and aquaculture to climate change,
647-732.
Pillay, T.V.R., and Kutty, M.N. (2005) 'Aquaculture: principles and practices.'
(Blackwell publishing)
Popper, A.N., Ramcharitar, J., and Campana, S.E. (2005) Why otoliths? Insights
from inner ear physiology and fisheries biology. Marine and freshwater
Research 56(5), 497-504.
R Core Team (2007) R: A language and environment for statistical computing. R
Foundation for Statistical Computing. (Vienna, Austria)
Sackett, D.K., Cope, W.G., Rice, J.A., and Aday, D.D. (2013) The influence of fish
length on tissue mercury dynamics: implications for natural resource
management and human health risk. International journal of environmental
research and public health 10(2), 638-659.
Schuster, W.H. (1960) Synopsis of biological data on milkfish Chanos chanos
(Forsskal), 1775. FAO, Rome.
62
Senta, T., and Hirai, A. (1981) Japanese Journal of 1chthyology 28(1), 45-51.
Senta, T., and Kumagai, S. (1977) Variation in the vertebral number of the milkfish
Chanos chanos, collected from various localities. Nagasaki University
Faculty of Fisheries Research Report 43, 35-40.
Silas, E., Mohanraj, G., Gandhi, V., and Thirunavukkarasu, A. (1982) Spawning
grounds of the milkfish and seasonal abundance of the fry along the east and
southwest coasts of India. In Proceedings of the symposium on Coastal
Aquaculture. Vol. 3. pp. 916-932. (The Marine Biological Association of
India).
SPC (2008) Vanuatu Aquaculture Development Plan 2008-2013. Secretariate of the
Pacific Community, New Caledonia.
Spennemann, D.H. (2002) Traditional milkfish aquaculture in Nauru. Aquaculture
International 10(6), 551-562.
Statistica Inc (2011) STATISTICA (data analysis software system).
Sulu, R.J., Vuto, S.P., Schwarz, A.M., Chang, C.W., Alex, M., Basco, J.E., Phillips,
M., Teoh, S.J., Perera, R., Pickering, T., and Oengpepa, C.P. (2016) The
feasibility of milkfish (Chanos chanos) aquaculture in Solomon Islands. pp.
1-94. (WorldFish: Malaysia)
63
Tampi, P.R. (1957) Some observations on the reproduction of the milkfish Chanos
chanos (Forskål). Proceedings: Plant Sciences 46(4), 254-273.
Tidwell, J.H., and Allan, G.L. (2001) Fish as food: aquaculture’s contribution:
Ecological and economic impacts and contributions of fish farming and
capture fisheries. EMBO Reports 2(11), 958-963.
Tzeng, W.-N., and Yu, S.-Y. (1989) Validation of daily growth increments in
otoliths of milkfish larvae by oxytetracycline labeling. Transactions of the
American Fisheries Society 118(2), 168-174.
Tzeng, W.N., and Yu, S.Y. (1988) Daily growth increments in otoliths of milkfish,
Chanos chanos (Forsskål), larvae. Journal of fish biology 32(4), 495-504.
Villaluz, A. Fry and fingerling collection and handling. In 'Aquaculture of milkfish
(Chanos chanos): State of the Art', 1986, pp. 153-180
Villaluz, A., and Unggui, A. (1983a) Effects of Temperature on Behavior, Growth,
Development and Survival in Young Milkfish, Chanos chanos (Forskal).
Aquaculture 35, 321-330.
Villaluz, A.C. Collection, storage, transport, and acclimation of milkfish fry and
fingerlings. In 'Advances in milkfish biology and culture: proceedings of the
Second International Milkfish Aquaculture Conference, 4-8 October 1983,
Iloilo City, Philippines', 1984, pp. 85-96
Villaluz, A.C. Milkfish fry collection and handling. In 'In: Tanaka, H., Uwate, KR,
Juario, JV, Lee, CS, Foscarini, R.(eds.). Proceedings of the Regional
64
Workshop on Milkfish Culture Development in the South Pacific, 21-25
November 1988, Tarawa, Kiribati. Suva, Fiji: Food and Agriculture
Organization of the United Nations, South Pacific Aquaculture Development
Project. pp. 77-87', 1990, pp. 77-87
Villaluz, A.C., Villaver, W.R., and Salde, R.J. (1983b) Milkfish fry and fingerling
industry of the Philippines: methods and practices. Aquaculture Department,
Southeast Asian Fisheries Development Center, No. 0115-4710.
Vuto, S.P., Oengpepa, C., R, S., T, P., M, P., Schwarz, A.M., R, W., S, S., A, T., and
C, T. (2014) 'Fry seasonality and feed trial studies of milkfish Chanos chanos
(Forskal, 1775) from the coast of Gizo, Solomon Islands.' In World
Aquaculture Meeting (University of the South Pacific/World Fish) Available
at https://www.was.org/meetings/ShowAbstract.aspx?Id=33203
Wainwright, T. (1982) Milkfish fry seasonality on Tarawa, Kiribati, its relationship
to fry seasons elsewhere, and to sea surface temperatures (SST). Aquaculture
26(3-4), 265-271.
Warner, R.R., Swearer, S.E., Caselle, J.E., Sheehy, M., and Paradis, G. (2005) Natal
trace-elemental signatures in the otoliths of an open-coast fish. Limnology
and Oceanography 50(5), 1529-1542.
Williams, L., and Rota, A. (2010) Impact of climate change on fisheries and
aquaculture in the developing world and opportunities for adaptation.
World Bank (2013) FISH TO 2030 Prospects for Fisheries and Aquaculture. (Ed.
TW Bank) pp. 1-102. (World Bank: USA)
65
66
Appendix
Appendix A. Table of p-values for the ANOVA test
The following data were tested with ANOVA for the level of significance in R- software Sites Compared
to Milkfish fry abundance (number)
Significance level = p-value (95%)
Kawenu Erakor 1.00 Mele Erakor 1.00 Teouma Erakor 0.00 Mele Kawenu 1.00 Teouma Kawenu 0.01 Teouma Mele 0.01 Month Compare
to Milkfish fry abundance (number)
Significance level = p-value (95%)
January November 0.00 February November 0.02 March November 0.03 May November 0.00 October November 0.00 December November 0.00 January April 0.00 February April 0.00 March April 0.00 May April 0.00 October April 0.00 December April 0.00 November April 0.67 October June 0.99 December June 0.94 January June 1.00 February June 0.16 March June 0.27 May June 1.00 October July 0.96 December July 0.82 January July 1.00 February July 0.03 March July 0.08 May July 0.99 October August 0.98 December August 0.89 January August 1.00 February August 0.08 March August 0.17
67
May August 1.00 October September 0.98 December September 0.90 January September 1.00 February September 0.08 March September 0.17 May September 0.99 Month Compare
to Milkfish fry length (mm) Significance level = p-value (95%)
January April 0.89 February April 0.02 March April 0.59 May April 0.03 October April 0.92 November April 0.04 December April 0.38 January December 1.00 February December 1.00 March December 1.00 May December 0.95 October December 0.17 November December 1.00 January February 1.00 March February 0.96 May February 0.97 October February 0.03 November February 0.99 March January 1.00 May January 1.00 October January 0.68 November January 1.00 May March 0.67 October March 0.29 November March 1.00 Month Compare
to Milkfish fry weight (mg) Significance level = p-value (95%)
January April 0.67 February April 1.00 March April 1.00 May April 1.00 October April 0.28 November April 0.61 December April 0.98 January December 0.91 February December 1.00
68
March December 0.96 May December 1.00 October December 0.96 November December 1.00 January February 0.97 March February 1.00 May February 1.00 October February 1.00 November February 1.00 March January 0.62 May January 0.79 October January 1.00 November January 0.91 May March 1.00 October March 0.28 November March 0.69 Moon Compared
to Milkfish fry abundance (number)
Significance level = p-value (95%)
Full 1st moon 0.86 New 1st moon 0.01 3rd Moon 1st moon 0.12 New moon Full moon 0.00 3rd Moon Full moon 0.02 3rd Moon New moon 0.44 Moon Compared
to Total length for fry (mm)
Significance level = p-value (95%)
Full 1st moon 0.60 3rd Moon 1st moon 0.10 New moon 1st moon 0.00 3rd Moon Full moon 0.30 New moon Full moon 0.00 New moon 3rd moon 0.00 Moon Compared
to Total weight of fry (mm)
Significance level = p-value (95%)
Full 1st moon 0.29 New moon 1st moon 0.00 3rd moon 1st moon 0.00 New moon Full moon 0.53 3rd moon Full moon 0.61 3rd moon New moon 1.00
Appendix B. The daily
median and standard
2017.
69
y SST from VMDG and monthly mean, maxim
d deviation beginning from February 17th 201
mum, minimum,
16 to February 17,
70
Appendix C. Milkfish fry mean length and weight plus the descriptive statistic for
Teouma over one year period
Descriptive statistics
Length (mm) Weight (mg)
Milkfish fry Mean 12.8 5.78 Standard Error 0.1 0.08 Median 13.0 6 Mode 13.0 5 Standard Deviation
1.2 1.6
Sample Variance 1.5 2.5 Kurtosis 0.4 -0.5 Skewness -0.6 0.2 Range 7 6 Minimum 8 3 Maximum 15 9 Sum 4709 2135 Count 369 369