You are on page 1of 25

CHAPTER 7

Flyingfish

Robert Gillett and James Ianelli

I. INTRODUCTION
Flyingfish represent an important resource in many parts ofthe world. Several
Pacific islands currently have developed flyingfish fisheries and many have a
history of traditional fisheries for flyingfish. Some Pacific islands do not have
flyingfish fisheries, yet the abundance of the resource appears to be at least as
great as other areas. As fishing pressure on limited reef resources increases, the
development of alternative fisheries is needed, particularly for small- scale
fishermen. Preliminary investigations suggest that flyingfish may also fall into
this category.
This chapter presents information obtained from a review of available
literature, discussions with fisheries workers, correspondence with flyingfish
authorities, and recent flyingfish fishing trials. This provides the basis for an
assessment of the potential for fisheries development for this resource in the
South Pacific.

II. BIOLOGY

In the following section, aspects of the biology of flyingfish are presented


with as much reference as possible to the Pacific islands situation. In many
cases, however, for lack of details specific to the Pacific islands, information on
studies from other parts of the world is provided.

IDENTIFICATION

In order to address fundamental questions on population dynamics and


biology of flyingfish, it is important to be able to identify clearly the species
involved. Flyingfishes (family Exocoetidae) are closely related to the garfishes
(family Hemiramphidae), longtoms or needlefish (family Belonidae), and
sauries (family Scomberosocidae). Although the adult forms vary consider-
ably, early developmental stages of these families have many features in
common. In particular, these fish typically have filaments on their eggs,
develop chin barbels at juvenile stages, and share a similar appearance at early
developmental stages.
178 Robert Gillett and James Ianelli

The taxonomy of flyingfishes is complex. Problems in properly identifying


flyingfish are due to 1) the large number of species that occur in the Pacific
islands, 2) the uncertain distinction between some genera, 3) the likelihood that
several species have yet to be described, and 4) juveniles that look significantly
different from adults of the same species. The difficulty in obtaining adequate
reference material further complicates the situation.
Parin (1960a) states that about 41 species of flyingfishes inhabit the western
part of the Pacific Ocean, which includes 29 species in the Coral Sea, 27 in
Indonesia-New Guinea, 19 in Marianas-Carolines-Marshalls,and 19 in Gilberts-
Table I. Flyingfish species that are likely to occur in the Pacific Islands.
Adapted from Parin (1960a), Kailola (1987) and Wass (1984).

Cheilopogon Exocetus
antoncichi monocirrhus
furcatus obtusirostris
unicolor1 olitans
arcticeps
atrisignis Hirundichthys
cyanopterus albimaculatus
intermedins speculiger
longibarbus
nigricans2 Parexocoetus
spilonotopterus brachypterus
spilopterus mento
suttoni
Prognichthys
Cypselurus sealei
angusticeps agoo
oligolepis3 albimaculatus
naresii brevipennis
pitcairnensis
poecilopterus
simus

1. The distinctions among Cheilopogon antoncichi, Ch. unicolor, and Ch. furcatus awaits
clarification and the names may have been used interchangeably by various authors.
2. Probably more than one species.
3. The distinction between Cypselurus angusticeps and C. oligolepis awaits clarification
and the names may have been used interchangeably.
Flyingfish 179

Samoa-Fiji-Tonga. Wass (1984), based on correspondence with Parin, indi-


cates that 16 species of flyingfishes occur near Samoa.
Among the references listed in the attached bibliography, 14 papers contain
primarily taxonomic information. Many authors indicate that six genera of
flyingfish occur in the Pacific islands (see Table I). These are: Cheilopogon,
Cypselurus, Exocoetus, Hirundichthys, Parexocoetus, and Prognichthys.
Two points should be noted:
(1) The genus Oxyporhamphus has, in the past, been included in the family
Exocoetidae but more recently there is agreement that it belongs in the
family Hemiramphidae;
(2) Cheilopogon includes species that were formerly placed in the genus
Cypselurus.
Because the same six flyingfish genera occur in the Indian Ocean, the 1984
FAO Species Identification Sheets for the Western Indian Ocean (Appendix 2)
can be used to identify Pacific islands flyingfish genera.
Identification to the species level is considerably more difficult. A revision
of the taxonomy of the group has been proposed by FAO and would culminate
in a comprehensive catalogue of flyingfish species found in the Pacific islands
and elsewhere.
To obtain taxonomic information on the species caught in some of the Pacific
island flyingfish fisheries, specimens were collected in 1989 and 1990 from the
nearshore fisheries in several Pacific islands countries and sent to the National
Marine Fisheries Service Systematics Laboratory in Washington, D.C. The
Laboratory Director, B. Collette, kindly made tentative identifications of the
species. In June 1990 the specimens were re-identified by N. Parin. The
results of these examinations are given in Table II. The discrepancies between
the opinions of the two taxonomists highlight the difficulty that fishery workers
are likely to encounter in correctly identifying flyingfishes.
Collette indicates that the Cheilopogon and Cypselurus species listed in the
table (which presumably constitute a major portion of the Pacific islands
flyingfish catch) can be distinguished using the following characteristics:
A) Pigment of pectoral fins
spotted: atrisignis, poecilopterus, suttoni
clear: unicolor
dark: spilopterus
dark with white margin: oligolepis, spilonotopterus
B) Pigment of dorsal fin
clear: unicolor, poecilopterus, oligolepis
with black spot: atrisignis, spilopterus, spilonotopterus,
suttoni
180 Robert Gillett and James Ianelli

DISTRIBUTION

Flyingfish common in the tropical Pacific have limits of distribution bounded


by 40° S and 40° N (Kovalevskaya 1982). Some authors partition flyingfish
species into two groups, neritic (coastal) and oceanic. "Oceanic" species can be
found in coastal areas; likewise, "coastal" species can also be found in oceanic
areas. An oceanic species does not require coastal habitat for any stage of its life
history, in particular, spawning. Coastal species on the other hand, use the
coastal environment for some stage of their life cycle, typically to spawn.
Parin (1960a) states that in the Pacific islands there are about 28 species in
the coastal group and 13 in the oceanic. He indicates that the proportion of
coastal to oceanic species is lower in the central Pacific and increases to the
west. Presumably this is related to the larger land masses in the western
Pacific. All of the species listed in Table II were obtained from nearshore fish-
eries in the central Pacific. Seven are described by Parin (1960a) as coastal
species while Cheilopogon suttoni is considered oceanic.
For fisheries involved in catching flyingfish at the surface at night, the depth
at which the fish are found will undoubtedly affect catchability. The vertical
distribution of flyingfish extends to 18 m during the night while roughly 86 per
cent are found in the shallow layer from 0-2 m (Nesterov and Bazanov
1986). The brightness of the moon may affect flyingfish depth, although no
studies of this phenomena are known.
Several visual survey methods have been developed to estimate the abun-
dance and distribution of flyingfish (e.g. Parin, 1983; Zuyev and Nikolsky,
1980; Oxenford et al, 1989). Problems with these methods include difficulties
in identifying species and determining absolute abundance. Nonetheless, these
methods can give some indication of the relative abundance and the suitability
of flyingfish habitat. Oxenford et al. (1989) found that in the Caribbean, the
distribution of//, affinis is considerably wider than the region where the current
fishery operates. Such definitive studies have not been conducted in the Pacific
islands region.

MIGRATION/DISPERSION

The extent of migration and/or dispersion of flyingfish will affect the degree
to which they can withstand fishing pressure. As noted above, there are oceanic
and coastal species of flyingfish. The oceanic group has a greater range but
fewer species than the coastal group.
In the Pacific islands region, no studies have been conducted to determine the
extent of flyingfish movements. In the Caribbean, tagging studies of
Hirundichthys affinis, a species that may be similar to H. speculiger found in the
Pacific, show that movement may be related to oceanographic conditions. These
Flyingflsh 181
Table II. Identification of flyingfish specimens from eight Pacific islands countries.

Origin of Collette's No. Parin's No.


Samples Identification Indiv. Identification Indiv.

Tonga 1: P. brachypterus 0) P. brachypterus 0)


(Jan-Apr) C. poecilopterus (2) C. poecilopterus (2)
1989 Ch. spilopterus (1) Ch. spilopterus (1)
Ch. suttoni (3) Ch. suttoni (3)
Ch. spilonotopterus 0) Ch. spilonotopterus 0)
Ch. atrisignis (2) Ch. atrisignis (2)
Ch. unicolor (1) Ch. antoncichi 0)
C. oligolepis 0) C. sp. 0)
Tonga 2: Ch. unicolor (3) Ch. antoncichi (3)
(Sept 89) Ch. spilonotopterus (2) Ch. atrisignis (2)

Tahiti: C. poecilopterus (6) C. poecilopterus (2)


(June 89) - C. pitcairnensis (4)

Cook Is: C. poecilopterus (1) C. poecilopterus (1)


(Feb 89) Ch. unicolor (2) Ch. antoncichi (2)
Ch. atrisignis (2) Ch. atrisignis (2)

Niue: Ch. spilonotopterus 0) Ch. spilonotopterus (2)


(Sept 89) Ch. atrisignis (2) Ch. atrisignis (2)
Ch. unicolor (2) Ch. antoncichi 0)
C. oligolepis (2) C. angusticeps (3)
(Unknown Species) (1)

Tokelau: Ch. atrisignis 0) Ch. atrisignis 0)


(Aug 89) Ch. unicolor (1) Ch. antoncichi (1)
Ch. spilonotopterus (5) Ch. spilonotopterus (6)
C. oligolepis 0) C. angusticeps (2)

Tuvalu: Ch. unicolor (3) Ch. antoncichi (3)


(Aug 89) Ch. spilonotopterus (4) Ch. spilonotopterus (6)
Ch. suttoni (2) -

Kiribati: - Ch. spilonotopterus (6)


(March 90) Ch. suttoni (3)

Papua New
Guinea: - C. oligolepis (4)
(Jan-Feb 90)

Total number specimens 53 Total number specimens 68


Total number species 9 Total number species 11
182 Robert Gillett and James Ianelli

studies revealed that H. affinis probably does not undertake extensive migra-
tions by life history stages. That is, the larvae and juveniles appear in the same
areas as the adults rather than moving from a "nursery" area. The greatest
recorded movement of a tagged fish was 200 nautical miles and the fastest
estimated speed was greater than 16 nautical miles per day (Oxenford et
al. 1989). The longest time at liberty for a tagged flyingfish was 121 days with
a mean time of 21 days. Twenty-two per cent of all taggedfishwere recaptured
in waters of countries other than those in which the fish were released. This
indicates that flyingfish can move considerable distances and can be subject to
capture in more than one fishery.

PREDATORS

As is well known, flyingfish escape predators by burst swimming that propels


them into the air where they can glide with their enlarged pectoral and pelvic
fins. Despite this elegant adaptation, flyingfish fall prey to many
species. Flyingfish were consistently found in the stomachs of skipjack tuna and
other large pelagics in the Pacific islands (South Pacific Commission, 1980-
1985). Olson and Boggs (1986) estimate that flyingfish make up from 4 to 13
per cent of the diet of yellowfin tuna in the eastern Pacific. No information is
available to suggest that the abundance of any flyingfish predators is limited by
the availability of flyingfish.

FOOD AND FEEDING

The food of flyingfish consists mainly of large zooplankton and small


fish. From analysis of stomach contents it has been found that feeding of
H. a//?«wintheCaribbeantakesplaceatnight(Lewisera/., 1962). Hall(1955)
examined 425 stomachs of H. affinis and found 22.4 per cent of the stomachs
contained copepods, 27.1 per cent contained other crustaceans, and 17.2 per cent
of the stomachs contained small fish. The study by Lewis et al. (1962) found
that fish occurred in 45.3 per cent of the stomachs examined, 32.2 per cent
contained copepods, and 21.0 per cent contained other crustaceans. Gorelova
(1980) found that the flyingfish he examined from the Pacific feed
opportunistically. Between species of flyingfish, there was no evidence of
specialized prey selectivity. He also concluded that larvae andjuvenile flyingfish
feed near the surface primarily during daylight hours at an average rate of 15-
20 per cent of their body weight per day.
Flyingfish presumably eat the same food items as other pelagic predators,
however, no information exists on how this potential competition may affect
flyingfish survival.
Flyingfish 183

GROWTH

Tropicalflyingfish,like many pelagic fish species, grow rapidly. To date no


growth studies are known to have been done on species supporting Pacific
islands fisheries. Studies on growth of tropicalflyingfishhave been carried out
in the Philippines (Dalzell et al, unpubl. m.s.), Indonesia (Watson, 1990), and
in the Caribbean (Storey, 1983). Flyingfish in the tropics generally live to about
2 years of age and are mature after 10-14 months. The growth rate and
maximum size varies by species with the oceanic species typically having a
smaller maximum size. Many of the commercially important species (e.g. from
the genera Hirundichthys, Cypselurus, and Cheilopogon) grow to about 20-
25cm and attain weights of 300-450g.

SPAWNING

Flyingfish deposit their eggs on debris, algae, or other substrate. Oceanic


species have fully pelagic free-floating eggs or deposit their eggs on the limited
debris found in the open ocean. In Japan, studies of Cypselurus opisthopus
indicate that this species offlyingfishspawns in association with sandy bottoms
and is vulnerable to capture with bottom gillnets.
As an example of the range and period over which some species of flyingfish
actively spawn, Breder and Rosen (1966) report that Parexocoetus mento spawn
off Japan in the warm months from May to September. The same species
collected off Fiji in October 1991 (where seasons are opposite to those in Japan)
was found in spawning condition and had fully hydrated eggs in the ovaries. The
Japanese studies cited by Breder and Rosen (1966) also state that most individu-
als of P. mento die after spawning.
In the Pacific, there are several anecdotal accounts of flyingfish spawning
behaviour. In the Cook Islands, the people of Atiu celebrate the arrival of the
flyingfish (Mokoroa, 1984). They observe three stages where capture is
possible: when the fish jump and glide predictably in one direction toward the
spawning grounds, when they lie still on the surface of the water to broadcast
their eggs and milt, and finally when they begin sinking to greater depths. It is
believed that a significant proportion of flyingfish die after spawning.
In Tokelau, similar behaviour thought to be associated with flyingfish
spawning is known as tuali. On Huahine, French Polynesia, theflyingfishhave
been observed to spawn in the sand in shallow water. Johannes (1981) gives the
following description of flyingfish spawning in Huahine, Society Islands:
During certain months, the most important being August, September, and
October, schools offlyingfishcome boiling through two reef passes near
the town dock at sunset, pursued by predators. The fish swim into a few
184 Robert Gillett and James Ianelli

inches of water close to shore, wriggle tail first into the sand and deposit
their spawn much like California grunion. Spawning takes only a few
seconds, after which thefishreturn to deeper water. No lunar periodicity
was noted.
For many of these observations, it is impossible to determine which species
is involved and how frequently these events occur. Some scientists have
expressed skepticism about accounts of flyingfish spawning in sand because no
other species in related families spawn in this manner. Furthermore, flyingfish
easily lose their scales, hence they may not survive spawning that involves
extensive contact with the substrate. Mature flyingfish in the tropics appear to
spawn several times over the course of a year. In studies on the Caribbean
species, H. qffinis, four distinct egg sizes have been found in the ovaries with
the largest being fully hydrated and ready to spawn (Storey 1983). This suggests
that not all eggs are spawned at once and that the spawning season is protracted.

FECUNDITY

Small species offlyingfish (Farexocoetus mento, certain species ofExocoetus


and Prognichthys) spawn between 400 to 1,100 eggs at a time (Kovalevskaya
1982). Larger forms have considerably greater fecundity, ranging between
16,000 and 24,000 eggs in certain species in the genera Cheilopogon, Cypselurus,
andHirundichthys. Flyingfish fecundity at a single spawning is not necessarily
a measure of the total reproductive potential of that individual. The fact that
many flyingfish species spawn several times per year suggests that total
fecundity is a better measure of reproductive output. Studies of//, qffinis egg
size indicate that they may spawn at least four times per year (Storey 1983).

EARLY LIFE HISTORY

The eggs of flyingfish have several distinctive characteristics. Flyingfish


eggs are negatively buoyant and typically have long sticky filaments that serve
to attach the eggs to floating objects. Yellowish white gelatinous masses often
observed on floating coconuts or other debris in the ocean are most likely
flyingfish eggs. Species within the genus Exocoetus are unlike most flyingfish
species in that their eggs are larger, do not have filaments, and are positively
buoyant. Presumably this is an adaptation to their oceanic spawning nature.
Among speciesof flyingfish, mean egg size is inversely related to the number
of eggs produced per unit of body weight. This implies that some flyingfish
species have evolved to invest in larger but fewer eggs, while others have been
successful with smaller but more numerous eggs. The survival offish hatching
from larger eggs would likely be higher than fish from small eggs under similar
conditions.
Flyingfish 185

Incubation of flyingfish eggs varies from about 4 days to 2 weeks with faster
development occurring in warmer regions. The size at hatch ranges from 3.5 to
6 mm depending on the species (Collette etal., 1984). Oxenford (1985) reports
that H. affinis absorb their yolk sack within a day after hatching. By the time
they are 7 to 10 days old the larvae are about 15 mm in length and are usually
dark blue on the dorsal surface and silvery on the ventral surface. The morphol-
ogy of different larval and juvenile stages changes in several species. Extended
chin barbels are the most common characteristic of flyingfish larvae and vary
considerably in shape and size among species. More primitive species of
flyingfish, e.g. from the genus Parexocoetus, have an elongated lower jaw
(Collette et al, 1984).

MORTALITY

A knowledge of mortality is important to understand the potential producti v-


ity of the resource, particularly from a fisheries development
perspective. Flyingfish appear to grow fast, be relatively short-lived, and spawn
profusely. These features suggest a relatively high natural mortality
rate. Inferences about Pacific islands species, however, can only be made from
studies carried out on similar species elsewhere.
Based on tagging studies in the Caribbean, H. affinis appears to have a high
natural mortality rate (Eastern Caribbean Flyingfish Project, 1989b). One
explanation is that most//, affinis die after the spawning season. Alternatively,
movement of//, affinis to areas where no fisheries occur may explain the pattern
of disappearance. In the Philippines, Dalzell et al. (unpubl. misestimated
natural mortality for Cheilopogon nigricans, and Ch. opisthopus to be in the
range of 70 to 81 per cent per annum but did not discuss the possibility of
extensive spawning-related mortalities. If flyingfish do have high natural
mortality rates, the stocks may be relatively resilient to fishing pressure. The
effect of spawning-related mortality requires further investigation, however.

RECRUITMENT

No information exists on flyingfish recruitment in the Pacific islands, much


less the factors that may be controlling the recruitment levels. In the Caribbean,
Mahon (1987) examined a suite ofbiological and environmental factors that may
affect flyingfish recruitment. He concluded that environmental conditions
influence recruitment more than the observed adult population size. This
suggests that current abundance levels are likely to be relatively independent of
past catches of previous generations of flyingfish. Abundance levels are
therefore likely to be variable, depending on past and present oceanographic
conditions as well as other factors such as predation.
186 Robert Gillett and James Ianelli

IMPLICATIONS FOR DEVELOPMENT AND MANAGEMENT

In summary, several biological characteristics of flyingfish appear to be


favourable from a fisheries development perspective: they appear to be short
lived, fast growing, highly fecund animals with a wide ranging distribution. This
would indicate that overfishing, at least to the moderate catch levels observed
in locally developed fisheries, is unlikely. Studies in the Caribbean show the
potential for interaction between fisheries in different islands based on the
dispersion of tagged fish. This situation may develop in the Pacific but the
distances among countries and the number of species involved are much
greater. Hence, the likelihood of significant interactions between flyingfish
fisheries in different Pacific islands countries is probably low.

III. FISHERIES FOR FLYINGFISH

PACIFIC ISLANDS FLYINGFISH FISHING

Traditional methods: Several techniques have been used in the Pacific islands
to catch flyingfish. Documented traditional flyingfish catching methods are
given in Table III. As with many aspects of Pacific traditional fisheries, much
information on fishing methods has never been adequately recorded.
The dipnet/torch technique is the most widespread, occurring from Palau to
French Polynesia. In areas where flyingfish fishing has developed beyond the
subsistence level, such as in Tahiti and Rarotonga, a modification of the dipnet/
torch technique is used. A good description of the traditional method in Vaitupu
Atoll, Tuvalu, is given by Kennedy (1930):
The canoes draw up in line (tamanga) facing north so as to sweep a front
parallel with the reef on the lee side of the island. The scene is one of
indescribable splendor, the village fires in the distance making dots of
light in the palm jungle. When all are ready the line commences to move
forward at a steady rate of about two knots. In the bow of each canoe
stands the bow-paddler with his dipnet held horizontally across his front
so that the bag is to starboard. That member of the crew immediately
behind the netter in the bow stands and holds aloft the blazing torch. It
is his duty to keep the torch well-trimmed and burning brightly. The other
members of the crew ply their paddles and keep a sharp lookout for
flyingfish lying or swimming near the surface in the vicinity of the
canoe. These appear light grey in colour in the glare of the torches, and
are easily seen at distances up to about forty feet. When the fish are
running well the uproar is deafening. To the neophyte all seems chaos,
but there is a definite thread of order and arrangement. Each canoe keeps
Flying fish 187

its place in line. Usually the netter scoops up one fish at a time and flings
it into a basket. In the thick of a school a netter may capture as many as
three fish, one after another, before emptying his net. When the fish is
lying dazed on the surface of the water, the mouth of the net is brought
down flat on the water with a resounding slap, and in such a manner that
the periphery ofthe bag surrounds the fish which, startled into flight leaps
up into the sack of the bag. Immediately after the slap, the mouth of the
bag is twisted quickly and lifted clear ofthe waterwith the fish inside. This
is the quickest method of taking fish from the water, but can be used only
withfishwhich are right on the surface. When thefishis swimming, the
mouth of the bag is thrust under water a few inches in front of it, in such
a manner that it will, unless it changes course, swim straight into the
opening.
Dipnet fishing at night is still done using torches in some areas. On Temana
Atoll, Kiribati, kerosene lamps have been forbidden because the people there
perceived that flyingfish stocks would be overfished.
Modern methods: Powell (1989a, 1989b) describes the gradual evolution of
the traditional dipnet/torch method into the technique used in Rarotonga
today. Important aspects of this development include the introduction of
kerosene lanterns in the late 1940s to replace palm frond torches, the use of skiffs
powered by outboard motors to replace paddled canoes, and the use of halogen
lamps to replace kerosene lanterns.
Dipnet/torch fishing has evolved to the greatest extent in French
Polynesia. The typical boat (calledpoti marara which literally means flyingfish
boat) used in that fishery is between 5 and 6 metres in length and uses a 60
horsepower outboard engine. In 1987 there were an estimated 272 of these
flyingfish boats in French Polynesia (Service de la Mer, 1989).
In both French Polynesia and the Cook Islands, small generators are used to
power the fishing lights. A high-powered light is affixed to a helmet worn by
the fishermen. This allows the fishermen to direct the light while still having use
of both hands to manoeuvre the boat and manipulate the dipnet. The boats in
these areas are specially designed so that the fisherman can stand in the bow
section of the boat to facilitate scooping. Steering is accomplished by the use
of an aviation-type "joystick" which may have an integrated throttle. The shape
of the hulls is such that they turn easily yet have enough V shape to be
comfortable in moderate seas. An important characteristic of these boats is that
they can easily be used for other types of fishing. In French Polynesia, only
about 40 per cent of thefishlanded from flyingfish boats are flyingfish. Recently,
in response to a request from the Cook Islands Government, the FAO/UNDP
Regional Fishery Support Programme (RFSP) contracted a naval architect to
produce an alternative flyingfish fishing craft which would require a smaller
outboard engine than the 40 to 60 hp versions that are presently being used.
188 Robert Gillett and James Ianelli

Most accounts of night fishing for flyingfish indicate that conditions for
catching are better during hours of maximum darkness. That is, the fisherman's
light is most effective at spotting and immobilizing fish if the moon is below the
horizon and there is no twilight. Calm conditions are often better because it is
easier to spot fish; if there is wind, it is usually best to fish downwind or in the
lee of an island. Scooping requires practice to become proficient and is done
while the fish is in the water, usually not when fish takes flight. The fisherman
must place the net so that the flyingfish swims into it and he must be quick so that
the fish does not change direction and avoid the net. When fish are not very
concentrated, fish that are missed can often be pursued until caught. Points of
reef or land are generally better than bays. Shallow open ocean seamounts are
also good, weather and distance permitting. Water clarity seems to affect catch
rates either because there may be fewer flyingfish in murky water, or they may
be more difficult to see.
Results from fishing introduction trials: In the recent past there have been
several attempts to introduce modern flyingfish fishing methods in the
region. Trials of night fishing for flyingfish using a kerosene lantern in
American Samoa by masterfishermen from the South Pacific Commission
yielded catch rates of 14.9 fish (3.63 kg) per hour over 3 nights (8 hrs total) of
fishing (A. Moana unpubl. m.s.). In Niue, an SPC masterfisherman averaged
45.3fishper hour over 16 trips during 1988 and 1989. In Yap, C. Friberg (Yap
Fishing Authority) reported an average of 23 flyingfish per hour were caught
based on 20 trips which took place during August and September. This project
was aided by an experienced Cook Island fisherman whose conclusion was that
catch rates in Yap were similar to those in the Cook Islands during the peak
season. There is one report of catching 15 to 20 fish per hour using a kerosene
lantern and household battery-powered torches in Solomon Islands (D. Ham,
pers. comm.). In Vava'u, catch rates from 19 trips between January 1987 to
November 1989 averaged 18.4 fish per hour using electric lights at night
(P. Mead unpubl. m.s.). In Fiji, Walton (1991) reports that initial trials held
outside of Suva Harbour entrance during August 1991, yielded roughly 10 fish
per hour. An experienced SPC masterfisherman from the Cook Islands assigned
to that project asserted that catch rates would probably be higher during the
warmer months (October to March). Despite promising indications from the
above tests, none of these fishing trials have resulted in the establishment of
viable fisheries.
DESCRIPTIONS OF FISHING METHODS FROM OTHER
REGIONS
Fishing practices from other areas of the world can provide some perspective
on alternative Pacific islands flyingfish fishing methods.
Flyingfish 189

Table III. Traditional Pacific Islands methods for catching flyingfish.


Location Method Source

Southern Scooping fish at night using dipnet Ministry of


Cook and torches Marine
Islands Resources, 1989

Scooping fish at night using dipnet and torches, Powell, 1989


hooking using floating lines with baited hooks. Mokoroa, 1984;
Catching fish by hand during occurence of inshore Powell, 1989
spawning aggregations.

Northern Scooping fish at night using dipnet and torches. Andrews,


Cook 1987;
Islands Beaglehole &
Beaglehole, 1938

Federated Scooping fish at night using dipnet and torches. Buck, 1950
States
of Micronesia

Tuvalu Scooping offish during day, scooping offish at Kennedy, 1930


night using dipnet and torches, herding fish into
apex of V-shaped net.

Scooping fish at night using dipnet and torches, Zann,1980


trolling using small hooks baited with coconut.

Tokelau Scooping fish at night using dipnet and torches. MacGregor, 1937

Kiribati Night fishing with torches and scoop nets when Turbott, 1950
the moon is full, night fishing with torches and scoop Lawrence, 1983
nets when there is no moon, fishing with torches and
scoop nets at sunset, fishing with hooks and floats, trolling.

Trolling using lure made of coconut mid-rib. Kennedy, 1930

Palau Scooping fish at night using dipnet and torch. Black, 1968;
Johannes, 1981
Marshall Scooping fish at night using dipnet and torch/lamps. Knight, n.d.
Islands

French Scooping fish at night using dipnet Bagnis el


Polynesia and head-mounted lamps. al., 1973
Catching fish by hand during occurence of inshore Johannes,
spawning aggregations. 1981

Solomon Fishing with carved floats, coconut mid-rib hooks, Wata, 1985
Islands and thin lines.

Mariana Catching fish using small hooks, thin lines, and Driver, 1989
Islands calabash floats.
190 Robert Gillett and James Ianelli

In the eastern Caribbean, flyingfish fishing is highly seasonal and targets on


the spawning behaviour of the principal species, H. ajjinis. Fishermen drift
alongside rafts of palm leaf or other vegetation which, together with a chum
basket containing chopped fish and fish oil held over the side, attract schools of
flyingfish. The flyingfish are then caught during the daytime with hand-held
dipnets or gillnets.
In the Indian Ocean, fishermen also take advantage of flyingfish spawning
aggregations. Some fishing is done with gillnets in conjunction with floating
vegetation, which presumably provide an attractive spawning habitat as a
lure. The general applicability of flyingfish fishing using gillnets is currently
being extensively tested in the Bay of Bengal (Pajot 1991).
In the Philippines, Martin (1938) described four methods of fishing for
flyingfish found there: using traps, large and small gillnets and purse
seine. Dalzell et al. (1990) describe the current use of gillnets and drive-in nets
used in the Philippines.
In Indonesia, fishermen collect the eggs of flyingfish off special bamboo
rafts; the eggs are dried and exported to Japan. A variation ofthese rafts also trap
the flyingfish which are sold on the local market (Basuki 1989).

CATCH STATISTICS

Pacific islands landings: Statistics on subsistence and artisanal fisheries are


often difficult to obtain and those which are available are sometimes
unreliable. This is especially true when dealing with flyingfish in the Pacific
islands. Data on catches of these fish are apparently only available for French
Polynesia, the Cook Islands, and Kiribati.
Service de la Mer (1989) indicates that 46poti marara fishing from the small
ports of Paea, Puunauia, and Arue, on the Island of Tahiti, caught a total of
213 t of flyingfish in 1988. It is estimated that there are a total of 272 poti
marara in French Polynesia. It would seem reasonable to assume that the
catches of those based furthest from the commercial centres would be smaller
than those of the boats from the three sampled ports because of the smaller
markets in more rural areas. Catches from the subsistence sector must also be
considered. A rough estimate of the total landings of flying fish from the 130
islands in French Polynesia might therefore be about 800 to 1,000 t.yr1.
In 1979 the Agriculture Unit of the Cook Islands government estimated that
46.11 of flyingfish were landed infiveislands in the southern Cook Islands. The
Ministry of Marine Resources (1989) indicated that subsistence and inter-island
trade of these fish average about 3 to 5 t.yr"' on Palmerston Atoll. Data supplied
to FAO (FAO 1989) show that in the years 1984 to 1987 the Cook Islands annual
catch was between 40 and 44 tonnes. Considering the more detailed data above
Flyingfish 191

and the active subsistence fishery for flyingfish in the northern Cook Islands, the
FAO figure appears low. About 60 t.yr"1 is perhaps a more realistic estimate.
Mees (1984) gives data from which estimates of flyingfish catches on 6 atolls
in the Gilbert Group of Kiribati can be made. These estimates range from 16 to
51 t per atoll per year (average 26 t.yr1). Extrapolating this to the 17 atolls of
the Gilbert Group gives a crude estimate of 453 t.yr1. Data supplied to FAO
(FAO, 1989) however, show that, in the years from 1984 to 1987, estimates of
flyingfish catches ranged from 161 to 2,455 t.
Anecdotal information from Tokelau, Tuvalu, the Marshall Islands, the
Federated States of Micronesia, Niue, and Palau indicate that active subsistence
fisheries for flyingfish exist in those locations. Although Melanesian catches
are likely to be small, at particular islands in Vanuatu, Solomon Islands, and
Papua New Guinea where there is a strong fishing heritage (e.g. Futuna, Belona,
Manus), flyingfish are taken. Apparently, few, if any, flyingfish are caught in
Fiji at present. Considering the above, a crude estimate of the total annual
landings of flyingfish in the Pacific islands is about 2,0001.
Catches from other regions: To give some perspective of flyingfish develop-
ment potential, it is useful to consider catches obtained from tropical flyingfish
fisheries in other parts of the world. In the Indian Ocean, recorded catches have
ranged from 5041 in 1986 to over 1,6001 by 1989 (FAO, 1991). Currently the
fishery is expanding through the use of gillnets on larger vessels. In the eastern
Caribbean, catches have steadily increased since the beginning of the fishery in
the late 1940s to present annual catches between 2,000 and 5,000 t. In the
Philippines, flyingfish catches have consistently been about 17,000 t per
annum. Indonesia reported annual catches of over 11,000 t from 1987-1989
(FAO 1991). Using the rough estimate presented in the previous section, less
than 6 per cent of the world landings of flyingfish are caught in the Pacific islands
region.

POST-HARVEST ASPECTS

Currently there are three known commercial markets for flyingfish


products. First in importance is the use of flyingfish for direct human
consumption. Flyingfish are also sold for bait, primarily for sport fishing. A
market also exists for flyingfish roe.
Regarding consumption, flyingfish is a nutritious food. It is high in protein
and, according to FAO (1990), has fewer calories and fat per gram than many
other fishes. Ciguatera poisoning has not been associated with the consumption
of flyingfish. Because flyingfish feed primarily on plankton and smallfish,they
are relatively low in the food chain and thus less likely to concentrate biotoxins
in their flesh.
192 Robert Gillett and James Ianelli

Flyingfish are currently processed as fresh, fresh frozen, salted/dried, and


smoked. Depending on the length of time required to deliver the flyingfish to
the market, they may be iced while at sea. In the Caribbean, a special "butterfly
fillet" is made whereby all bones are removed and a single one-piece fillet
remains. FAO (1990) produced a manual illustrating this filleting method, as
well as several aspects of handling and preserving flyingfish. Frozen flyingfish
are commonly filleted first and fast frozen in vacuum packs for maximum
quality. Preservation of flyingfish is particularly important in fisheries which
are highly seasonal.
Flyingfish represent a significant source of revenue in many areas. In the
Cook Islands, the 1990 price for a string of seven flyingfish sold for US$2.80
(approximately US$2.50 per kg). In the eastern Caribbean, the annual landed
value of flyingfish in recent years has been about US$5.5 million (Mahon
1989).
In a study of processingflyingfish,Herborg (1971) found that it was possible
to preserve flyingfish by marinating them for three months at room
temperature. The final product, presented in "pickled" form, has a shelf-life of
several months under refrigeration. Fish jerky made from tuna has recently
been successfully produced in the Tokelau Islands and requires minimal
technology1. This type of processing may be appropriate for flyingfish as
well. Such value-added products increase locally earned revenue.
Flyingfish are considered one of the best bait species for deepwater pelagic
sport fishing. Some claim that flyingfish can be up to ten times more effective
than artificial lures (e.g. P. Mead, 1991). They keep well when frozen and can
even be re-frozen successfully without becoming too soft to be useful. Mead
describes one technique for rigging and fishing with flyingfish as trolling bait
(ibid.). Preston etal. (1987) present another method.
There are fisheries for flyingfish eggs, especially in Indonesia. Fishermen
there use unique bamboo rafts that attract flyingfish during periods when they
are known to be actively spawning. The flyingfish deposit their eggs on the
rafts; the fishermen collect, dry, and eventually ship the eggs to Japan. Redmayne
(1989) reported that in the United States during 1988 whole flyingfish sold for
US$2.24/kg while the processed roe sold under its Japanese name, tobiko,
brought an average price of US$25.20/kg. He describes in detail the qualities
of flyingfish roe as a food item:
Tobiko...is a flaming fluorescent orange. The color is artificial, looks
artificial, and is intended to add fun to other food. Not only does tobiko
supply a psychedelic touch to sushi, but it adds hundreds of small mouth
explosions as the berries burst on your palate.. .color and pop are essential
to the product... Tobiko without the pop is like champagne without the fizz.
Flyingfish 193

IV. DISCUSSION

INDICATIONS OF FISHERY POTENTIAL

There are several indications that the Pacific islands flyingfish resources
have potential for further development.
On a biological basis, commercially important flyingfish species found in the
tropics appear to be short-lived, fast growing, highly fecund animals with a wide-
ranging distribution. This suggests that their populations should be fairly
resilient to fishing pressure.
Development potential is also suggested by the fact that areas where no
flyingfish fisheries exist have similar climates and geography to areas with
developed fisheries.
For example, French Polynesia has a substantial flyingfish fishery while in
Fiji, at the same latitude, virtually no flyingfish fishing occurs. There is no
indication that the productivity of the waters surrounding Tahiti is greater than
those surrounding Fiji.
Comparisons of Pacific islands flyingfish fisheries with fisheries outside the
region also reflect the potential of the resource. An estimated 6 per cent of the
world landings of flyingfish comes from the Pacific islands region, yet geo-
graphically, the Pacific islands region is substantially larger than the other
primary "regions" where flyingfish are caught (Indonesia, the Philippines, the
Indian Ocean, and the eastern Caribbean).
The magnitude of pelagic fish habitat can also be shown by comparing
catches of skipjack and yellowfin tuna, two species that are common where
flyingfish are found. Catches of these tuna species from Pacific islands waters
represent about 46 per cent of the world landings2 while flyingfish landings are
almost an order of magnitude smaller. This represents some evidence, albeit
circumstantial, that the capacity to harvest flyingfish on a sustainable basis in the
Pacific islands may be large enough to withstand increased fishing effort and
development.
There is no published evidence that flyingfish stocks in the Caribbean have
been overfished, even though the fishery has produced steady catches since the
late 1940s. Dalzell et al. {unpubl. m.s.) estimated the flyingfish exploitation
rates for the principal species in the Philippines to be over 50 per cent per annum,
suggesting that the stocks are very heavily exploited. Despite this, catches have
remained stable at over 17,000tsince 1987. Some researchers caution however,
that flyingfish populations may behave similarly to clupeoids such as the
Californian sardine or the Peruvian anchovy and may collapse when a threshold
level of exploitation is exceeded.
194 Robert Gillett and James Ianelli

SUGGESTED FUTURE WORK

Taxonomy: In order to address fundamental questions on population dynamics


and biology of flyingfish, a reliable method of identifying the species is
required. A taxonomic revision of the family followed by the publication of a
catalogue of flyingfish species is needed.
Tagging: The tagging of flyingfish could help resolve certain constraints to
development. For example on Temana Atoll, Kiribati, where gear restrictions
have been established due to perceived local over-exploitation, a tagging study
similar to that carried out in the Caribbean may resolve the need for local
regulation by revealing the range of the target species.
In areas where the fishery is developed, such as the Cook Islands and Tahiti,
tagging studies would also be useful to estimate the overall productivity of the
resource. This information would be useful in determining the need for man-
agement with a view to maximizing long term catches.
Investigations on markets for flyingfish: In areas where local consumers are
unfamiliar with flyingfish, a programme may be needed to introduce the
product. This might consist of supplying the market with flyingfish for trial
sales and, if needed, providing cooked samples for potential consumers to
sample.
Areas in which flyingfish are abundant but where the market is limited would
benefit from investigation of alternative products. The processing of flyingfish
for long-term storage through appropiate technology such as pickling or drying
may increase the marketability of flyingfish and add value. These methods
would be particularly applicable in outer island areas without freezers.
Flyingfish have proved to be a superior bait for trolling but their utility as
longline bait has yet to be tested. Quantitative trials of flyingfish as tuna longline
bait are needed to gauge its effectiveness. Presently, the standard imported
saury tuna longline bait sells for about US$2.70/kg and its availability has been
a major constraint to small-scale longline fishing in the Pacific islands.
Demonstrations of development potential: In areas where flyingfish are not
caught, the abundance and seasonality of the flyingfish resource need to be
assessed. Following this, demonstrations of the fishery potential to local
fishermen are needed. This could consist of making fishermen aware of the
relatively low capital expense required to start fishing for flyingfish and the
possible advantage of this fishery as a supplement to other fishing activities. The
actual fishing methods also need to be demonstrated. This could be carried out
through a masterfisherman extension programme. Alternatively, a video of
fishing methods could be produced.
Investigations into alternative fishing methods: There are several successful
flyingfish fishing methods from other areas that may be worth trying in the
Pacific islands region. The practice of "chumming" flyingfish schools during
Flyingfish 195

their spawning season, as is done in the eastern Caribbean, may warrant


investigation. This method would require less fuel than the current commercial
fishing method and may be possible to carry out in conjunction with other
fisheries such as tuna handlining.
In the Indian Ocean and Indonesia, the flyingfish fisheries also take advan-
tage of flyingfish spawning aggregations. Adapting fishing methods which
target spawning aggregations of flyingfish in the Pacific islands region thus
appears to be an area of research that may be fruitful. It should be noted,
however, that successful fishing methods for a particular species in one area may
not necessarily work in another area, even for the same species. In many
instances fish behaviour is specific to certain areas and environmental condi-
tions.
With regards to fishing for flyingfish using lights at night, some innovation
on different lighting systems may be useful.

V. CONCLUSIONS
There appears to be potential for the development of flyingfish fisheries in
the Pacific islands. This contention is based on what is known of the biology of
the fish, locations in the Pacific islands where considerable development has
occurred, and comparison with other regions of the world. Development of a
flyingfish fishery for small-scale fishermen seems possible because of the
relatively low technology required and, in many cases, the proximity and
apparent abundance of the resource. Development is presently constrained by
the difficulty in properly identifying flyingfish species, the limitations of
existing markets, and the lack ofawareness of the flyingfish resource and fishing
techniques.

ACKNOWLEDGEMENTS
Hugh Walton and Mose Pelasio are to be thanked for their assistance during
this project. Paul Mead, David Itano, Foua Toloa, Andrew Wright, and Mike
Savins generously provided flyingfish samples. Drs. B. ColletteandN. Parin
identified the various species. Several people provided information
including: Louise Wrobel, Randy Veterli, Robert Mahon, Hazel Oxenford,
Robert Pitman, and Kurt Schaefer. Michael Batty, Edward Lovell, Jay Smith,
and Sarah Langi made useful comments on drafts of this report.

NOTES
1. SPC(1991)ReportonSPCcoastal fisheries programme activities in 1990/1991. Work-
ing Paper 5,23rd Regional Technical Meeting on Fisheries, South Pacific Commission,
Noumea, 20 pages.
2. Based on FAO (1991) world landings and estimates reported in the Regional Tuna
Bulletin, First Quarter, 1990. South Pacific Commission.
196 Robert Gillett and James Ianelli

REFERENCES

Abe, T. (1978). Keys to the Japanese fishes. Hokuryukan, Tokyo, 358 pp.
Agricultural Planning Unit. (1979). Statistics ofartisanal fishing in the southern group 1978.
Statistical Working Papertio. 13, Ministry of Agriculture and Fisheries, Rarotonga. 38
pp.
Agricultural Planning Unit. (1980). Statistics ofartisanal fishing in the southern group 1979.
Statistical Working Papertio. 15, Ministry of Agriculture and Fisheries, Rarotonga. 38
pp.
Amesbury, S., Gushing, F. and Sakamoto, R. (1986). Guide to the coastal resources of Guam
- Volume 3 - fishing on Guam. Contribution Number 225, University of Guam Marine
Laboratory, University of Guam Press, 110 pp.
Andrews, G. (1987). Marine ecology survey of Pukapuka Atoll. Commonwealth Science
Council and Australian Institute of Marine Science, Townsville. 133 pp.
Bagnis, R., Mazellier, P., Bennett, J. and Christian, E. (1973). Poissons de Polynesie. Les
Editions du Pacifique, Papeete, 368 pp.
Basuki, M. (1989). Suitable gears to collect the eggs of flyingfish. 5 pp. unpubl. manuscript.
Beaglehole, E. and Beaglehole, P. (1938). Ethnology of Pukapuka. Bulletin 150, Bernice P.
Bishop Museum, Honolulu, 119 pp.
Black, P. (1968). The fishing lore of Tobi. 134 pp. unpubl. m.s.
Blanchet, G. and Borel, G. (1988). The "pod marara" or the success of a small fishing-boat
fitted to many needs. Working Paper, World Symposium on Fishing Gear and Fishing
Vessel Design, 7 pp. Food and Agriculture Organisation, Rome.
Breder, C. and Rosen, D. (1966). Modes of Reproduction in Fishes. Natural History Press,
Garden City, New Jersey. 308 pp.
Buck, P. (1932). Ethnology of Tongareva. Bulletin 92, Bemice P. Bishop Museum,
Honolulu.
Buck, P. (1932). Ethnology of Manihiki and Rakahanga. Bulletin 99, Bernice P. Bishop
Museum, Honolulu. 238 pp.
Buck, P. (1950). The material culture of Kapingamarangi. Bulletin 200, Bernice P. Bishop
Museum, Honolulu. 291 pp.
Chakalall, B. (1992). Flyingfish from the Caribbean. INFOFISHInternational 1/92,48-50.
Collette, B., McGowan, G., Parin.N. and Mito, S. (1984). Belonifonnes: development and
relationships. In Ontogeny and Systematics of Fishes - based on an international
symposium dedicated to the memory of Elbert Halvor Ahlstrom. Special Publication
Number 1, American Society of Ichthyologists and Herpetologists.
Dalzell, P., Corpuz, P., Arce, F. and Ganaden, R. (1990). Philippine small pelagic fisheries
and their management. Aqu. Fish. Manag. Vol. 21, 77-94.
Dalzell, P., Dumlao, R. and Pauly, D. (unpubl. m.s.). Fisheries biology of Philippine
flyingfish (Families: Exocoetidae and Hemiramphidae). ICLARM Contribution, Small
Pelagics Management Project, International Center for Living Aquatic Resources
Management, Philippines. 35 pp.
Dalzell, P., Lindsay, S.R. and Patiale, H. (1991). Fisheries resource survey of the island of
Niue. A report prepared in conjunction with the South Pacific Commission Inshore
Fisheries Research Project and the FAO South Pacific Aquaculture Development
Project for the Government of Niue, July 1990.
Flyingfish 197
Driver, M. (1989). The account of Fray Juan Pobre's residence in the Marianas 1602.
Micronesian Area Research Center, University of Guam, 33 pp.
Eastern Caribbean Flyingfish Project. (1988). News Bulletin. Number 3, January-June 1988,
Bellairs Research Institute of McGill University, Jamestown University, Barbados, 11
pp.
Eastern Caribbean Flyingfish Project. (1989a). News Bulletin. Number 4, July-February
1989, Bellairs Research Institute of McGill University, Jamestown University, Barba-
dos, 8 pp.
Eastern Caribbean Flyingfish Project. (1989b). News Bulletin. Number 5, March-August
1989, Bellairs Research Institute of McGill University, Jamestown University, Barba-
dos, 14 pp.
Fabres, B. (1985). Bibliography on flyingfish biology and related topics. In Development
Strategies for Flyingfish Fisheries of the Eastern Caribbean, (R. Mahon, H. Oxenford
and W. Hunte, eds.) pp. 43-50. International Development Research Center, Canada.
Food and Agricultural Organization. (1983). FAO Species Identification Sheets: Fishing
Area 51 (W. Indian Ocean). Food and Agriculture Organization of the United Nations,
Rome.
Food and Agricultural Organization. (1989). FAO Yearbook- Fishery Statistics Catches and
Landings 1987. Food and Agriculture Organization of the United Nations, Rome, 490
pp.
Food and Agricultural Organization. (1990). Flyingfish: the art of fishing and processing.
FAO Technical Cooperation Project. TCP/RLA/4509(T). 32 pp.
Food and Agricultural Organization. (1991). FAO yearbook-fishery statistics catches and
landings 1989. Food and Agriculture Organization of the United Nations, Rome, 490 pp.
Gorelova, T. (1980). The feeding of young flyingfishes of the family Exocoetidae and of the
smallwing flyingfish, Oxyporhamphus micropterus, of the family Hemirhamphidae../.
Ichthyol. 20(4), 60-71.
Hall, D. (1955). Recent developments in the Barbadian flying-fish fishery and contributions
to the biology of the flying-fish Hirundichthys affinis (Gunther 1866). Fishery Publi-
cation No. 7,1955, London, 41 pp.
Herborg, L. (1971). Possibilities for marinating flyingfish (Exocoetidae) and robins
(Carangidae). UNDP/FAO Caribbean Fisheries Development Project, Barbados. DP/
CAR/REG/189. 5 pp.
Hornell, J. (1922). The flying-fish fishery of the Coromandel Coast. Madras Fishery
Department, Report No. 4 ofl922. Madras Fishery Bulletin, Vol. XV, 99-112.
Imai, S. (1959). Studies on the life histories of the flyingfish found in the adjacent waters
of Japan - 1 . Mem, Fac. Fish. Kag. Uni. Kagoshima, Japan, 1-85.
Imai, S. (1960). Studies on the life histories of the flyingfish found in the adjacent waters
of Japan - II. Mem. Fac. Fish. Kag. Uni. Kagoshima, Japan, 8-45.
INFOFISH (1989). INFOFISH Trade News. No. 2/89, Kuala Lumpur.
Inoue, Y., Tawara, Y. and Matsuo, M. (1986). Diurnal migration offish schools relevant to
set-net. Bull. Jap. Soc. Sci. Fish. 52, 55-60.
Inshore Fisheries Research Project. (1988). Workshop of Pacific Inshore Fishery Re-
sources. South Pacific Commission, Noumea, 80 pp.
Inshore Fisheries Research Project. (1989). The marine resources of Palmerston Atoll, Cook
Islands. SPC Inshore Fisheries Research Project Technical Report No.2, and FFA
Research Coordination Unit Report No. 89/16, South Pacific Commission, Noumea, 57
pp. (draft).
198 Robert Gillett and James Ianelli

Johannes, R. (1981). Words of the Lagoon: Fishing and Marine Lore in Patau District of
Micronesia. University of California Press, Berkeley, 245 pp.
Jordan, C. (n.d.). The flyingfish industry of Trinadad and Tobago: a continuing case study.
4 pp. unpubl. m.s.
Kailola, P. (1987). The fishes of Papua New Guinea: a revised and annotated checklist -
Volume 1: Myxinidae to Synbranchidae. Research Bulletin No. 41. Research Section,
Department of Fisheries and Marine Resources, Port Moresby, Papua New Guinea. 194
pp.
Kennedy, D. (1930). Fishing for flying-fish (fai isave). In Field Notes on the Culture of
Vaitupu, Ellice Islands. J. Polyn. Soc. 39(1), 61-70. Avery Press, New Plymouth.
Knight, J. (n.d.). Flyingfish jojo. In Guide Book to the Marshall Islands. 30 pp.
Koch, G. (1983). The Material Culture of Tuvalu. Institute of Pacific Studies, University of
the South Pacific, Suva. 215 pp.
Kojima, S. (1969a). Studies on the migration and spawning habit of the flyingfish,
Cypselurus opisthops hiraii Abe - II - behavior of the school approaching land for
spawning. Bull. Jap. Soc. Sci. Fish. 35(3), 284-288.
Kojima, S. (1969b). Studies on the migration and spawning habit of flyingfish - III - behavior
of the fish schools in the spawning ground. Bull. Jap. Soc. Sci. Fish. 35(11), 1055-1059.
Kojima, S. (1971). Studies on the migration and spawning habit of flyingfish - IV -
distribution of eggs in the spawning ground. Bull. Jap. Soc. Sci. Fish. 37(4), 284-288.
Kovalevskaya, N. (1982). Superfluous reproduction and development of flyingfishes of the
family Exocoetidae../. Ichthyol. 22(4), 48-54.
Lawrence, R. (1983). Temana. In Atoll Economy: Social Change in Kiribati and Tuvalu, 99-
108. Report No. 4, Development Studies Centre, Australian National University,
Canberra. 261 pp.
Lewis, J., Brundritt, J. and Fish, A. (1962). The biology of the flyingfish Hirundichthys
affinis (Gunther). Bull. Mar. Sci. GulfCaribb. 12(1), 75-94.
MacGregor, G. (1937). Ethnology of Tokelau. Bulletin 146, Bernice P. Bishop Museum,
Honolulu. 183 pp.
Mahon, R. (1989). Fishery management options for the lesser Antilles. Field Document 4,
FAO, FI:TCP/RLA/6776. 117 pp.
Mahon, R. and Mahon, S. (1987). Seasonality and migration of pelagic fishes in the eastern
Caribbean. In: Report and Proceedings of the Expert Consultation on Shared Fishery
Resources ofthe Lesser Antilles Region. FAO Fisheries Report .No. 383,192-273. FAO,
Rome.
Mahon, R., Oxenford, H. and Hunte, W. (1985). Development strategies for flyingfish
fisheries of the eastern Caribbean. Proceedings ofanlDRC-sponsored Workshop at the
University of the West Indies, Cave Hill, Barbados, 22-23 October 19H5. International
Development Research Centre, IDRC-MR128e, 148 pp.
Martin, C. (1938). The flyingfish industry of the northwestern and southwestern coasts of
Cebu. Philipp. J. Sci. Vol. 67, 177-187.
Matsuda, M., Ainaoka, K., Araga, C , Uyeno, T. and Yoshino, T. (1984). The Fishes of the
Japanese Archipelago. Tokai University Press, Tokyo. 379 pp.
Mead, P. Draft report of masterfishermen gear development project. South Pacific Commis-
sion, B.P. D5 Noumea, New Caledonia, unpubl. m.s.
Mees, C. (1985). The Fisheries of Marakei Island. Fisheries Division, Ministry ofNatural
Resource Development, Kiribati, 23 pp.
Flyingfish 199
Mees, C. (1984). The fisheries of Temana and Arorae. Fisheries Division, Ministry of
Natural Resources Development, Tarawa, Kiribati. 14 pp.
Ministry of Marine Resources. (1989). The fish and fishes of Palmerston Atoll. Rarotonga,
Cook Islands.
Moana, A. Report of second visit to American Samoa, July 1988. Deep Sea Fisheries
Development Project, South Pacific Commission, Noumea, unpubl. m.s.
Mokoroa, P. (1984). 'Maroro-Tu: the flyingfish return.' n Atiu - an island community.
Institute of Pacific Studies, University of the South Pacific, Suva, 114-125.
Monte, S. (1965). A contribution to the study of the flyingfish reproduction (Hirundichthys
affinis). Boletin Inst. Biologia del Marino, Universidade Federal do Rio Grande do
Norte, Vol. 1,1-15.
Munro, I.S.R. (1967). The Fishes of New Guinea. Department of Agriculture, Stock, and
Fisheries, Port Moresby, 651 pp.
Nesterov, A. and Bazanov, S. (1986). Vertical distribution of flyingfishes (Exocoetidae).
Voprosy Ikhtiologii, No. 4, 685-687.
Nichols, J.T. and Breder, CM. (1935). New Pacific flyingfishes collected by Tenipleton
Crocker. Am. Mus. Novit. 821, 1-4.
Nystrom, U. (1989). Capture of flyingfish - techniques old and new. Bay of Bengal News,
Issue Number 36, December 1989, pp. 20-21.
Olson, R.J. and Boggs, C.H. (1986). Apexpredationby yellowfintuna (Thunnusalbacares):
independent estimates from gastric evacuation and stomach contents, bioenergetics,
and cesium concentrations. Can. J. Fish. Aquat. Sci. 43, 1760-1775.
Ouchi, O. (1985). A story of Yap Island. Asahi Newspaper Company, Tokyo, 258 pp.
Oxenford, H. (1985). Synopsis of the biological data on the four-winged flyingfish
Hirundichthys affinis Gunlher. In Development Strategiesfor Flyingfish Fisheries ofthe
Eastern Caribbean, (R. Mahon, H. Oxenford and W. Hunte, eds.), 51-88. International
Development Research Centre, Canada.
Oxenford, H. (1988). Preliminary tagging results on the four-wing flyingfish Hirundichthys
affinis in the Eastern Caribbean. 41st Annual Meeting, Gulf and Caribbean Fisheries
Institute, St. Thomas, 13 pp.
Oxenford, H., Mahon, R. and Hunte, W. (1989). Relative abundance and distribution of
adult flyingfish in the eastern Caribbean. 42nd Annual Meeting, Gulf and Caribbean
Fisheries Institute, Ocho Rios, Jamaica, 10 pp.
Pajot, G. (1989). Flyingfish Gillnet. Bay of Bengal Programme, Food and Agriculture
Organisation, Madras. 3 pp.
Pajot, G. (1991). Capture of flyingfish: Findings of BOBP trials and studies. Bay of Bengal
News, Issue Number 41, Bay of Bengal Programme, Food and Agriculture Organisation,
Madras, 4 pp.
Parin, N. (1960a). Distribution of flyingfish in the western and central parts of the Pacific
Ocean. Trudy Instituta Okeanologii, 41, 153-162, [in Russian]. Translation No. 52,
Ichthyological Laboratory, U.S. National Museum, Washington, 15 pp.
Parin, N. (1960b). The flyingfishes (Exocoetidae) of the northwest Pacific. Trudy Instituta
Okeanologii, 31, 205-285, Akademiya Nauk SSSR, [in Russian]. Translation IPST
Catalog No. 618, Office of Technical Services, U.S. Department of Commerce,
Washington D.C.
200 Robert Gillett and James Ianelli
Parin, N. (1961a). Contribution to the knowledge of the flyingfish fauna (Exocoetidae) of
the Pacific and Indian Oceans. Trudy Instituta Okeanologii, 42,40-91. Translated from
the Russian by Laurence Penny, Ichthyological Laboratory, Fish and Wildlife Service,
U.S. National Museum, Washington, D.C.
Parin, N. (1961b). A new species of the flyingfish from the western Pacific - Cypselurus
vitiazi, sp. Nov. (Pisces, Exocoetidae). Trudy Instituta Okeanologii, 42, 1412-1416.
Parin, N. (1961c). The basis of classification of flyingfishes (families Oxyporhamphidae
and Exocoetidae). Trudy Instituta Okeanologii, 43, 92-183.
Parin, N. (1983). Assessment of the abundance of flyingfish by visual observation. Biol.
Oceanogr. 2(2-4), 341-355.
Paxton, J., Hoese, D., Allen, G. and Hanley, J. (1989). Zoological Catalogue of Australia
- Volume 7-Pisces: Petromyzontidae to Carangidae. Australian Biological Resources
Study, Australian Government Publishing Service, Canberra.
Powell, R. (1989a). The evolution of the boats used at present in the Cook Islands for
catching flyingfish. 21 pp. unpubl. m.s.
Powell, R. (1989b). The evolution of nightfishing gear for catching flyingfish in the Cook
Islands. 13 pp. unpubl. m.s.
Powell, R. (1989c). Notes on the spawning of flyingfish at Atiu, Cook Islands. 3 pp. unpubl.
m.s.
Preston, G., Chapman, L., Mead, P. and Taumaia, P. (1987). Trolling techniques for the
Pacific islands. Handbook'No. 28, South Pacific Commission, Noumea, 162 pp.
Redmayne, P. (1989). Flyingfish and tobiko (roe). Seafood Leader, March/April 1989,53-
58.
Service de la Mer. (1989). Bulletin statistique du secteur de la mer annee 1988. Ministere
de la Mer, Polynesie Francaise.
SPC. (1980-85). South Pacific Commission's Skipjack Survey and Assessment Programe
Final Country Reports No. 1-19. Noumea, New Caledonia, par. var.
Storey, K.W. (1983). Aspects of the biology and fishery of the flyingfish, Hirundichthys
affinis, at Barbados. MSc. Thesis, University of West Indies, Barbados, 161 pp.
Suda, A. (1973). Development of fisheries for nonconventional species../. Fish. Res. Board
Can. 30(12), Part 2, 2121-2158.
Titcomb, M. (1952). Native use offish in Hawaii. MemoirNo. 29,Suppl. J. Poly. Soc. Avery
Press, New Plymouth, 162 pp.
Turbott, I. (1950). Fishing for flyingfish in the Gilbert and Ellice Islands. J. Poly. Soc. 59(4),
349-367.
Walton, H. (1991). Preliminary investigation of flyingfish fishing, Suva, Fiji, July 23 -
August 24, 1991. FAO/UNDP Regional Fishery Support Programme 91/46. 7 pp.
Wass, R. (1984). An annotated checklist of the fishes of Samoa. NOAA Technical Report
SSRF-781, U.S. Department of Commerce, 43 pp.
Wata, A. (1985). Traditional use and knowledge of marine species in Santa Ana Island,
Makira, Solomon Islands. Fisheries Division, Ministry of Natural Resources, Honiara,
Solomon Islands. 7 pp.
Watson, D. (1990). Life history of flyingfishes (Exocoetidae) in the Flores Sea, Indonesia.
[abstract of Ph.D. dissertation], University of Guelph, 2 pp.
Wrobel, L. (n.d.). Tableau especes Marara en Polynesie Francaise. EVAAM, 1.
Yacapan, N. (1991). Growth, mortality and recruitment of short-wing flying fish
Oxyphorhampus micropterus in Bohal Sea, Philippines. Fishbyte 9(2), 10-12.
Flyingfish 201

Zann, L. (1980). Tuvalu's subsistence fisheries. Effects of Energy Crisis on Small Craft and
Fisheries in the South Pacific, Report 4, Institute of Marine Resources, University of the
South Pacific, Suva, 13 pp.
Zuyev, G. and Nikolsky, V. (1980). Procedure of quantitative recording of flyingfish
(Exocoetidae)../. Ichthyol. 20(6), 147-149.

You might also like