Flyingfish predators, prey and

research methods: Lessons

learned in the eastern Caribbean
• Hazel A Oxenford
• Centre for Resource Management and Environmental
Studies (CERMES), University of the West Indies,
Cave Hill Campus, Barbados
 Flyingfishes – Exocoetidae
Order: Beloniformes
8 genera 67 species

• Cheilopogon • Fodiator
• Cypselurus • Hirundichthys
• Danichthys • Parexocoetus
• Exocoetus • Prognichthys

Source: Fishbase. Photo by C. Jesson

 Two-winged flyingfishes

Fodiator (2 spp)

Exocoetus (5 spp)

Parexocoetus (3
spp)
 Four-winged flyingfishes

Cypselurus (12 spp) Hirundichthys (8

spp)

Prognichthys (6
Cheilopogon (30 spp)
spp)
 Distribution and characteristics
of flyingfishes

Source of image: SeaWifs

• Global distribution in tropics • Most restricted to single
and subtropics ocean (some circum-global)
• Strictly epipelagic • Small (most < 30 cm)
• Oceanic and neritic species Capable of gliding flight
•
 Importance to global fish yield
• Majority of species (> 30) support small scale
fisheries
• Annual landings globally > 73,000 mt
• Best known flyingfish fishery is in Barbados
• The target species is the Atlantic fourwing
flyingfish (Hirundichthys affinis)
 Traditional fishery
in Barbados

• Ligon 1651
• Hughes 1750
• Schomburgk 1848
• Up to 1950s
sailing boats
Image removed
Economic importance of flyingfish to
Barbados
• 300 day-boats, 150 ice-
boats
• 1,100 fishers, 450
persons in post harvest
Economic importance of flyingfish to
Barbados

• Landings > 2,000 mt

per year
• Represent > 60% of
annual fish landings
• Ex-vessel value of US
$ 2.65 million
• 6 fish processing
companies
Cultural importance of flyingfish to Barbados

• National dish
• Dollar coin
• Definitive stamps
• Promotionary material

Source: Fishbase
Cultural importance of flyingfish to Barbados

Barbados Port Authority

Cultural importance of flyingfish to Barbados

Image removed

A way of life for many Barbadians

 Fishing Techniques in Barbados
• Surface floating gillnets
• FADs (“screelers”)

screelers

Courtesy of the Fisheries Division

 Fishing Techniques in Barbados
• Dipnets
• Chum basket

bait basket screeler

Courtesy of the Fisheries Division

Predators

Marine
mammals
• dolphins
• sea lions

Sea birds
• boobies
• noddies
• frigate birds

After Parin 1968

 Prey

• small finfishes
• pteropods
• ostracods
• amphipods
• decapods
• chaetognaths
• ascidians
• siphonophores hyperiids
• salps

euphausiids copepods

After Parin 1968

 Flyingfish prey studies
Flyingfish species Common name Prey items Age Tropic References n
level
Exocoetus monocirrhus barbel flyingfish planktonic copepods, pelagic gastropods, juvenile Gorelova 1980
radiolaria, salphidae, chaetognatha
Prognichthys sealei sailor flyingfish planktonic copepods, ostracods, gastropods juvenile Gorelova 1980
Hirundichthys affinis fourwing flyingfish small finfishes, planktonic copepods, euphausidsadult Hall 1955, Lewis et al 1962 441, 293
amphipods, chaetognaths, siphonophores, Barroso 1967 317, 931,
gastropods, pteropods, squids, salps, fish eggs, Cruz & Araujo 1971
macroalgae fragments, marine insects
Cheilopogon cyanopterus margined flyingfish planktonic crustaceans, other zooplankton juv/adult Heemstra and Parin 1986
Parexocoetus brachypterus sailfin flyingfish zooplankton juv/adult Lewis 1961
Masuda and Allen 1993
Cheilopogon heterurus Mediterranean flyingfish planktonic copepods, euphausiids, larvae 3.1 Lipskaya 1987 17, 22
fish eggs/larvae
Exocoetus obtusirostris oceanic two-wing flyingfish ascidians, planktonic copepods larvae 3 Lipskaya 1987 20, 10
Exocoetus volitans tropical two-wing flyingfish planktonic copepods, ascidians larvae 3.0 - 3.1 Lipskaya 1987 70, 10, 4, 3, 150
Hirundichthys speculiger mirrorwing flyingfish planktonic copepods larvae 3 Lipskaya 1987
Cheilopogon agoo Japanese flyingfish zooplankton juv/adult Masuda and Allen 1993
Cheilopogon doederleinii n/a zooplankton juv/adult Masuda and Allen 1993
C. pinnatibarbatus japonicus n/a zooplankton juv/adult Masuda and Allen 1993
Cypselurus hiraii n/a zooplankton juv/adult Masuda and Allen 1993
Cypselurus poecilopterus yellow-wing flyingfish zooplankton juv/adult Masuda and Allen 1993
Cheilopogon rapanouiensis Easter island flyingfish small finfishes, zooplankton juv/adult Parin 1996
Cheilopogon furcatus spotfin flyingfish finfishes, zooplankton juv/adult Parin 1999
Cheilopogon intermedius n/a small finfishes, zooplankton juv/adult Parin 1999
Cheilopogon pitcairnensis n/a small finfishes, zooplankton juv/adult Parin 1999
Cheilopogon spilonotopterus stained flyingfish small finfishes, zooplankton juv/adult Parin 1999
Cheilopogon spilopterus manyspotted flyingfish small finfishes juv/adult Parin 1999
Cheilopogon suttoni Sutton's flyingfish small finfishes, zooplankton juv/adult Parin 1999
Cypselurus hexazona n/a small finfishes, zooplankton juv/adult Parin 1999
Cypselurus oligolepis largescale flyingfish small finfishes, zooplankton juv/adult Parin 1999
Cypselurus opisthopus black-finned flyingfish small finfishes, zooplankton juv/adult Parin 1999
Cheilopogon pinnatibarbatus Bennett's flyingfish zooplankton, finfishes juv/adult Parin and Gibbs 1990
C. pinnatibarbatus melanocerusn/a zooplankton, planktonic crustacean juv/adult Paulin et al 1989

Primary source: FishBase

 Detailed diet salps
salps
larval fish
larval fish

composition
Hirundichthys affinis copepods
other copepods
other
crustaceans
Early studies used crustaceans Hall 1955
frequency of occurrence
in stomachs
larval fish

Lewis 1964
 Feeding Habits
• Feed exclusively on small zooplankton
• Actively hunt mostly for small finfishes,
copepods and other crustaceans
• Also eat other planktonic invertebrates
• Can be considered tertiary-level feeders
• Feed mostly at night
• Feed actively through the spawning season
Competitors
For Euphausiids:
• fishes of the ‘shifting
layers’
• small deep water fishes
(Chiasmodon)
• Tuna

For Copepods:
• euphausiids
• shrimp
• shifting layer fishes
• hyperiids
• myctophiids
• molas
Hyperiids
For Hyperiids:
• myctophids
• small deep water fishes
ephausiids copepods
• tunas
• snake mackerels

After Parin 1968

 Research Methods
• Eastern Caribbean Flyingfish Project
• Funded by IDRC
• 5-year duration (1986-1991)
• University of the West Indies and Bellairs
Research Institute
• Fishery Divisions of 7 countries
• 3 scientists, 4 MPhil students
 Research Methods
• Distribution and abundance of adults,
juveniles, larvae and eggs
• Movements
• Age determination
• Spawning behaviour
• Genetic population structure
 Distribution and Relative
Abundance
Adults and Juveniles
• Acoustic gear
• Visual survey
• Night-lighting
• Predator gut contents
• Fishery catch data

Eggs and Larvae

• Neuston net tows

Survey vessel
 Sonar Technology
Adults and juveniles

• Hull mounted fish finder

(vertical and side scan)

• Surface towed sonar

(looking up, looking down)

Conclusion
Significant failure!
Visual Survey
Adults and juveniles
 Visual Survey

Distribution and
size of flyingfish
patches

Image

From: Oxenford et al. 1995a

removed
 Visual Survey
Relative abundance
of all flyingfishes
across the eastern
Caribbean

From: Oxenford et al. 1995a

 Visual Survey
Parexocoetus brachypterus
From: Oxenford et al. 1995a

Cheilopogon cypselurus
 Visual Survey

Note: Conclusions
• Assumes a constant • Provide reliable data on
proportion of the school relative abundance
flies • Easy and relatively
• Close correlation with inexpensive
other indices (dipnet • Suitable for individual
data) species
• Objectivity could be
improved with additional
instrumental recording
(e.g. video)
Night-Lighting and Dipnetting
Adults and juveniles

Image removed

Traditional flyingfish fishing in Tokelau

 Night-Lighting and Dipnetting

• 200 watt spot lamps

• 5 mm mesh dipnet
• 4 replicate stations
• 40 mins each

From: Lao 1989

Arrangement of nightlights
on small fishing vessel
 Night-Lighting and Dipnetting

• Partitioning by species

From: Oxenford et al. 1995b

From: Oxenford et al. 1995b

Exocoetus volitans

Hirundichthys Exocoetus volitans

affinis
 Night-Lighting and Dipnetting

• Partitioning by size

From: Oxenford et al. 1995b

From: Oxenford et al. 1995b

Parexocoetus brachypterus

small juveniles large juveniles

Night-Lighting and Dipnetting
Comparing distribution of four-winged
flyingfishes among years

graphic removed

From: Pitman et al. 2002

 Predator Gut Contents: Adults and juveniles

image removed

From: Storey 1983

Monthly size frequency of

Hirundichthys affinis sampled by
the fishery (gillnet and hooks)
and by dolphinfish predators

from dolphinfish
from fishery
 Fishery Dependent Catch Data: Adults

Grenada (3 yr)

Martinique (13 yr)

Barbados (7 yr)

Dominica canoe

Dominica (1 yr)

From: Hunte 1987

St. Lucia (3 yr)

MONT HS

Barbados day launch

Neuston Net Tows: Eggs and larvae

From: Oxenford et al 1986

 Neuston Net Tows: Eggs and larvae
Exocoetus volitans

From: Hunte et al 1995

Day collections
• Larvae more abundant
Parexocoetus brachypterus
• only small larvae captured
 Adult Movements: Tagging Hirundichthys affinis

Off Barbados
• Mulloney 1961 (n = 762)
• Lewis 1964 (n = 1,288)
Off Brazil
• Barroso 1967 (n = 552)

Results
• Local recapture of 1 – 6.4%
• Time-at-large up to 50 days From: Barroso 1967
 Adult Movements: Tagging

• 7,019 flyingfish tagged • Tagged and released

• across the eastern
Floy fingerling tags Caribbean
• Hand-stitched in 30 sec • 1988 and 1989 fishing
• Captured by dipnet and seasons
barbless hook
Photo by H. Oxenford
 Adult Movements: Tagging

Public awareness campaign

 Adult Movements: Tagging
Results
• 4.5% recaptured
• Mean time-at-large: 3 weeks,
maximum:121 days
• Greatest recorded
displacement: 200 nmi
• Fastest speed > 16 nmi/day
• 10% recaptures showed
transboundary movement
• Transboundary movements
same for males and females

From: Oxenford
• Greater movement by maturing

1994
fish than running ripe or spent
fish
• No tag returns in following
season
 Age Determination: Adults and juveniles

Otolith growth checks

• Otoliths must be ground
and polished
hatching
• Viewed under 500 x check

• Good agreement
between readers
• Presumed daily rings
clearly visible in lapilli of
juveniles and adults Lapillus otolith from
Hirundichthys affinis
• Rings cannot be
counted beyond 120
 Age Determination
Validating daily rings
• Marking captive adults with OTC
• Marking, tagging and releasing (n = 946)

Tagged fish marked with OTC

 Age Determination
Validating daily rings
Laboratory rearing
• Hatched in 3-6 days
• Fed on brine shrimp

Otoliths in newly hatched H.affinis larva

Collecting eggs of Hirundichthys affinis

Flyingfish larvae
 Age Determination
Validating daily rings
Laboratory rearing
• Sacrificed every day for counting rings
• Maintained in laboratory for 52 days
• Rings and days strongly correlated

From: Oxenford
1994

Laboratory reared larvae of

Hirundichthys affinis
Otoliths growth checks vs known age
 Age Determination: Further validation
Modal progression Radio chemical dating
• Fishery independent • Th-228/Ra-228 radio
sampling isotopic pair

Conclusions
• Otoliths show daily rings suitable for
aging juveniles
• Larvae easy to hatch in laboratory
• Juvenile growth is fast (1.4 mm/day)
From: Oxenford

• Growth rate slows markedly after

1994

sexual maturity
• Radiochemical dating feasible but
expensive
Von Bertalanffy growth curve for
• Fishing gear very size selective
Hirundichthys affinis
Reproductive Characteristics:
Spawning behaviour

image removed

Flyingfish spawning on surface substrate (screeler)

 Spawning Behaviour

The mystery?
• Scarcity of spawning
substrata (flotsam)
• Scarcity of eggs
• Newly hatched larvae
swim upwards
• Anecdotal evidence

From: Oxenford et al. 1996

from fishers

The test
• Provided substrates at
different depths
• Provided surface
substrates for sinking
Experimental spawning substrata
moorings
 Spawning Behaviour
Results
• Flyingfish did spawn on
substrata at 20 m depth

From: Oxenford et al. 1996

• Strong preference for
surface

Results
• Flyingfish spawned on all
3 substrata
• Experiment inconclusive
as insufficient time
 Population Genetics
Mitotypes for mtDNA of Hirundichthys affinis

From: Gomes et al. 1999

• 360 fish
• mtDNA RFLPs
• gDNA RAPDs

Frequency of composite mitotypes of H. affinis

 Population Genetics

From: Gomes et al. 1999

• Consistent results for • 3 distinct unit stocks
RFLPs and RAPDs • Shared management
• Clear separation of in eastern Caribbean
populations