Environmental Biology of Fishes Vol. 24, No. 4, pp.

295-300, 1989
0 Kluwer Academic Publishers, Dordrecht.

Pelagic spawning of the hawkfish Oxycirrhites typus (Cirrhitidae)

Terry J. Donaldson 1,4& Patrick L. Colin2,3

’ Section of Ichthyology, Museum of Natural Science and Department of Zoology and Physiology, 119 Foster
Hall, Louisiana State University, Baton Rouge, LA 70803, U.S.A.
2Motupore Island Research Station, University of Papua New Guinea, Papua New Guinea
3 Present address: Caribbean Marine Research Center, cfo Florida State University Marine Laboratory, Rt. 1,
Box 456, Sopchoppy, FL 32358, U.S.A.
4Reprint requests.

Received 30.3.1988 Accepted 22.7.1988

Key words: Behavior, Courtship, Eggs, Habitat availability, Site-attachment, Social group, Spawning ascent

Synopsis

Pelagic spawning of the deep slope coral-dwelling cirrhitid Oxycirrhites typus was observed for two social
groups at Papua New Guinea. This species was previously reported to be a demersal spawner in an aquarium.
Courtship in social groups consisting of a single male and one or two females commenced just prior to or after
sunset among the branches of gorgonian or antipatharian corals. Males and females occupied separate corals;
males either visited females at their corals or met them at an adjacent coral just prior to courtship. Courtship
was sequential and consisted of two or more bouts with each female that culminated in a rapid ascent into the
water column and the release of floating eggs. Fertilized eggs, taken from a third social group, were spherical
and averaged 0.69 mm in diameter. Spawning pairs sought refuge in their resident corals or in the coral where
courtship occurred immediately after spawning was completed.

Introduction Thresher (1984) questioned demersal spawning

The long-nosed hawkfish Oxycirrhites typus is a
distinctive tropical to sub-tropical cirrhitid distrib-
uted from Baja California, Mexico west to the Red
Sea. This species is typically found on deep slopes
of coral and rocky reefs and inhabits gorgonian and
antipatharian corals at depths of 25-100 m (Randall
1963). There is some published information con-
cerning its reproductive behavior. Takashita (1975)
and Tanaka et al. (1985) briefly described apparent
courtship behavior in aquaria. Another observa-
tion suggested the possibility that this species might
lay demersal eggs (reviewed in Thresher 1984).
This suggestion has been cited elsewhere (i.e. Ran-
dall 1985).
of this species as a result of field observations of
pelagic spawning by another cirrhitid, Cirrhitich-
thys oxycephalus, in the Gulf of California. He
believed the existence of two modes of spawning
within a small, monophyletic family of 34-35 spe-
cies was unlikely. Additional species of cirrhitids
have been spawning pelagic eggs, including Cirrhit-
ichthys falco (Donaldson 1986, 1987), C. aprinus
(Donaldson, unpublished), C. aureus (Y. Yogo,
personal communication), and Paracirrhites arca-
tus, P. forsteri and Cirrhituspinnulatus (Donaldson
unpublished). Recently, we observed courtship
and pelagic spawning of Oxycirrhites typus near
Motupore Island, Papua New Guinea, where this
species occurred in relatively shallow (20-26m)
depths.
296
Methods and study sites
Observations of courtship and spawning were
made using scuba between 6-15 November, 1986 at
times to include sunset and dusk (1730-1830 h).
Additional observations of behavior and habitat
use were made during occasional daylight dives
within these dates. Data were recorded on plastic
slates, and underwater photographs of courtship
and spawning were taken. Time of sunset was de-
termined from meteorological tables. Water vis-
ibility was estimated from known distances to sub-
jects at approximately 20-25 m. Spawning was ob-
served at two sites located about 16 km SE Port
Moresby. The first site was located on the slope of a
broad fringing reef ca. 500m south of Loloata Is-
land at a depth of 22-23 m. This site had poor water
visibility, ca. 3-6m and water temperatures be-
tween 2627” C. The second site was located at part
of the Papuan Barrier Reef, locally called Horse-
shoe Reef, ca. 6 km south of Motupore Island.
Water temperature was 285°C.
Body sizes were estimated in the field by mea-
suring the distance between two points on the sub-
stratum relative to the observed position of the tip
of the snout and the edge of the caudal fin for a
given individual. Fertile eggs were obtained by col-
lecting adults with quinaldine-alcohol solution
shortly before (within one hour) spawning. Fish
were returned to the laboratory alive, their body
sizes measured, and their gametes stripped by gen-
tle hand pressure shortly after sunset (when they
would have naturally spawned) the same evening.
Gametes were mixed in a petri dish containing
seawater and later embryos that hatched were mea-
sured using an ocular micrometer. Eggs, embryos
and larvae were preserved in 3% formalin and
specimens deposited in the Australian Museum,
Sydney (AMS) .
Results
Three social groups were studied. Courtship and
spawning were observed from a group consisting of
a male (72 mm TL) and two females (71 and 67 mm
TL, respectively) at the Loloata Island site and
from a pair (male- 65 mm TL; female- 55 mm TL)
at the Horseshoe Reef site. In the former group,
the male and the largest female lived among the
branches of separate antipatharian corals; the
smaller female lived on a large Subergorgia sp. sea
fan located less than 0.5m away from the larger
female’s coral. In the latter group, both male and
female were observed at 23-26m in two Suber-
gorgia sp. sea fans, located 3m apart. Gametes
were taken from a second pair collected from an
antipatharian coral near the Loloata Island site
(male- 99.5 mm TL; female- 92 mm TL). Courtship
and spawning of 0. typw was observed off Loloata
Island on five consecutive nights, 6-10 November,
1986 and at Horseshoe Reef, 15 November. Court-
ship began 23 minutes before to 22 minutes past
sunset at Loloata Island and 13 minutes past sunset
at Horseshoe Reef.
The male at the Loloata Island site left his coral
and made his way, either swimming between the
coral branches or along the bottom, to the first
female, located less than 0.5 m away. Once there,
he waited, dorsal fin erect, until the female arrived
from the upper branches of her coral. Occasional-
ly, the male swam into the upper branches of the
coral in search of the female. As the male ap-
proached, the female swam toward him, dorsal fin
erect. Daily courtship consisted of several bouts of
activity during a period which ranged from 12-19
(x = 5.3) minutes in duration. Generally, the male
and female circled horizontally or vertically around
one another among the branches of the coral and
occasionally settled upon the branches to rest. The
pair rested by aligning their bodies l-3 cm apart in
parallel or anti-parallel position (see Donaldson
1986, 1987, for description of cirrhitid motor pat-
terns). As courtship progressed, the female led the
male in short swimming movements (‘lead and fol-
low’; Donaldson 1987) through the branches of the
coral, each movement separated by periods, 3-5
seconds in duration, of parallel or anti-parallel rest.
During rest periods of longer duration the male
moved his body close to the female and nudged her
abdomen or flank below the dorsal fin with his
snout (Fig. 1). Then the male often mounted the
female by placing his body along her upper flank,
adjacent to or touching her dorsal fin; balance was
 297

Fig. 1. Nudging behavior by a male Oxycirrhites typus (the male is the lower fish).

Fig. 2. Descent after pelagic spawning by a pair of Oxycirrhires fypus.The female is partially obscured by the edge of the coral.
298
maintained by extending a pectoral fin against her
body while he beat his caudal fin vigorously. The
male also swam from his position parallel to the
female, looped to pass in front of her and exposed
his flank perpendicularly to her snout, before com-
ing to rest parallel to her opposite flank. Just prior
to spawning, the pair assumed parallel or near-
parallel positions, lifted their snouts upward by
raising their bodies with extended pelvic fins,
raised their dorsal fins and fin spines, and quickly
swam up or out from the coral to a point 20-25 cm
away. There, they turned their snouts downward,
flexed their bodies rapidly, and released a small
cloud of gametes, which remained visible for 2-3
seconds in the fading light; no egg predation was
observed although egg predators, usually poma-
centrids, were present nearby. Immediately after
spawning, the pair quickly swam back to the
branches of the same coral (Fig. 2) and briefly
assumed a position parallel to one another before
the male departed.
After spawning with the first female, the male
swam to the second female on the upper edge of the
Subergorgiu sea fan OSm away. Daily courtship
bouts with the second female commenced 4-12
minutes past sunset and lasted 2.1-9.7 minutes be-
fore spawning occurred. On one occasion, the first
female did not spawn with the male and in turn he
did not join the second female, who remained on
the edge of the sea fan until total darkness had
fallen (ca. 1840 h). Courtship and spawning behav-
ior was the same as with the first female, although
fewer bouts of courtship (x = 3.3) were observed
per period before spawning. On one exception (6
November), however, the male and the second
female were observed at parallel rest, their bodies
1 cm apart, on the edge of the sea fan. There, they
positioned their vents out past the edge over open
water and both quivered together during three pe-
riods of l-2 seconds each over the space of one
minute. Gametes may have been simultaneously
released by each during these brief periods. After-
wards, the pair repositioned themselves a few cm
away and repeated the behavior four more times in
succession without ascending into the water co-
lumn.
Large predatory fishes were observed in close
proximity to the courting 0. typus, and it is possible
that the pair spawned from the edge of the sea fan
rather than making a spawning ascent above the
coral and exposing themselves to the risk of preda-
tion. Alternately, these behaviors may merely have
been a demonstration of male responses to an ap-
parent lack of readiness on the part of the female,
After spawning had been completed with the
second female, the male slowly swam back to his
coral by way of the first female’s coral; the second
female sought shelter within the sea fan.
Courtship and spawning behavior of 0. typus at
Horseshoe Reed was similar, although the onset of
courtship was after rather than before sunset. Prior
to the onset of courtship, both male and female
rested and fed in separate Subergorgia sea fans.
Sometime just after 1730 h (32 minutes before sun-
set), the female moved slowly from her sea fan and
swam upward along the reef slope, past the male’s
sea fan and two adjacent but shallower sea fans,
and eventually moved into the branches of a large
antipatharian coral ca. 6 m from her starting posi-
tion. This coral was located at the base of a steep
reef face at 23 m depth. At 1800 h the male traced
the female’s path to the large antipatharian coral
where the female was sitting. However, the male’s
progress was much slower as he moved from one
point of cover to the next before arriving at 1812 h.
Courtship commenced at 1815 h (13 minutes past
sunset) when the male ascended into the upper
branches of the coral and joined the female. Court-
ship consisted of two bouts, each in separate parts
of the coral, and was brief (3 minutes and 4 seconds
total duration) compared with courtship durations
of the Loloata Island social group. After the
spawning ascent, the male and female moved into
shelter within the branches of the coral.
Fertilized eggs taken from the third social group
were spherical and averaged 0.69mm in diameter
(+ 0.23; N = 17), slightly larger than that of ripe
ova of C. oxycephalus reported in Thresher (1984).
Eggs taken later from a fourth group, collected 5
May, 1987 and stripped at 2000 h, hatched after 15
hours incubation at 27°C. Attempts to rear the
embryos were unsuccessful.

Discussion
Courtship and spawning behavior of 0. typus was
similar to that described from field studies for the
cirrhitids (Thresher 1984, Donaldson 1986, 1987).
Aspects in common included: (1) onset of courtship
just before or after sunset with spawning after sun-
set, (2) successive courtship and spawning by the
male in social groups containing one or more fe-
males, (3) courtship and spawning at a site common
to both the male and female or at a site located
within the female’s home area, (4) pelagic spawn-
ing with a rapid but relatively short ascent into the
water column by both male and female, (5) the
return of the male and female to resident corals
after spawning, except at Horseshoe Reef where
the pair moved into shelter in the same coral used
for courtship. Pelagic spawning at or near dusk is
found among many tropical reef fishes. Spawning
ascents into the water column above the substra-
tum allow the release of planktonic eggs some dis-
tance above the bottom, minimize immediate egg
predation by benthic organisms, and facilitate
transport of eggs away from the spawning site (see
Thresher 1984 for review). Spawning adults are at
risk from piscivores, but this threat may be reduced
by rapid dashes to minimal heights where eggs can
be released (Moyer 1987). Oxycirrhites typus
spawning was rapid (ca. l-2 seconds in duration)
and ascent lengths were relatively short (<0.25
m). Observed ascent lengths were low compared to
other cirrhitids (C. oxycephafus, 0.3-l.Om,
Thresher 1984, personal observation; C. fulco, 0.2-
0.6m, Donaldson 1986,1987; and C. uprinus, 0.3-
0.6m, Paracirrhites arcatus, 0.4l.Om, P. forsteri,
1.0-2.5 m, and Cirrhitus pinnulatus, 1.52.0m,
personal observation), and may reflect the initial
distance from the reef where the spawning ascent
starts from a large gorgonian or black coral. Al-
though planktivorous and piscivorous fishes were
present at both spawning sites, no attempted pre-
dation on spawning adults or released eggs were
observed. At the Loloata Island site water visibility
was considerably less and this, because of reduced
ability to detect predators, could have influenced
the distance moved by fishes between resident cor-
als and spawning sites. At Loloata the male trav-
299
elled only 0.5 m between corals while at Horseshoe
Reef the male and female moved slightly over 10 m
to the shallower spawning site although both resid-
ed in corals located less than 2 m apart. Movement
between corals by 0. typus at Horseshoe Reef was
slower compared with those at Loloata but the
distances travelled were much greater. Movement
of 0. typus at Horseshoe Reef consisted of quick
dashes between patches of cover (corals, rocks and
epifauna) alternating with periods of rest under
cover for up to five minutes. Both male and female
maintained erect dorsal spines while moving. This
behavior has been observed in other cirrhitids, in-
cluding C. f&o (Donaldson 1987) and C. aprinus
(personal observation).
Popular accounts of 0. typus suggest that this
strongly site-attached species occurs in pairs, may
be monogamous, and seldom, if ever, leaves the
gorgonian or antipatharian coral where it resides.
In Papua New Guinea, this species occurred in
social groups consisting of a single male and one or
two females. This species used one or more corals
within a given area, moved freely between those
corals, and formed pairs prior to courtship. Al-
though the social organization and reproductive
behavior of only a few social groups were described
here, the patterns of organization and reproduc-
tion were consistent with those observed for other
cirrhitids (Thresher 1984, Donaldson 1986, 1987).
Social organization and reproductive behavior of
0. typus may depend upon habitat availability (i.e.
suitable corals). The subsequent acquisition of this
habitat is by: (1) emigrating adults, in search of
potential mates, or by (2) newly-settled larvae,
which may be incorporated into existing social
groups when they reach maturity.
Acknowledgements
We thank G.B. Constantino, J.M. Fitzsimons, J.T.
Moyer, J.E. Randall, and R.E. Thresher for their
comments on the manuscript or useful discussion.
L.J. Bell, A.E. Trumble and M.C. Wilkins are
thanked for their kind assistance. The comments of
two anonymous reviewers are appreciated. Re-
search by the senior author was generously sup-
ported by an award from the Coypu Foundation.
Field operations were funded by the University
Research Committee, University of Papua New
Guinea.
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