BULLETIN OF MARINE SCIENCE,3l(3): 558 573, l98l

ASPECTS OF SEXUAL REPRODUCTION AND LARVAL

DEVELOPMENT IN THE SHALLOW WATER
HERMATYPIC CORAL, GONIASTREA AU STRALEI'{ SIS
(EDWARDS AND HAIME, 1857)

Barbara L. Ko.iis and Norman J. Quinn

ABSTRACT
Sexual reproduction and larval developmenl in Goniastrett au.strttlensi's, a shallow water
hermatypic coral, were studied from September 1977 to November 1980 on Heron Island
reef. Australia. G. australensis is a simultaneous hermaphrodite with ovary and testis in-
termingled on the same mesentery. Gonad development occurred synchronously within and
between colonies. An annual spring spawning season of approximately 2 days was observed'
Gametes were released between 1600-1800 h on neap low tides between the full and last
quarter moon phases. Fertilization is probably external; eggs from a single polyp were re-
leased in clumps held together by mucus and sperm were freely released. Egg clumps were
negatively buoyant and sticky, adhering to objects when contact was made. Sperm and eggs
were usually released simultaneously from the same polyp. Laboratory experiments indi-
cated that self-fertilization was possible with apparently normal larval development and
settlement. In aquaria ciliated larvae developed in approximately 2 days and settlement
commenced 17 days after spawning. Reproduction began in colonies with an arithmetic mean
radius of 1.5-2.0 cm, aged 4+ Years.
While G. australensis did not planulate, the mode and timing of spawning and larval
development substantiated Stimson's (1978) hypothesis that the mode of reproduction in
hermatypic corals may be related to habitat, but not his prediction that corals characteristic
of the reef flat would begin to sexually reproduce at an early age and planulate.

It has been assumed from studies of sexual reproduction in hermatypic corals

that they are viviparous and release planulae (Hyman, 1940; Wells, 1954). How-
ever, it Las been noted that the release ofplanulae has not been observed in most
species of coral, including species in which gonads were present at some time
during the studies (Connell, 1973; Stimson, 1978).
The most comprehensive report of sexual reproduction in a non-planulating
hermatypic coral was the study of Favia pallida (Marshall and Stephenson, 1933)
(nomenclature of Faviidae is in accordance with Veron and Pichon' 1976). The
presence of gonads in November and absence in December 1928 indicated a
ielatively brief spawning period, contrasting with the long, often year-round'
period oi larval rilease found in known planulating species, and suggested that
gamete release and external fertilization may have occurred.
Stimson (197S) synthesized information from his own and previous studies on
the presence or abience of planula release and habitat distribution of corals and
compared reproduction among corals in shallow and deep water. He hypothesized
that the mode of reproduction in hermatypic corals is related to habitat and
predicted that corals characteristic of the reef flat would begin to sexually repro-
huce at an early age and characteristicaliy planulate while deep water corals
would have alternate modes of reproduction, probably seasonal gamete release.
The sexual reproduction, larval development, distribution and abundance of
Goniastrea austialensis were studied on Heron Island, Australia, to augment the
meager data on the life history of hermatypic corals and to test Stimson's hy-
poth-esis. G. ctustralensis (Fig. 1) is a massive hermatypic coral widely distributed
in the Western Pacific (Veron, 1974 Yeronetal.,1977). It has been recorded as
558
 KOJIS AND QUINN: SEXUAL REPRODUCTION IN GONIASTREA .559

lOcm
I

Figure I Reefflat colony of (]oniastrea australensis on Heron Island reef

present in a variety of habitats from the reef flat to the deeper water
of the reef
slope (veron et al., 1977). on Heron and wistari Reefs of t'he capricorn Group,
Great Barrier Reef (.23"27's, r5r"155'E), G. austrarensrs is largeiy restricted io
major shallow water habitats: lagoon, inner and outer reef flai and crest. It is
rarely found on the reef slope and then only in water less than 5 m deep.

M,qrE,Rrer_s eNo ME-rnoos

Gonads
Reproduction was studied from September 19'77 to November 1980 on the reef
flat at Heron lslancl.
Ten colonies (>15 cm diameter) at each of three different sites (total :o cotonieg
were inoiviauailf
tagged on the inner reef flat (Fig' 2). Sites were more than 200 m apart
trl-oioy7 cletermination of
variation in the timing and mode of reproduction within the same general truuitut.
In addition, large
untagged colonies from the outer reef flat, crest and lagoon were impled
to assess variation in the
99{e and timing of reproduction with habitat. Coloniei *er. ,urnpled'rnon,nty o"t*..n Septembei
1917 and November 1978.by re^moving samples of approximately j
to zo poifps -itt a hammer and
chisel' Samples were fixed in 10vc sea watei formalin in the fielcLand 24 h'lateitransferred
to a fresh
solution of l07o formalin for storage.
P^reserved whole samples were tlecalcifiecl in Gooding and Stewart's
solution and initially treated
as follows: (1) single polyps were embedded in "Tissue Prep," sectioned
at a nominal thickness of
8 pm, stained either with Mayer's hematoxylin and eosin or Fieidenhain's
hematoxylin, and examined
for the presence ofgonads; (2) other polyps from the same samples were then ciissected
-aiumeter) and observed
using a binocular dissecting microscope. It was found that eggs
1z-s:o fnl and .sferm
clusters could be seen in both calcifiecl and decalcified potypr iittt. rn.r.rii".i.,
were removed ancl
gently squashed on a slide. Eggs ancl testes nearing maturity were visible
in the field with the naked
eye when the mesenteries were exposed
Five entire colonies with ripe gonacls were repeateclly split and examined to determine
whether
gonad development was synchronized throughout a colonv.

Spawning
Additional colonies from the reef flat were maintainecl in aquaria with unfiltereci,
running sea water
and plankton netting filtering the run-off from the outlets. Colonies on the
reef ffat were plriodically
sampled to determine if they spawned at the same time.
The convention ofnrrmhsling days according to the lunar cycle has been adoptecl
purposes (Atoda, 1947a; Stimson, 1978), i.e. new moon is lunar
for comparative
day l: first quo.i.r, clay g; fuil moon,
day 15: last quarter, day 22.
 560 BULLETIN OF MARINE SCIENCE,
VOL. 3I, NO. 3. I98I

Figure 2. -----........-
Map of sample and transect
sites on Heron Island reef_

Larval Settlement and Development

time
."[r"'::r#iir.:lr"r:5;r^trate -from spawning to development of mobire,
--- -.- 'rom aquaria the ciriared rarvae, a'
ia. morning forio*ing ,pu',''ni"r,l."ri"rln. spawn in the aquar_
In 1979, ciriatetr rarvae were transferred
to and .n.11^r",*d in 2.-4.rclean plastic
fl:i;,.*liH:f,:l;:::,:l-ti.J;..;r ;;;;"1'Xou,,.," were ven,irared ro,prorideaquaria containins
air and increase
uor o," uni ,ia;: ;i';;;;;J?";il:li;",s";:;x
film was placed over lhe tops t*1j:l;1,i'-;.ffiJx,;.T;
ti.ili;'_xT and keep the conrainers
olaquaria,;..1;;; evaporaljon
In 1979' anolher laboratory trial was ru.r.-,o-,r.,...rine.rhe inlervai crean.
menr and whelher normar^raivae
,j.";i.;.;;;;1'eir-tenitizea ;.;;;; spawning and settre_
with singre coronies lserr-fertirizeJf .r*, irnr ,"")i,u.u".
covered 2J large-mouth jars ";;il';;:ll ."r"ri"r-roi:;;bj;'J.";ri:rtirized) from
were
aquaria
and p."rro.J *it'lr'uii'io. ,r,or, herd in
aquarra were conditioned in sea i.,L.uulr.,priJ.,io,pru.rng rarvae
water fo^eu"rul duy' on." inside,
the polvps were fed bv .rnning
y"fili;;r." *?rJlir,.
tu-uJ r,u;;il; and metamorphosed,
wirh potyps were cteaned periodica,y
b;,.;;;;;; aquaria fo.;;;";';;-urs dairy. The aquaria
To compare rates of a.".rop."nl, ule". r;i;;,,il;,1;:
r.i"".'*#'?.rr;"t.d";Ji.o- ,rJ';;;j;;i:'rr...undingfingerrips.

5#:i'.i::i;!"#::i,:U:',*llt';*:ru"*"#;i;;"'T;.'x^';'JT:, coronies
and were-.,u*.,"0,
samprer of rarvae were coile*J.
n^.j'i.':-+{!r,.ioto.nya. in phosphate
water' \lored in 0 r M rnH 7 2,t pnorpt'rr.
r,uri.,. ui.i.ru.o,l.o i" sprli:, buffer or firtered sea
il.jir]m. semi_trr;n :ecrions
1,",i fi"iliii,;5:lx:l *lil*r;ffl#ilil1.i,il,T;:,ylii,l#'.j,ii,. ; ,.it.,r,-'j,, ji
Colony Size and Reproduction
To determine the approximate
size at the onset of reproduction,
sampled. The arirhmeric mean
raoiu.'ix;;i';;t;;;.r.in,iru l0r colonies were measured and
w,a.s r"""i,il"n-'.^suring the heighr
l
KOJIS AND QUINN: SEXUAL REPRODUCTION IN CONIASTREA 561

Table 1. The arithmetic mean radius (i), standard deviation (SD), and coefficient of variation (CV%)
for repeated size samples of 6 colonies of Goniastrea rtuttrolensis

Colonies

i(cm) I .89 3.67 5.63 6.21 6.56 7.56

SD 0.07 0. l5 0.29 0.10 0.21 0.11
cv% 3.'7 4.1 5.2 1.6 3.2 1.5

(distance from the point of attachment to the tip of the colony), length (greatest diameter at right
angles to the height) and width (greatest diameter at right angles to both the height and length).
Wooden calipers and ruler were used to measure coral colonies. The calipers prevented damage to
coral skeletal projections.
Hughes and Jackson (1980) found that partial colony mortality distorted the linear relationship
between size and age among reef corals. Partial colony death commonly occurred tn G. australensis
(personal observation). Thus the arithmetic mean radius (i) was used in preference to the geometric
mean radius (G-) (Loya, 1976b) because the G* decreases in relation to x as differences in l, w and
h measurements increase. While we used x to consider the effect of age, present data indicate only
the effect of size on reproduction, irrespective of age, and do not indicate whether age will induce
the onset of reproduction or increase fecundity at a smaller size than would occur if growth were not
impaired.
Six in sitzr colonies were consecutively measured eight times by the same person to determine the
error inherent in the size measurements. The colonies were measured four times on one day and four
more times the next day. The mean error aI Ihe 957o confidence level was +0.3 cm (Table 1). No
signiflcant difference between the variance of the measurements was observed (F Test; P > 0.05).
The mean coefficient of variation was3.17o with measurements of large colonies being as variable as
small colonies. Variation in colony measurements was probably associated with the errors in mea-
suring irregularly shaped colonies.
Each measured colony was sampled for gonads shortly before spawning. When gonads were abun-
dant, samples were examined in the field: when there were few or no gonads, samples were examined
using a dissecting microscope. Each sample was assigned a gonad index value based on the criteria
in Table 2.

Distribution, Abundance and Size

Five transects were made perpendicular to the beach, extending from the beach rock, across the
reef flat to the inner edge of the crest (Fig. 2). All coral colonies l0 cm either side of the transect
were identified to species or genus and the distance each colony paralleled the line was measured. A
20 cm wide transect was used to reduce the effect ofpatchiness on measurements ofdistribution and
abundance. Additionally, the length, width and height of all colonies of G. australensis in Transect
2 were measured.

Table 2. Female and male gonad abundance values for Goniastrett australensis

Gonad Female Gonads: Male Gonads:

Abundance Average No. of Average Size of
Values Eggs Mesentery ' Testis Mesentery-'

0 Non-reproductive Non-reproductive
0.5 <1 Few mesenteries with testes
1.0 t-<3 Surrounding 1-3 eggs or equivalent in size
1.5 3-<5 Surrounding 3-5 eggs or equivalent in size
2.0 <-1 Surrounding 5-7 eggs or equivalent in size
2.5 1-<9 Surrounding 7-9 eggs or equivalent in size
3.0 9-13 Surrounding 9-13 eggs or equivalent in size
 562 BUI,I,ETIN OF MARINE SCIENCE. VOL. 3I. NO, ]. 198I

u.
1
&
&' F"
f;
'ry,

Figure 3. Polyps of Goniastrea australensis split through the center exposing oocytes and sperm
clusters: o, oocytes; sc, sperm clusters.

Rr,sulrs
Gonads
G. uttstralen.tir is a simultaneous hermaphrodite with ovary and testis inter-
mingled in the same mesentery (:gonad) (Figs. 3 and 4) (Favia pallicta; Marshall
and Stephenson, 1933). Gametogenesis occurred over approximately 9 months of
the year. In most colonies oocytes were first observed in March, except in one
instance when they were present in a sample collected in the beginning of Feb-
ruary. oocytes did not begin developing in all mesenteries of a colony concur-
rently, although nearly all mesenteries of sexually mature colonies contained ova
shortly before spawning. As vitellogenesis proceeded, the eggs changed in color
from white to brown. The mean diameter of 20 eggs measured in 197g immediately

{}-153.
*' t *,

Figure 4 Cross section through the mesenteries of a polyp of Goniustrea uustrulen5is showing male
and female gonads approximately 2 weeks before spawning: m, mesentery; n, nucleolus; nu, nu'al"ur;
o, oocytes; sc, sperm clusters.
 KOJIS AND QUINN: SEXUAL REPRODUCTION IN
GONIASTREA .563

Goniastrea ctustralensis in
Table 3. Tide timesl fbr Heron Island during the spawning periotl of
1918, 1979 and 1980

1978 lg'7g* l98o

t'4oon' Spaun
-'i"g N4oon\ Iimeof Spawn
Moon' Ti..of slu*-
PiJg" "fi; Dare .fimeo[
r"*iiJt {ge -Dare Lowlide ins
Age Date ro* t,,1"
15 16 Nov. 15.11 NA 15 5 Nov. l5.03 15 24 Ocr. 14.33

l6 17 Nov. 15.43 NA 16 6 Nov. 15.24 NA 16 25 Oct. 15.18

t7 Nov. 16.15 A 17 7 Nov. t6.24 NA 11 26 Oct. 16.05

18
19 Nov. 16.52 A F 18 8 Nov. t7.07 NA 18 21 OcI. 16.54 t
18
19 9 Nov. 18.00 A 19 28 Oct. 17.50 AF
t9 20 Nov. t7.37 AF
20 NA
20 21 Nov. 18.43 A 20 10 Nov. 19.09 A R 29 Oct. 19.03

21 22 Nov. 19.58 A 2l ll Nov. 20.30 A 21 30 Oct. 20.30 NA

22 23 Nov. 2r.05 A 22 12 Nov. 21.35 A 22 31 Oct. 21.45 NA
23 24 Nov. 21.58 A 23 13 Nov. 22.24 A 23 I Nov. 22.39 NA

@ervedinthefie1d;R'remnantsofspawningobservedinthefield:NA.co1oniesin
aquaria did not spawn
6:;;;i;;; oiirittout' and lvlarine Queensland Ausrratia leTS. ls7e l980
ro rt'i tuni' calendu u'ed bv Aroda {.ls47r'
'" 5;i"-;;;;;;;;il1."-o'oine
i:ii"f,';.;:";;;;ii'rt *inoip*'."rto \pawnins obt'rvation\,on rhe reel frat
+ Fieli.ample' indiiated thar 'ome colonie' \pawneJ on lhr\ oate

present until late Sep-

after release was 510 g,m (450-530 pm).Testes were not
tember/earlY October.
1978 had ripe
Five colonies examined for intra-colony synchrony in Nov€mber
in nearly all polyps, including those
go".A, i' approximatety ttre same_abundance
regions
polyps bordering
ilth";Jg;;tthe colonies. Gonads were only absent in
iamaged or severely infected by algae'
^-;il;i";it"?onaO
recently
development synchronized in all colonies, was apparent rn
throughout one year from individually tagged colonies'
-;;,ht sampi;s 1uken in.tudlng
All colonies oUrerueJ, tagged tolonies at all three sites and untagged
coloniesfromtheouterreefflat,crestandlagoon,showedSynchronyofgonad
development.
SPawning
and 1919 and
G. australensis released ova and Sperm in November 1977,1978,
on 6 November had
Orrrirrg tn" end of October 1980. In i977, samples collected
was observed in
ripe gonads; no gona;;;re present on- 6-Declmber' Spawning
gametes
and in the field (.1978;1980). In all observations
iiSTA ; liTS;1980)
were released duringih; uit"-oon low tide commencing at approximately
"i""?i" 1600
h EST, with the maj.ority being released
t u.,O until 1800
jl. ln 1,'9'78,the bulk of spawning.in the field probablv
"ppio^i-"t"ly
berween 1600-1700 rrdJr"
"o.rti.ruing
was observed in
occurred between t3lZg November. Spawning in this species
mid-January 1977
the field at Lord Uowe Island at approiimatelt 1630 h during
(T. Donne and L. Zell, personal communication)'
spawned annually' In tagged sampled over
Sexually mature
successive years ";i;;i;t
(N : tO,- tsTl, 19'78, l97g;N : 20' -c-olo-nies gonads were
1977',1978)
pr"r"rt i" all samples at itre beginning of November but absent at the end of
^Novemberibeginning
- of December'
stained,
i" nxea in firmalin immediately after spawning, sectioned and prob-
";; vesicle *"r pt"r"tt and ceil division had not begun.
g"iil"i""i Thus ova
"o O"r"f"ped shortiy Lefore spaw_ning. and fertilization could have occurred
was seen
prior to, Ouiing or after spawning. No distinct fertilization membrane
"UfV
at any stage.
l
BULLETIN OF MARINE SCIENCE, VOL, 3I, NO. 3, I98I

4tfn 's*
; d*"*U

Ir
w

Figure 5. Goni'strea australen.si.s, eggs and sperm being released from polyps: e, eggs; s, sperm.

During a single daily spawning period, ova and sperm may be released simul-
taneously (Fig. 5) or separately. Field colonies spawned in i or 2 days. In con-
trast, colonies held in aquaria for 3 or more days prior to spawning released
ga,metes for up to I week, though not on every day. Addition-ally, po"lyps from
different sections of the same colony spawned on different days. Lengthy-spawn-
ing in aquaria may have been an experimentally induced artifact.
In the field and in the aquarium, eggs were retained in a mucous matrix on the
surface of each polyp (Fig. 5) until the polyp had completed egg release. The egg
clt'mps were sticky and negatively buoyant adhering to the subiiratum on contact,
often adhering to the short strands of filamentous algae surrounding many of the
colonies (Fig. 6). Egg color was similar to the sediment trapped between algal
strands and to the foraminiferan epifauna. Clumps free in the water sank slorily
and in still water rested on the sand surrounding spawning colonies (Fig. 7j.
caught in currents, eggs were carried until they came in cJntact with the sub-
stratum and adhered.
Light microscope observations of live sperm indicated that sperm had a single,
motile flagellum and triangular head, and were released unconitrained insteal of
in discrete packets as occurs in some alcyonarians (Gohar, 1940).

Larval Development and Settlement

rn 1979, eggs from all aquaria developed into mobile, ciliated larvae in approx-
imately 2 days irrespective of whether the eggs were cross-fertilized or rilf-f".-
tilizred. water temperature was measured and compared in the aquaria and on the
reef flat on 2 days and no significant difference was found (diily range 23.5-
27.5"c). Larval development was similar in aquaria and on the reef flat.
Larval development was similar to the anemone Arictmsia palliata (Gemmill,
1920). Developing larvae of G. australensis became ciliated and mobile at the
,

565
KOJIS AND QUTNN: SEXUAL REPRODUCTION IN GONIASTREA

Figure 6. colony of Goniastreu ttu.strulensis on the reef flat surrounded by recently spawned eggs
adhering to the substratum: e, eggs.

gastrula stage. Larvae sampled 65 h after spawning.had small scattered mucous

llanAs in the zone of future ectoderm. By day 7, larvae had a well developed
lctodermal layer separated from the embryonic endoderm by a layer of mesoglea,
a stomodeum and mesenterial filaments. Mucous glands were large and abundant
in the ectoderm, especially in the thickened aboral region. Larvae sampled on
days 12, 15, l7 and 22 showed less rapid, but continuing, development. Zooxan-
thellae were not observed in eggs or larvae.
In 1978, larvae were held in aquaria for 23 days' but none settled.In 1979'

ff..!. ,*
'..t
.P )

lr'',
F
tr* ,1crn,
-n
Figure 7. Negatively buoyant egg clumps resting on the sand: ec, egg clumps
l
566 BULLETIN OF MARINE SCIENCE, VOL' 3I' NO. 3' 198I

Developing larvae of GoniLstrea australensis adhering to algae and sand:

dl, developing
Figure 8.
larvae.

three larvae which had developed from self-fertilized eggs settled 17 days after
settled'
spawning and 3 days later 20 larvae from potentially cross-fertilized eggs
dttr"., ,l"ttt"O on iubsequent days. When the experiments were terminaled 47
J"t; ;ft"r the initial spawning, most juvenile corals and a number of unmeta-
morphosed larvae were alive. No budding had occurred'
In the laboratory experiment to test how long larvae remained benthic, colonies
,p"*..a ""d egg'clumps attached to filamentous algae on5the bricks or settled
on tt" sand. Moit deveioping larvae remained benthic for days after spawning
(Fig. 8). On the morning of AuV 6, larvae were seen swimming and from then on
ihe"nu-ber of benthic l"uruu" decreased markedly. It is not known if the larvae
were lost at the outlet, died or settled'
Colony Size and ReProduction
This species is a protandrous hermaphrodite. Colonies smaller than 2.5 cm
were usually male (Fig. g), or if hermaphroditic, eggs were only sporadically
fresent inteimingled *ith ,p"t- clusters. As the size
of colonies increased' the
of with gonads increased (Table 4) as did the size of testes
i"i"l"t"g" co'ionies
and the number of eggs Per gonad.
Distribution, Abundance and Size
The transect data indicated that G. australensis is common and widely distrib-
uted on the reef flat of Heron Island (Table 5), where it is the fifth most abundant
,pl"i"r una co-pti ses 4.47o of the live coral cover. Similarly' Grassle (1973)
found G. australensis ranked sixth in abundance on the reef flat, but was not
common on either the crest or slope'
- -lirting"irhable
colonies ranged irom 0.5 cm to smallmicroatolls approximately
30;; lriatest radius. Coloniei measured in Transect 2 ranged from 0'5-13'9 cm
i with a mean size of 4'5 cm x (N : 45)' (Table 6)'
 KOJIS AND QUINN: SEXUAL REPRODUCTION IN GONIASTREA

Figure 9. plot of gonad abundance against the arithmetic mean radius of Goniastrea australensis:
(xl, colonies with testes only: (o) hermaphroditic colonies
Figure 10. Oceanic surface temperatures near Heron Island (averaged over 12 years, 1966-1977)
aniLordHowelsland(averagedover2years, lg77-1978)(CSIRO,Cronulla,Australia)andspawning
times for Goniastrett australensis (.L).

DrscusstoN
This is the first report of a study of sexual reproduction, larval development
and life history strategy of a hermatypic coral that releases gametes' Gamete
release may be widesfread among the Scleractinia. There have recently been
;;;;rt; oi gu-"t" release and external development of larvae in two species of
-andcoral from
ufr"i11utypi" temperate waters, Astrangia danae -(|zmant-Froelich et
al., 1980) Poracyathus siearnsii (Fadlallah and Pearse, MSt)' Gamete release'
simultaneous hermaphroditism and annual protogynous gonad development may
be common amongst the members of the Faviidae. Eggs were released in aquaria
colonies of Faviifauzs (Shlesinger in Rinkevich and Loya, 1979a). At Heron
irlulo, Kojis and euinn (1980) observed a brief annual spawning season in Fav-
ites a;dtti and Leptoria phrygict. Simultaneous hermaphroditism and
protogy-
,ro.rr, unnuul gonad devetopment have been rep'orted in Favia pallida aI Low Isles.
Greui su.rier- Reef (Mars^hall and Stephenson, 1933), Favia fragun (Duerden,
1902) and Faviafavus (Rinkevich and Loya, 1919a)'
Associated wiih simultaneous hermaphroditism is the possibility of self-fertil-
ization. G. australensis may commonly self-fertilize, since ova and sperm are
released simultaneously. Self-fertilization in sessile animals, such.as corals, ensures
ugui^i reproductive #asbge, since an "individual will always be certain of suc-
cissful contact with a reproductively capable individual, namely itself ' (Tomlin-
son, 1966), regardless of population density. Thus, it m.ay .be especially im-
;;.i";i in'ifre Jolonization bf ne* habitats and the recolonization of old habitats
uft", u catastrophe which eliminates all but a few widely spaced individuals'
A strong indi-cator of reproductive mode in hermatypic corals may be the num-
ber of eggi p"r gonad (Rinkevich and Loya, 1979a). Faviid species that release
eggs per gonad shortly before
!u-et"r?." stroigty fecund having more than seven per gonad. In contrast,
ipawning. G. auitralensis had a maximum of 13 eggs
itytophiro pistillata, a planulating species had only one or, rarely, two mature
eggs per gonad (Rinkevich and Loya, 1979a).

I Fadlallah, y., and J. s. pearse, MS. Sexual reproduction in solitary corals: synchronous gametogenesis and broadcast spawning
in Parac|dthus stearnsii r
 RIII-LETIN OF MARINE SCIENCE, VOL. ]1, NO. 3, I981

Table 4. Percentage of sexually reproducing colonies in similar size classes for Goniastrea austra'
/ensis and Stl,lophora pistillata (Rinkevich and Loya, 1979b) (i : arithmetic mean radius, Gm =
geometric mean radius)

G oniasttea aust rale nsi.t Sttlophoru pislil[0lu

Size Classes 7oof Colonies Size Classes 7" of Colonies
x (cm) Reproductive G- (cm) Reproductive

0.51-1 .00 19 0
1.01-1.50 4 0 1.01-1.50 33 3.0
I .5 1-2.00 l5 3 3.3 l .51-2.00 25 32.2
2.01-2.50 l0 40.0 2.01-2.50 22 54.2
2.5 1-3.00 l4 35.1
3.01-3.50 l5 60.0 2.5 1-4.00 13 69.2
3.5 1
-4.00 7 100.0
4.01-4.50 9 77.8 4.01-4.50 10 90.0
4.51-6.00 9 88.9 4.5 -6.00
1 Jb 86. I
6.01-20.00 18 I 00.0 6.01-8.50 l4 92-9

It has been suggested that the failure to detect planulation in corals may be
because (1) colonies of some species may not spawn every year (Connell, 1973),
(2) death may follow reproduction (Marshall and Stephenson, 1933) and (3) col-
onies studied were sexually immature (Stimson, 1978). Data from this study dis-
count all of the above. Colonies of G. australensis have been shown to spawn
annually, 3 years in succession, and thus colony death is not related to repro-
duction. Also, while planulating species, e.g. Srylophora pistiLlatd, become sex-
ually reproductive at an earlier age than G. australensis, the colony sizes of both
species are similar. The size at the onset of reproduction is small (t : 1.51-2.00
cm) and if this is true for other species, the possibility that all colonies inspected
were juveniles is unlikely. Indeed, failure to detect planulation may be because
(1) many corals do not planulate, they release gametes, and (2) gamete release is
brief, occurring annually.
Korringa (1947) listed three factors which determine the time of reproduction
in marine invertebrates: annual temperature variation, lunar tidal cycle or vari-
ations in nocturnal illumination, and the day-night (light/dark) cycle. The timing
of spawning in G. australensis appears to be influenced by all three factors.
Annual periodicity of spawning has been correlated with "a definite tempera-
ture, which is a physiological constant for the species, or . . a definite temper-
ature change, at either the maximum or minimum temperature of the locality"
(Orton, 1920). Gonad maturation in G. australensis may be influenced by the
rapid spring temperature rise beginning in September, with final ripening occur-
ring only when a minimum temperature of approximately 23*24C is reached (Fig.
l0). At Lord Howe Island the rapid spring temperature rise begins in November

Table 5. Relative abundance of Goniastrea australensis in transects l-5 on Heron Island reefflat

Transect No Vo I ive Coral Cover

I 4.1
2 4.9
3 3.1
4 8.2
5 3.7
l
KOJIS AND QUINN: SEXUAL REPRODUCTION IN GoNIASTREA

Table 6. Size distribution of Goniastreu rtustralensis on Heron Island reef flat along Transect
2 (i : arithmetic mean radius)

Size Classes

x (cm) % of Total N

0- l .00 2 4.4
1.01-2.00 6 I3.3
2.0 I
-3.00 9 20.0
3.0 I -4.00 9 20.0
4.0 1
-5.00 4 8.9
5.01-6.00 5 ll.1
6.01-10.00 5 lt.l
10.01-14.00 5 11.1

and 2f-24"c is not reached until January. This could explain why spawning at
Lord Howe Island is 2 months later than at Heron Island.
Timing of spawning is associated with the lunar cycle in a number of inverte-
brate specieS, €.g., the echinoid centrechinus (Diadema) setosus (Fox, 1924) and
the polychaete PLcttynereis dumerilii (Hauenshild, 1955, 1956). Timing of spawn-
ing is also correlated with the lunar cycle in a number of planulating coral species
(stimson, 1978). colonies of G. australensis spawned for 3 consecutive years
between the full and last quarter moon phases at the end of october or in No-
vember on the afternoon low tides. The lunar cycle appears to influence the
particular time of the month, while the day-night cycle may influence the time of
day or on which low tide in the semi-diel cycle spawning occurs (Table 7).
Time to settlement of larvae in hermatypic corals is probably influenced by
three factors: (1) presence of an appropriat-e^substrate, 121 suitable environmental
conditions (Harrigan, 1972; Chia and Bickell, 1977) and (3) the time from release
of planulae or gametes to the development of mature larvae (i.e., ready to settle)
(Rinkevich and Loya, 1979a). Settling and metamorphosis in many planulating
hermatypic corals occurred only when larvae were at an advanced developmental
stage, i.e., three to six complete mesenteries (e.g., Siderastrea radians, Isophvllia
dipsaceo, Duerden, 19021 Fungia octiniformis var. palawensis, Abe, 1937 po-
ciLlopora damicornis, Atoda, 1947a', and Acropora bruggemunni, Atoda, 1951a).
The larvae of Stvlophora pistilluta were released with or without mesenteries,
but settled only at the Halcampoides stage (Rinkevich and Loya, r979a). Some
larvae of S. pistillata were ready to settle immediately upon release, while set-
tlement in others was delayed until development was completed. Few of the
species studied had larvae able to settle at an early developmental stage, e.g.,
Galaxea aspera had no mesenteries (Atoda, 1951b) and Favia fragum had only
one pair of complete mesenteries before attachment (Duerden, 1902).
Although the stage of larval development at which settlement occurs is not
known in G. australensis, the relatively Iengthy period from larval release to
settlement as compared with many larvae releasing species may be attributed in
part to time to larval maturity. It may be that the minimum time to settlement of
17 days in aquaria is comparable with that occurring on the reef flat since Abe
(1937) found in Palau that Goniastrea aspera settled in aquaria between 16 and 23
days after release.
rn G. australensis gonads were present in most polyps irrespective of their
position in the colony. This contrasts with the distribution of gonads in branching
species. In Pocillopora damicorai.r (Harrigan, 1972) and Stvlophora pistillat6
l
BULLETIN OF MARINE SCIENCE' VOL'
31' NO' ]' 198I
570

species of hermatypic coral

Table 7. Reproductive periods for eight planulating

Lat
'.TG:
Annual Larval
Release Period
Author Location
Species
Hawatt All year
Cyphttstrea ocellina Stimson, 1978 All year
Caribbean 22"N
Favia fragum Duerden, 1902 20'N 8 months
Abe. 1937 Palau
Fungia actiniformis Palau 8'N All year
Gaktrea asPera Atoda, 195lb 8'N All year
Duerden, 1902 Caribbean
Manicina areoLatrt Palau 20"N All year
Pocilloporo da mic or nis Atoda,1947b
caespitosa
Marshall and StePhenson, 1933 Low Isles, Great 8'N All Year
illo portt mic ornis
P oc da
Barrier Reef 17'S
Harrigan, 1972 Hawaii 22"N All Year
P ocillopora damic orni s
Palau 8'N All Year
Seriatopora hYStix Atoda, 1951c 8'N All
Atoda,1947b Palau Year
Styktphora Pistillata
Rinkevich and LoYa, 1979b Eilat. Red Sea 29'N 8 months
S tt lophora Pis till ttt a

(RinkevichandLoya,lgTgb)gonadsaTemoreplentifulinthemid-sectionofthe
be absent tp'!9 1'5 cm from a
branch. while in Ar;;;;;;'pZlifera thev..mav mav be
!ll'l'lli;;;'i;';;;;;i Ju'""i"ii'inr' rni''oiir'ience in sonad distribution
localized growth in
of m-assive species vers;:
relared to the unlform growth jor,"s, 1g10; Buddemeier and 1l: Kinzie, 1976). For ex-
branching species (wo-oa- Porites lobata indicated that
ample, skeletal A"rrriiili"it rf tfrr"iassive (Isedale,
coral
there are no
srowth occurred ,"";;;;i;itne .1977)..Since
concentrated (cf. branching species), enersv
growth is"otutt,rm
;i-:JJi#;;#;ht"h is directed both to gonad devel-
in all polyps of reproiu;,i"il -ut"." colonies
;;;;;i";t growt'h' r .-r. :- -^-^^ in
Decreaseingrowthratehasbeencorrelatedwithincreaseinsizeinanumber
calcification in corals is most rapid in
of corals and it is g";;;lli;g*pr".g that Buddemeier and. Kinzie, 1976)' In
vounser, smalter, ;i;;;; iconn"il, 1973;
ie33;, stytophora pistittata (Rinkevich
'iiiii'r;,i#;..'i1il;;li-u,ro'st"pr'".'.or,,. .laigei colbni'es.were more fecund than
and Loya, tglgb) ""I'C-.^)"tti.trrris, rate is caused by the onset of
smaller colonies. rt -uv u" ttr"t decreasing irowthassociated with increasing size'
sexual reproductio" ffitd ft'il;-g
f;ffditv
oii"cts all its energy toward maintenance and growth until it
A colony initially o*i"g to competition and
attains a size *t"."-t-# ;h;;"t of de*ath ,orii" of_p.redation, itt energy to reproduction'
overgrowth diminishes'JnJli"u,'upportion of
As a colony loi.r"r"ur" in rir", li"an channel a larger proportion
"ontinrrlJ
energy to reProduction' faclors must
When deter.ining"it'te annual fecun<Jity of a coral ' a number of
(i.e., number of esgs per gonad)
be taken into considJ.;;ir;:'Fil, polyp fJcunoitv fecundity varies among pol-
may increase u, tt gro*, urg".. S""o"i,'if
f
""ioiony parts of a colony
yps of a colony, tt" rlii"'.r-i"proa.rJtiu"lo.ror-r"productive with sizelage'
whether thii ratio varies
will need to be detei.i""a as well as larvae for long periods (Table 7)' gamete
Third, while planuh;i;;;;;";: "bu'" plan-
releasing faviids t uue u-UJef annual
,pu.'r,in! period. individual colonies of
a reproductive season to deter-
ulating species t uu"-rit [""n ioUo*lO
tnroirgtr
turuu"";;;; ;;i;ased by a iotyp, ind thus pfesent knowledge
mine how -u'y in" nrr.nu"r oriuruu" found in a polyp at any one
time'
of fecundity l, in polyps of stylophora
"or.,nrrliTo of eggs, ,p"r- and larvae
The simultaneous presence
-anJ-ioyu,
pisti,ata Gint<evictr Tglg"l uii iorio fragum lDuerden, 1902) indi
I

KOJIS AND QUINN: SEXUAL REPRODUCTION IN GONIASTREA 571

cates that there may be more than one cycle of larval development and release
within a polyp in a year.
Stimson (19':.8) hypothesized that the release of planulae in shallow water corals
enables them to retain recruits in fast moving shallow water by allowing rapid
settlement. G. australensis is a shallow water reef coral that does not planulate
although it exhibits a mode of spawning and larval development that encourages
the retention of recruits on the reef flat. Spawning occurs before and during low
tide when the current is least and water depth shallow. During low tides, water
only just covers the coral heads ensuring that egg clumps will remain near the
substrate and not be carried by currents above the coral heads, a possibility
during high tide when the reef flat may be covered by 2 m of water. Egg clumps
are sticky and negatively buoyant, adhering to the substrate. Thus, developing
larvae remain near the parental colony. Ectodermal mucous gland development
enables the larvae to adhere readily to surfaces (Chia and Bickell, 1971). The
color of developing larvae provides effective camouflage in the benthic habitat
and possibly diminishes predation.
Stimson (1978) also suggested that corals characteristic of reef flats differ from
deeper water corals in simple demographic characters, such as age, at the onset
of reproduction. Stylophora pistillata, an r strategist (Loya, 1976a), characterizes
the reef flat in the Red Sea (Loya, 1972) and appears to be typical of the shallow
water corals discussed by Stimson. A comparison of size at first reproduction
between S. pistillata and G. australensis suggests that while linear size is similar,
age differs. S. pistillata first reproduces when less than 2 years of age (Rinkevich
and Loya, 1979b) while G. australensis, extrapolating from Woodhead's (1971)
growth data, does not begin reproducing until 4 to 7 years of age. With increasing
linear size, the differences in age between similar sized colonies of these two
species become more marked.
Species characteristic of reef flats that primarily form new colonies by means
of sexual reproduction may have various modes of reproduction that reduce
planktonic life to a minimum and retain larvae on the reef flat. Some species,
such as Stylophora pistillata (Loya, 1976a) and PocilLopora damicornis (Harri-
gan, 1972), planulate and are r strategists. They have a relatively small maximum
colony size, early age at the onset ofreproduction and possibly large reproductive
effort (year-round presence of gonads and/or larvae). Others, such as G. austra-
lensis, release gametes and are K strategists. They have a relatively large maxi-
mum colony size, delayed age at first reproduction and smaller reproductive effort
(i.e., gonads present only part ofthe year). Such species may be adapted to the
range of environmental conditions occurring on the reef flat over decades, if not
centuries. Edmondson (1928) tested a number of Hawaiian reef flat corals for
their ability to resist fluctuations in salinity, temperature and exposure and found
species of the family Faviidae to be among the most tolerant.
These data do not invalidate Stimson's hypothesis that different modes of re-
production are adaptive in different habitats. However, results show that plan-
ulation is not the only mechanism that can achieve this end and that demographic
characteristics of shallow water corals and thus their position on the r and K
continuum may vary.

AcrNowr-nDGMENTS
We thank Drs. R. Endean and C. Plowman of the University of Queensland, Brisbane, Australia;
Dr. D. C. Potts of the University of California, Santa Cruz, and Dr. R. Reichelt, Australian Institute
of Marine Science, Townsville, Australia, for their critical reading of the manuscript. We would also
like to thank Dr. R. Endean for his suggestions regarding experimental design and Mss. V. Harriot,
l

BULLETIN oF MARINE scIENcE'

vol- 3l' No 3' 1981
572

L.Keyes,l.Stejskalantlmanyothersfortheirassistanceinthefield'Thisstudywassupported
tr'6**tr"'a' in" Great Barrier Reef Marine Park
Au-
financially bv grants fit- ;il:"i;;;;t"t facilities
n"ton Island Research Station
s*t",y;l;;;i.Ju..-1t " r,r! olt1t" gratefully
thority and the Ecological Australia' is acknowledged'
and oceanographic data rt"t EdfnO' Cronulla'

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Dere Acclprpo: March 24, 1981.

Aoonesses: Zoologv Department, Universitt,of Queensland, Suint Lucia, Queensl.and 4067, Aus-
lralia;PnessNT ADDREss: (N.J.Q.) Fisheries Department, Papua New Guinea Universit,t of Tech-
nologl-, P.O. Bor 793, Lae, Papua New Guinea.