Proceedings of the Fourth Internationsl Coral Reef Symposium, Manila, 1981, Vol.

NEPRODUCTIVE STRATEGIES IN FOUR SPECIES OF

PORNDS (SCLERACTINIA}

Barbara L. Kojie
Tnlogy Department, University of Queensland, Saint Lucia, Queensland, Australis
and
Norman J. Quinn
Fisheries Departmeut, Papua New Guinea University of Teqhnologr, Lae, Papua, New Guinea

ABSTRACT

The mode and tining of reproduc'tion, abundance and distribution were studied in four congeueric species of
hern"matypic coral on Heron Island reef, Great Barrier \*f.. Poites and.rewsi, P. lobota, P. lutea and P. ,nunayen'
sis have stable gonochorism with the former three species having a short annual period ofegg and sperm release,
fertilization being external, and the latter'having internd fertilization and an ertended larval release period. P.
andrcwsi, P. lafutc and P. luiea reproduce asexually when pieces &om large colonies are broken off and dispersed
by waves and currents. The abundance of the four species ol Paites on Heron Islend reef flat is rel,ated to tbe
relative success of sexual and aserual recruitment (fragrnentation).

II\ARODUCTION the periphery. Colonies are shades of brown, green

Only a few studies have attempted to interpret
ecological data while considering the reproductive
ecology and life history phenomenon of hermatlpic
coral species (Loya 1976, Bak and Engel 1979,
Lewis 1974). Distribution, species diversity and
zonation were frequently studied aad related to
physical parameters such as wave action, water
depth (Yonge 1963, Stoddart 1969, Loya 19?2) and
sedimentation (Morrissey 1980). Part of the reason
for the lack of.attention to reproductive ecology was
the assumption that herrratypic corals were uni-
formly viviparous and released planulae (Hyman
f940). This assumption was questioned (Connell
1973, Stimsoa 1978) and shown to be incorrect
(SchlesingBn in Rinkevich and Loya 1979, Kojis aud
Quinn 1980, 198U.
&eproductive ecology, relative abundance and
maximum coloay size in four sympatric species of
Poites were studied on HeroD Island reef {23"2?'S},
Great Barrier Reef, Australia- Differences in abun-
dance and nerimrrm colony size are interpreted in
the light of variations in reproductive strategies
among the species.
SPECTES DESCRIPTIONS
P- andlewsi Dana, 1846, P lobata Dana, 1846, P.
lutea Milne Edwards and Haime, 1851 form large
microatolls >2 m in diameter on the reef flat. Small
and deepwater colonies of. P. andrewsi are ramose,
whereas microatolls, corrmon oa Heron Islaud reef
flat, have massive centers with short branches on
or grey. The massive colonies of P. lobota and P,
Iutea are purple and yellow-brown resp€ctively. In
contrast, P. nunuyensds Vaughan 1918 forms
small, massive or ensusting colonies with a max-
im 'm diameter of 20 Cm and is sirnila1' in color to P.
lutea Massive colonies of, P. murrayensds can be
easily mistaken in the field for small eolonies of P.
Iutes-
MATERIAI-S AND METHODS
Samples were colected at approrirnately moathly
intervals between September 1977 aad January
1979 from 30 taggpd colonies oI Poites andrewsi
(ten at each of three sites), ten tagged colonies ofP.
lobata (one site) and 20 tagged colonies of P. lutea
(two sites) (Fig. 1) as described in Kojis and Quinn
(1981). In addition, untagged colonies from various
regions of the reef flat were sampled periodically
during the year to assiess variation in the timing of
reproduction between habitats. A poriion of
colonie were sampled at intervals of one week or
Iess immediately prior to, during and after spawn-
ing. A rninimnm of six different colonies of P. mu*
rayensis were sampled randomly from inner reef flat
site A (FiS. 1) in Jaauary, March, May, September,
November and December 19?9.
Samples were fixed and decalcified as detailed in
Kojis and quiun (f98U. A 1-2 polyp thick slice of
decdcified tissue was plaeed on a slide and gently
squashed with a coverslip. With a dissectingor com-
pound microscope the presence or absence and sex
146 KOJISANDQUINN

of gonads, 2-Bpm diameter could be determined.
Representative samples were treated histologically
to verify observations of decalcified tissue with
regard to s€x and stages of gametogenesis, i.e. iime
of spermatozoan development and whether oocytes
developed into ova, zygotes or embryos in situ. It is
important to use a fixative and preservative that
does not harden tissue, e.g. formalin. Alcohol
hardened decalcified tissue making it rubbery and
non-amenable to squashing.
Shortly after spawning, the eggs of P. ondrewsi
were pipetted onto a slide with no coverslip and
measured. Since spawning was not observed in P.
lnbata and P. lutea, eggs were measured in
decalcified tissue collected close to the time of
spawning. The tissue was carefully dissected and a
coverslip placed over the tissue. Only larger eggs
from the latter two species were measured since
small eggs \il'ers lslqined after spawning. All eggs
were measured with an eyepiece micrometer on two
perpendicular axes and the average d.iameter deter-
mined.
Colonies were saurpled from the top, middle and
bottom on different sides of P. andrewsi (9 colonies;
3{ samples from each), P. lobata (1;?} and P. lutea
g; -12l.to determine if gonad development was syn-
chronized within colonies.
To observe spawning and larval development,
colonies of. P. andrewsi and P. lobata were held in
aquaria during the end of November and December
19?8 and 1979;P. luteainJauuary 1979 and,P. Mur-
rayensis between the end of November 1978 and
February 1979, mid-November and end of December
1979 and the end of October and beginning of
November 1980.
The method of determining distribution and abun-
dance of species from tratrsects (Fig. 1) is descriH
in Kojis and Quinn (1981). The size and mortality of
fourteen tagged colonies of P. nurrayensis were
recorded from 1978 to 1980.

EF CREST

l'hron lstand

transectsl and ?

transecl 3

transect tr

, 500m .

Figure 1. Map of sample and transect sites on Heron Island reef flat. Sample sites: J andS Poites lutea;2 P. laba,",,^ 4,5
and 6 P. andreusii A, general locale of P. muroyensis collections.
 REPNODUCTIYE STRATEGIES IN P,ORITES

RESULTS

T\ro modes of reproduction were observed in the

four speeies of Parttes studied. Annual broadcast
spawnng over approximately one week- was ob-
served only in P. iid.rewsi and is assumed to occur
in P. tobata and P. Iutea since samples revealed very
siniLar gonad development and spawning periodici-
ty. In contrast, P. murrayensis releases well-
developed larvae over five months or more.

BROADCAST SPAWNING IN P. AIVDI?EI'T1S/,

P.LOBATA ANDP. LUTEA

All thre sp,ies have stable gonochorism; entire

colonies are either male or female. In P. and.rewsi
the sex ratio between male and female colonies Figure ?. Several weeks prior to spawning, zoo8anthellae
sampled randomly was 2:1 (N:771. There was a low were abundant in the tissue surrounding oocytes
incidence of hermaphroditism: four colonies out of (Pontes labota): o, oocytes: a zooxanthellae.
IO7 (3.7Vol had an equal ratio of eggs to sperm
clusters and three colonies were predominately male
l2.9%ol with a few eggs in tissue samples' There was
t no hermaphroditism in P- lobata or P lutea All
t
tagged colonies were female and of 43 and 44 co
:.' lonies of P. Iabata and P. Iutearespectively sampled
I
:
frorn various regions of the Heron Island reef flat,
:
:,
only three in each species were male.
-* Gametogenesis of both sperm and eggs in allthree
species occurred over approximately four months of
the year. No gonads were present in samples from
tagged colonies of P. andrewsi and P. Iabata from
mid-January to the end of August 1978. Gonads
were observed in the majority of samples collected
the last week of September and spawning occurred
in December .ln P. lutea, a few oocyies, remnants of
spawning, were present in samples collected on 6
and ? February and no gonads were present in Figure 3. Prior to spawning, zoo'qanthell,ae e'ntered the
samples Som March to August. Only two of 13 ooplasm of oocybes (Porites lutea): o, oacyte; gu,
samples collected at the end of September had germinal vesicle: a zooxanthellae.
gonads, while all samples collected on 6 November
and examined had gonads. Spawning occurred diameter of 338 and 29? pm (N:20) respectively.
between the end of January and beginning of Spermiogenesis is similar to that describd for
February. Stylophora pistilla.ta (Rinkewich and Loya 19?9) and
In large reproductively mature colonies, ali Para,cyathus stearnsii tFadlallah 19811 with the ex'
mesenteries had gonads. In all three species, only ception that U all clusters in a colony are at the
ooc5rtes were present in histological sections of same stage of development at any one time {Figs. a
ovaries sampled before and afber spawning. Eggs of and 2) clusters containing sperln are present only a
P. andrewsi shortly after spawning, were white. few days prior to spawning. Sperm clusters in
Although the oocytes of. P. Iabuta andP. luteswere decalcified iissue were distinguishable from oocytes
observed ouly aftLr they had been fixed in formalin primarily by the absenee of a germinal vesiele. A
and decalcified, the color is probably similar. Zo+ testis initially developed in a mesentery as a row of
xanthellae, abundant in the endoderm surounding discrete, translucent clusters. These clrlsters en-
the oocytes {Fig. 2), were incorporated into the larged and coalesced prior to spawning.
ooplasm several weeks before release (Fig. 3). The Within a colony, gonad development, was closely
spawned eggs of P. andrewsi had an average synchronized and gonad abundances similar.
diameter of
231 am {N=19} and range of Among conspecific colonies gonad development was
ZI5-242 sm. Eggs dissected from decalcified tissue initially more variable, but proceeded to synchro-
of. P. tnibota and P. lutea had a maximun egg nized spawning.
l{8 KOJISANDQUI}{5

Figure 4. Sperm clusters within a colony were.at the same Figure 5, Just spawned.eCgs of Porites ondruwsii ;
&ag" of development {Poites andrcwsii); s' qperm above the sPawning colonY: e' eggs'
clusters: e, zooxanthellae.

In 19?8, no spawning was observed i:r aquarium
held colonies. A comparison of histological prepara-
tions of testes fron; colonies of P. attdrewsi held
se;e.al w€eks in aquaria and from freld colonies in-
dicated delayed development in the former.
In 19?9, spawning was observed in aquarium-held
colonies of p. onarewsi only. Dense clouds of sperm
were released on 1,2,4,5,7 aad 8 December between
ZttOO-Zt:gO hrs. On ? December 19?9 at 21:45 hrs a
f*g" i"*Uo of tiny white eggs were s-een floating
in an'aquarium {Fig. 5}. Eggs were also present
throughiut the water colurnn owing to coutinued
spawning and water movement. Fewer eggs were
released-during spawning on 8 December. Mobile,
ciliated larvae-developd approximately 23 h after
spawning. Tbenty tarvae were pipetted into a 2-1
taree mouth jar (Kojis and Quinn 1981) and four set-
In 19?8, P- lutea spawned betwren 20 Ja*-*:::
u"Jz F"t*uty (last qua*er moon, 1 FebruarS't;:..:;
i" igrs Gtwein'rs Jiluarv and 5 Februarv-(Fje'
with most spawning occurring before l! Jn::::s1:'
(last quarter-moon' 21 Jaurrary),
-
S"iiuffv mature colonies of all three -sp+;:r:r
;;;"""""4t. In tagged colonies sampleri-;-'
sic"""sirr" y"atj {P. ondrewsi: N= 13' -1977^' r: " :
i5ig n: 11, l9??, 19?8;P la-baca N :,3. !=l*
""a p.-l"iin tt:i6;1et8, le?e, 1er80 *-':
igia,-ieffit
lq-;a, 1-9y& 19?9) gonads were pre€ent ea€h 5*-'
Also,'since no colony changed sex during sanpi::"'
fi-;;;;" not a function of colony s'ze' a ccl*:".
pi"Utlfv does not change sex during iis lifetin:-
l\

tlel within ooe *eek of spawning- The remainder of

rii
! --
I
the larvae died or were lost inwater changes and the I

til/ iI
polyps survived onlY a few weeks. I
' Aiproximate dates of spawning wery t
I
I

iiit
{educed
trom-the pr€sence or absence of gonads in field c
t
samples and, in the case at P. andrewsl, aquarium
spawaing observations. It is suspe+ted that spawn-
s.
i:ng in these species, as in Goniastrea australensis
8.'
{Kojis aud Quintl 1981}, is influenced by the lunar
cvcie. In 1977, P. andrewsi spawned irnmediately llll [n
tFIr!
prior to 6 December (last quarter moou 4 December)'
io 19?8, spawning oicurred between l'l-23 Finure 6. Period of gonad development and spawnir-'
- -poirn"
December (last quarts moon' 23 December) tFig' 6) andrewsiiind P. lutea on Heron Island reei :--
and in 19?9 between 3-9'December {last quarter a"ti"" and summer 19?8-19?9' A'different *":':
moon, 1l December). l+;"November l9?8 to earlv Februarv lui:'
;;;? "oti"e
' Sampling for P. labata during the - summer .'..:

lg77-15 spawning season occurred only on 7 PTANULATION IN P. MURNAYENSIS

November 19?? when all colonies had gonads and 19
Jaauary 19?8 after all colonies had spawned. In P. murrayensis releasd larvae and was dioecj*::'*'
19?8, spawning occurred between 13-24 December rriift "
indication of hermaphroditiss'
a:C in 19?9 during mid'December.
"o
Gametogenesis probably began in August i:
BEPNODUCTIVE STRATECIES IN ISO&IrSS 149

September {all colonies collected 21 September had species in each transect with only P. and'rewsi and
gonads) and larval release began as early as mid- P. tnurrayensrs presant in all frve lrausects.
November. Reproductive colonies have either The patchy distribution o:f, P. lobata and P. lutea
sperm, eggs or eggs and larvae. The presence of was reidily apparent on the reef flat. Colonies ofP.
eggs and larvae simultatreously within a colony in- Iuteawete eoutmon ouly near Transects 1 and 2 (Fig.
dicated that larvae may develop and be released 1l where this species forms interconnected "step-
repeatedly over a single larval release s€ason. ping siones" from the mid-reef flat to the crest.
Larval release at Heron Island reef occurs from Microatolls af P. lobata were often clumped in wide
November to April, over approximately five months ly separate locations. On sandy substrate on tbe reef
of the year. flat SE and NE of Heron Island, they often formed
On 8 December 19?8, settled polyps and free mono-specific patches. Small, drscrete colouies of
swimming larvae were observed in aquaria in which these two species were found infrequently. In con-
colonies bad been placed several days earlier. Lar- trast, small colonies of P. and'rewsi and P- m.urro,yen-
vae continued to be released from colonies through' sis were cornmon on the reef flat' These species are
out December 19?8 to the beginning of February found in almost half the 15-m segments
19?9 when the field trip ended. Also, three out of six
field samples collected on 27 March 1979 had larvae; Table 2. The percent occlurence of four Pontes species in
while aone of 6 samples collected on 13 May had Tlansects 1-5 on Heron Islandreefflat. Tbansects were
gonads. Larvae were released on 20 November 1979 divided into 15-m segments. Figures refer io the percen-
tage of segments in which a species was recorded.
from eolonies placed in aquaria several days earlier
and continued to be released until the end of the
field trip in the last week of December. In 1980, col- Speies
onies placed in aquaria the first week of November
did not release lanrae, Thus, larval release at Heron P. andrewsi 4L.4
Island reef occurs from November to April over ap- F. mundyensis 40.5
proximately five months of the year. P. Iabato 7.6
The larvae of P. murmyensis were brown with P. lutea 6.0
abundant z,ooxanthellae ia the endoderm and had an
average dismeter of 6?9 rm (range 401'8?2 pm;
N:11). Released at an advanced developmental
stage, i.e. well developed stomodaeum" mesenteries Table 3. Relative abundaace lVo Eve coral cover) of four
and mesenterial filaments, they generally settle Poites speeies in Tlansects l'5 on Heron Island reef
rapidly, many within the first 24 hrs. flat.
the species apparen[ly rcproduces more than
once in its life time since colonies from 1.3-5.5 cm Tlansect No.
radius.had larvae. It is unknowB whether colonies Species
relea# larvae annually, since no colonies were
tagged and sampled over successive years.
P. andretusi 6.35 ?.36 9.51 6.22 6,82
DISTRIBUTION AND ABUNDANCE
Table 1 shows the relative abundance of the four
P. murrayensis
P. Iutea
1.25
1.20
2-84
2.88
2.05
0.58
I.27
'j'
P. lobeta 1.85 2-O3
Porites species in terms of perceat live coral cover
and Table 2 the number of 15-m segments in which
eaeh species occurs; the latter measurement gives
the relative number of colonies for each species. ASEXUAL COLONY REPRODUCTTQN BY
Table 3 shows the percent live coral cover of the FRAGMENTATION
Table l. The relative abuadance of four Pontes species on
Heron Island reef flat.
Recruitment by fragmentation may commonly oc-
cur among corals, especially those achieving a
Species Vo Live Coral Cover relatively large size. Connell (19?3) related obsenva-
tions of asexual recruitment in species of Acrcporu
andPoites, and Highsmith (19801 reported asexual
P- ondrewsi 6.02 colony "multiplication" in P. Iutea" On Heron
P. murtzyensis 1.?4 Islard, new colony formation by fragmentation
P. tutea 0.83 commonly occured in P. and.rewsr, less frequently
P. Iabata 0.78 in P. lobata and P. lutea arrd rarely, if ever, in P.
murrayensis.
150
Formation of fragments begins with bioerosion oI
eolonies at least partially separatingportious in situ
and is often completed by physical forceg sueh as
storms acting to separate and disperse fragments.
Fragmentation works most obviously on the
microatolle ol Poites.
Large microatolls of P. andrewsiwere at times in a
partiat state of collapse with fragments resting on
the sand around the collapsed portion of a colony.
Small branched fraguenti were sometimes totaliy
invested in tissue indicating frequent turning by
strong tidal currents or wa,re-actiott, and thus, c6nti-
nuing dispersal. When size prevented frequent turn-
ing or colonies lodged betwen oEhers, the basal
region became buried in sediment and supported the
rest of the colony above the substrate. Invariably
reef fl,at colonies were attached to substrate of. P. an-
drewsi origin indicating they had been derived from
asexually produced fragments or were in situ rem-
nants of onee much larger colonies.
Very large aud old microatolls of P. lpbata and P.
lateawere seldom single, discrete colonies. Death of
parts of microatolls separatd portions. Bioerosion
and solution acting on the dead tops lowered the
center in relation to the periphery and, eventually,
regrowth occurred in the center with subsequeat
separation (Fig. 7). Storms probably detach and
disperse portions of colonies.
Figure ?, A microatoll of Parites lutea on Heron Island
reef flat: n, regrowth into center of microatoll from live
periphery; N, new eolony. formed through separation of
regrowth.
Further substantiathg the common occurrence of
asexual reproduction in these three species is the
freguency with which a single color and sex
dominated local areas. Tbe ten tagged colonies of P.
:tJrewsi at site 5 were all male and bright green as
;*:e eigbt of the ten tagpd colouies at site 6. While
...-:;- i ir other localities (M. Pichon, pers. comm.),
KOJISANDQUI]{X
colonies on Heron reef flat were all of a single coiar
and most were female.
DISCUSSION
Withiu the family Poritidae and the genusPorires
two modes of reproduction have been observe*
Parites murrayeisis, P. stephensoni (Crpssland
1952) {: P hpddani, Marshall and Stephenson
1933), P. asrewides, P. cl.auada {Vaughan 1d10} and :::
Goniopora queensland.iae decimd Eernard l{Yama-
zato, pers. comm.l released Larvae, whileP. ondrewsi
i*
and by inference P. lobata and P. /rztea, spawued ::;
eggs and sperm, Gonochorism is the rrrle in the fami-
ly Poritidae as Goniopora qu,eenslandioe deeime
(Yamazato et al. 1975ll, P. lobata, P. compressa
(Stimson 19?6) and the species in this study are
dioecious. ::1
The relative abundance of the four species of *S
Poites studied on Heron Island is related to the :]$
relative success of sexual and asexual reandtment
(fragmentation). P. andrewsi" the most abundant
$
species in terms of percent coral cover and frequen' ..n
cy of occurence in 15-m segments, reproduces sex- .i*
ually by seaeonal broadcast spawniug, once'a year.
The two least abundant species, P loban and P.
larea, probably spawn sirnilarly. Successful local
sexual recruitment in all three species appear€ to be
infrequent. This may be owing in part to extremely
high mortality of eggs, larvae and juveniles; a conse
quence of their small size (Chia f97a). Sexual
recruitment in P. Iobata and, P, lutea may be further
hampered by unsuccessful fertilization of ova owing
to the faucity of male colonies. This assumes that
development is not parthenogenic.
In all three species, fagnentation is probably the
primary mode of recruitment on the reef flat.
Fragmentation by mechanical forces rccurred more
frequently and fragments probably survived more
readily in P. andrewsi owing to its branching habit :t:
than in P. labata and P. /a*ea which form massive
colonies. SmaII massive colonies are more easily
buried in sediment than branching colonies owing to
inherent differences in shape and patterns of
o3$r"*
and recruitment among planula releas-
ing corals, such as P. murrayensiq are primarily by
sexual means. Asexual reproduction plays little or
no role. The percent live eoral cover of P. rnuralen-
srs was considerably less thanP. andrewsi, but more
than P. Iobato and P lutea Recruitment in P. an'
d.rewsi and.P. rnunayeasrs was equally successful as
both occur in a similar number of 15'm segpents,
but was achieved in P. andrewsi largely by asexual
new colony formation and in P. murrayensis by sex'
ual recruitment- Sexual recruitment was more suc'
cessful in the latter owing to 1) large larval size
REPBODUCITIVE STRATEGIES tN roiIT&S 151

$79 pm versus 241 pp,l and 2) the extended larval Connell, J.H. f973. Population ecology of building
release psriod. Larvae of P. murrayensrs were corals. /n
O.A. Joaes and R. Eodean Biology
available for settlement for at least 5 months of the and Geology of Coral Reefs. VoL 2, 2A5"248.
New York: Academic Press.
year comparad with a few weeks far P. andrewsi
Fadlallah, Y. 1981. Tbe repreduetive biology of three
The low percent coral cover of P. murzuyensds com' species of corals from Centrd PN.D.
pared with P. a.nd.rewsi was due to the relatively Thesis, University of California at Santa r9s
small colony siee of P. munayeasrs rather than a p.
low numbs of colcniee. Highsmith, R 1980. Passive colonizetion end ualcol-
Species which achieve a small adult size relative ony multiplication in the nrsssive Poites
to others in the same taxa frequently produce large luteo, J. Exp. Mar. BioL EcoL 45(1h5$6?
eggs or larvae and develo,pment of eggs is lecitha' Hyman, L,N. 1940. lbe invertebrates. l.
trophic, bd protectine or viviparous {Chia 197a). through Ctenophora, New York 726
p.
P. murrayensr} releases relatively large planulae Kojis, B.L. and N.J. Quinn. 1980. Mode and Lg of agx-
and achieves a nraxirnum diamets of 20 cm, while ual "reproduction in some members of the her-
the other three species release small eggs and col- matypic coral family Faviidae. Am.Znl. 819.
onies achieve a size of >2 m. 1981, Aspects of sexual repro snd
Maximum size achieved by corals may be related larval dwelopment in the sballowwater
to two factor$ l) the amount of energy expended oa coral, Goniastrea australensis
growth and reproduction as indicated by the Haime 185?). Bull. Mar. Sci. {in prees).
iumber of montis per annum gonads and/oriarvae L€wis. J.B. 1974. Settlement and growth factr influone-
are present and 2) the longevity of colonies of a ing the contagious distribution of some Atlantic
reef cords. Prrc. 2nd Ini. Coral Reef
species. Both P. munvyensis and P stephensoni
2t
201-205-
{Stephenson 1933} release planulae over 4n extend- Loya, Y. 1972. Community structureatrd species
ed p€riod and are small in size. AlBo, the maximum of hermatypic coralg at Eilat, B€d Sea.
age-of P rnurrayensis may be relatively young. The 13:1fi}'123.
largest of 14 tag8Bd colories ( > 10 cm diameterl did 19?6. Settlement, mortality recruit-
in situ, whereas the smaller colonies survived. In ment of a Red Sea coral population. .lz G. . Mackie
coatrast, there was no morbality among the larp {ed.). Coelenterate Ecology and Behavior. 8F99.
taCged colonies af P. andrewsi P. lpbato and P. New York Plenum Publishing Corp.
lztea These species pmbably achieve a larger size Marsball, S.M. and T.A. Stephenson. 1933- The
than P. murrayemsis and P. stephensoni because of the corals. Sci. Bep. Great Barrier
1928-1929. 3:2L9-245.
they grow faster and are longer livd. Morrissey, J. 198t). Community struciure and nof
nacroalgae and hermatypiccorals on a reef
flat of Magaetic Island (queenslard,
Aquatie Botaay 8:91-139.
ACKNOWLEDGAMENTS Rinkevich, R. and Y. Loya. 1979. The of the
Red Sea Coral, StybpftomPistilhta l. and
firis r€search was supported by grants from the planulae. Mar. EcoL Prog. Ser. 1:133-144.
University of Quee.n"leld, tbe Great Barier Reef Marine Stephenson, TJq,. 1939. Developmenl and fon of
Park Authority and the Ecological Society of Australia- colonies of Po cillopotz and Porfees, Part l.
Rep.
We *ould Iike to thenk Ms. L. Daddow for assisting with Great Barrier R€ef Exped. 1928-f929, 3:1 3-134.
histological preparatious, Prof. Ir{. Pichon for iudentifying Stimson, J.S. 1976. Rsproduclioa in
some coEmor
specimens of Porites and Mes. V. Harriott, L. Keyes, I. Hapaiian reef corals. /n
G.O- (ed.l.
Stejskal and many others for their assistance fu the fieH- Coelenterate Ecology and tsehavior. pp. 2?1-180.
The uee of lleron Islaud Research Station is Sratefully New York: Plenum Press.
ackaowledged . 19?8. Mode and timing cf rel bion in
some common bermatypic corals of ii and
Enewetak. Mar. BioI. 48:1?3-184.
Stoddsrt, D.B" 1969. Ecology and morpbologr recent
REFEBENCES coral reefs- Biol. Rev. 44:438-498.
Vaughan, T.W. f910. lbe recent na of
Bak, RP.M. aqd Epgel M.S 19?9. Distribution, abun southern Florida. Yb. Carnegie Wash.
dance and survival of juvenile hermatypic corals 9:135-144.
(seleracbiuia) and the ot i!6 history Yamazato, K., M. Oshiro and E. Oshiro. 192S.
strategy in tbe parent coral community. Mar, Biol. tive biology of a scleracbinisn coral
54(4):341-352. qtaensbndioe decima^ 13th Pacific
Chia, F.S. 19?4. Clagsificatiou and adaptive sis.ifcance greee Proc. 1:135,
of developnental patt€rns in marin* inve'rtebrates. Yonge, C.M. 1963. The biology of coral
Thalassi:a Jugoslavica lO: 121.13O. Mar. Biol. l:20S260.