MARINE ECOLOGY PROGRESS SERIES
Mar. Ecol. Prog. Ser.
Vol. 60: 185-203, 1990
Published February 8
REVIEW
Reproduction and recruitment of corals:
comparisons among the Caribbean, the Tropical
Pacific, and the Red Sea*
Robert H. Richmond, Cynthia L. Hunter
Marine Laboratory, University of Guarn, UOG Station, Mangilao, Guam 96923, USA
ABSTRACT. Detailed reproductive data are now available for 210 of the ca 600 identified scleractinian
reef coral species. The majonty (131 species) are hermaphroditic broadcast spawners, although
hermaphroditic brooders (11 species), gonochoristic broadcasters (37 species), and gonochoristic brooders (7 species) have also been reported. Characteristics of sexuality and mode of reproduction are
generally conservative within species, genera, and even families, although some exceptions occur.
Variation in timing or mode of reproduction in allopatric populations may represent adaptations to local
environmental conditions or indicate problems in the taxonomy of some groups. Synchronous spawning
of numerous species occurs on the Great Barrier Reef, while asynchrony among and withln species has
been observed in the Red Sea, Caribbean, Central Pacific, Hawall, and southern Japan. Sexual
reproduct~onis the primary means for successful recruitment for some coral populations, while asexual
processes may be the dominant or sole means of recruitment for these same species at the limits of their
ranges. Recruitment success of different reproductive strategies may vary within and between localities,
and is mediated by both biotic (predation, competition) and abiotic (environmental variability, disturbance) factors. Data on reproductive patterns and recruitment success may be applied to coral reef
management practices.
INTRODUCTION
Until the last decade, the majonty of data on coral
reproduction were anecdotal and incomplete observations based on short-term and sporadic studies (see
review by Fadlallah 1983). This situation is not surprising, in light of the remoteness of tropical coral reefs
from most universities and research facilities, and the
logistical difficulties of studying corals in situ. However, a number of recent investigations have been
published based on continuous monitoring of field
populations, as well as histological and laboratory
examination of individuals. Previous generalizations
This review originates from a UNESCO/COMAR workshop
held in Fiji comparing Atlantic and Pacific tropical coastal
ecosystems
8 Inter-Research/Printed in F. R. Germany
and perceived trends may now b e re-examined, as data
for a greater number of species over a wide geographic
range have become available.
Detailed reproductive data have been reported for ca
4O0/0 of the known species from the tropical Pacific
(studies from the Great Barrier Reef, Guam, Palau,
Enewetak, Hawaii, Okinawa, a n d Panama), 3 0 % of
Caribbean coral species, and 6 % of Red Sea species.
These studies provide information on coral sex (hermaphroditism vs gonochorism), mode of reproduction
(brooding vs broadcast spawning), and timing of reproduction (seasonality, periodicity, a n d synchrony). Certain patterns of reproduction and recruitment are now
discernible from these data. In this paper, w e review
data for 210 scleractinian species, a n d compare reproductive processes observed in the Caribbean, eastern
Pacific, Hawaii, Central Pacific, southern Japan, Great
Barrier Reef, and Red Sea. As data for more taxa and
186
Mar. Ecol. Prog. Ser 60: 185-203, 1990
regions become available, additional (or different) patterns and trends may emerge.
The study of coral reproduction has advanced
through numerous theses and dissertations over the
last 10 yr, as well as from concerted group efforts, most
notably on the Great Barrier Reef of Australia. The
major pattern that has developed from the Great Barrier Reef studies is one of remarkable similarity and
synchrony of reproductive activity among coral species.
The majority (90 %) of species studied there broadcast
spawn gametes annually, during the week following
the full moon in the austral spring (Harrison et al. 1984,
Willis et al. 1985, Babcock et al. 1986). Data from other
regions (the Caribbean, Red Sea, Central Pacific,
Hawaii, and southern Japan) show different patterns,
with considerable variation in mode, timing, and
synchrony among species. In addition, populations of
the same species reported from 2 or more regions may
display different reproductive traits. Globally, corals
display great plasticity in their life history characteristics. These data are summarized for each region in
Table 1, and across regions in Tables 2, 3 and 4.
REPRODUCTION IN GENERAL
Corals reproduce both sexually and asexually. Sexual reproduction involves the process of gametogenesis, which may require from a few weeks for
sperm, to over 10 mo for eggs. Spawning and subsequent fertilization of eggs by sperm results in small,
presumably genetically unique, dispersive propagules
(planula larvae) which may settle, metamorphose and
develop into primary polyps. Asexual reproduction is
also common in many scleractinian species, and may
occur through fragmentation (see review by Highsmith
1982), polyp bail-out (Goreau & Goreau 1959, Sammarco 1982), or asexual production of planulae (Stoddart 1983). Asexual processes result in clonal propagules (genetic replicates of adult colonies) which, if
derived from fragments, have the apparent advantages
of large size and locally adapted genotypes.
Stylophora pistillata; Loya 1976, Rinkevich & Loya
1979a).
Corals can be simultaneous or sequential hermaphrohtes (see discussion in Fadlallah 1983). Two
species, Stylophora pistillata and Goniastrea favulus,
exhibit protandrous development (Rinkevich & Loya
1979b, Kojis & Quinn 1981a).The only report (Duerden
1902) of true protogyny, in which the colony functions
first as a female and becomes hermaphroditic in subsequent years, has been questioned (Szmant 1986). Most
hermaphroditic corals exhibit annual protogyny, where
eggs develop prior to spermary formation during a
reproductive season.
Mixed breeding systems have been described for a
brooding species, Pontes astreoides, in which 26 % of
colonies examined were hermaphroditic, 28 O/O had only
female gonads, and 46% were sterile (Chornesky &
Peters 1987). In Galaxea fascicularis, some colonies are
female and some are hermaphroditic, but eggs in the
latter apparently serve only to provide buoyancy for the
sperm packets (Heyward et al. 1987, Hamson 1989).
Hermaphroditism is advantageous when the probability of finding (or, in the case of sedentary corals,
proximity to) members of the opposite sex is low, and
self-fertilization is possible. Heyward & Babcock (i986)
found varying levels of success in self-crosses in 4 coral
species (0% in Montipora digitata, 1.5 to 1 6 % in
Acropora tenuis and Goniastrea aspera, and 26 to 89 %
in Goniastrea favulus). In experiments performed during spawning events on Guam during the summers of
1987 and 1988, no viable planulae developed from
mixing gametes from the same individuals in Acropora
irregularis or A. humilis, while nearly 100 % of the
embryos resulting from self-fertilized eggs of Acropora
tenuis developed successfully (Richmond 1989,
unpubl.). Barriers to self-fertilization apparently break
down with time after spawning for some species, but
not for others (Heyward & Babcock 1986, Richmond
unpubl.)
Numbers of gonochoristic and hermaphroditic
species within each region are summarized in Table 2.
Within regions, hermaphrodites range from 60 to 100 %
of reported species. Globally, the majority (68 %) of
coral species studied are hermaphroditic (Table 3).
Sexual reproduction in corals
Hermaphroditism vs gonochorism
Brooding vs spawning
In hermaphroditic species, ovaries and spermaries
may develop on the same mesentery (most favids and
mussids), on different mesenteries within the same
polyp (most pocilloporids and acroporids; see Fig. l ) ,in
different polyps within the same colony (e.g. Cladopsammia rolandi; de Lacaze-Duthiers 1897 in Fadlallah
1983),or at different times within the same colony (e. g.
Fertilization may take place within the maternal
polyp (brooding), or externally in the water column
after gametes are shed (broadcast spawning). Species
which broadcast spawn outnumber brooders in the
Pacific regions and the Red Sea (Table 2 ) . However,
brooding may be the predominant mode of reproduction in the Caribbean. Overall, for the present data
h c h m o n d & Hunter: Reproduchon a n d recruitment of corals
187
T a b l e 1. Reproduchve charactenstics of corals from t h e Caribbean S e a , Great Barner Reef, Central Pacific, Hawaii, O i u n a w a ,
eastern Pacific, a n d Red S e a . Symbols - Sex: H, h e r m a p h r o d l t ~ c ;G , gonochoric, X , u n k n o w n . M o d e : S , s p a w n e r , B, brooder;
possibly stenle. Timing: month a n d lunar d a y of g a m e t e release (spawners) or planulation (brooders) [ m o n t h is divided into
p h a s e s 1 , n e w moon, 3, first quarter, 5 , full moon, 7 , last quarter; 2, 4 , 6 a n d 8 indicate intermediate lunar p h a s e s (after Shlesinger
& Loya 1985));W , winter, s p , s p n n g ; sr, s u m m e r ; f , fall; yr, year-round; X, u n k n o w n
Species
Sex
Mode
Timing
Source
Caribbean
ACROPORIDAE
Acropora cenricorn~s
Acropora palmata
H
H
S
S
JuVAug, 6-7
Aug
Szmant-Froelich (19841, Szmant (1986)
Szmant-Froelich (1984). Szmant (1986)
H
SP
SP
sr
Yr
Duerden (1902), Van Moorsel (1983)
Vaughan (1910)
Mavor (1915)
Van Moorsel (1983)
AGARICIDAE
Agaricla agancltes
Agaricia crassa
Agaricja fragills
Agarjcia humihs
FAVlIDAE
Diploria stngosa
Favia fragum
H
B
B
B
B
H
H
S
B
Aug, 7
Yr
yr, 3-5
~Manicinaareolata
H
B
~Montastreaa n n d a n s
Mon tastrea ca vernosa
H
G
S
S
SP
Yr
Aug, 7/Sep, 7
A"g
Szrnant-Froelich (1984)
Duerden (1902),Vaughan (1910)
Szmant-Froelich (1984), Szmant (1986)
Szmant-Froelich et al. (1985)
Wilson (1888) ~n Fadlallah (1983)
Duerden (1902)
Szmant-Froehch (1984),Szmant (1986)
Szmant-Froelich (1984). Szmant (1986)
MEANDRINIDAE
Meandnna ( = ~Meandra)areolata
X
B
JulIAug, 3-8
Boschma (1929),Yonge (1935) in Fadlallah (1983)
G?
H
B
B
SP
Feb-Mar
Duerden (1902)
Szmant-Froelich (1984). Szmant (1986)
H
B
May-Jun
Jan-Sep
yr, 6 8
Nov-Feb
Vaughan (1910),Szmant-Froelich (1984)
Szmant (1986)
Chomesky & Peters (1987)
Tomascik & Sander (1987)
MUSSIDAE
Isophyllia sp.
Mycetophyllia ferox
PORITIDAE
Pontes astreo~des
Porites pontes
SIDERASTREIDAE
S~derastrearadians
X
X
(or female only)
B
G7
(some hermaphroditic)
H
G
G
B
B
S
F
yr?
Jul-Sep
Duerden (1902)
Szmant-Froelich (1984), Szmant (1986)
Szmant-Froelich (1984),Szmant (1986)
G
6G 10H:3x
S
12B 7 s
Aug
Szmant-Froelich (19841, Szmant (1986)
Pacific
Great Barrier Reef
ACROPORIDAE
Acropora aculeus
Acropora aspera
Acropora austera
Acropora cereaLis
Acropora cuneata
Acropora cytherea
Acropora digitifera
H
H
H
H
H
H
H
S
S
S
S
B
S
S
Acropora divancata
Acropora elseyl
H
H
S
S
Acropora flonda
H
S
Nov, 6
seasonal
Nov, 6
Nov, 6
sp-sr
Oct/Nov, 6
sp-sr
Oct, 6
Oct, 6
Nov, 6/Dec, l
Oct, l/Nov. 6
Nov, 6
Acropora formosa
H
S
Acropora gemmifera
Acropora grandis
H
H
S
S
Babcock et a1 (1986)
Bothwell (1981)
Babcock et al.(1986)
Babcock et al.(1986)
Bothwell (1981)
W~llise t al. (1985), Babcock et al. (1986)
Bothwell (1981)
Wlllis et al. (1985)
W ~ h et
s al.(1985)
Babcock et al. (1986)
W i h s et al. (1985)
Willis et al. (1985),Wallace (1985b),
Babcock et al. (1986)
Babcock et al. (1986)
W d l ~ set a1 (1985)
Wlllis et al. (1985), Babcock et al. (1986)
W i h s et a1 (1985). Babcock et al. (1986)
S~derastreasiderea
TROCHOSMILIIDAE
Dendrogyra cylindrus
Nov. 5-6
Oct, 6/Nov, 6
Nov, 6
Nov, 6
8
M a r . Ecol. Prog. Ser 60: 185-203, 1990
T a b l e l (continued)
pecies
Sex
acific
reat Barrier Reef
CROPORIDAE
cropora granulosa
cropora horrida
cropora humlLis
cropora hyacjnlhus
cropora laOstella
cropora longlcyathus
cropora loripes
cropora l u t k e n ~
cropora rnlcropthalrna
cropora rnlllepora
cropora nasuta
cropora nobdis
cropora pahfera
cropora pulchra
cropora robusta
cropora sarnoensls
cropora sarmen tosa
cropora
cropora
cropora
cropora
secale
selago
sol~tanensls
tenuis
cropora v a l e n a e n n e s ~
cropora valida
cropora cf varia bills
cropora cf vaughanl
cropora yongei
streopora rnlcrophthalma
ontipora aequ~tuberculata
ontlpora dlgltata
on lipora fol~osa
onlipora hisplda
ontipora lnforrnis
ontlpora rnonaslenata
ontlpora spurnosa
ontlpora tu berculosa
ont~poraturgescens
GARICIIDAE
chysens rugosa
Mode
Timing
Feb/Mar
sr?
sp-sr
Oct, 6
Nov. 5-7
sp-sr
Oct/Nov, 6
Nov, 5-6
Sep. 6/0ct, 6
Nov, 1
Nov. 5
Source
sp-sr
Oct, 6-7
Nov. 5
sp-sr
Nov, 6
Nov, 6
Nov, 6
Feb/Aug/Nov?
Nov, 7
Nov. 6 7
Nov. 6
Oct, 6
Oct, 6-7/Nov, 6
Nov, 5-6
Nov, 6
Oct, 6-7/Nov, 6
Nov, 5-6
sp-sr
Nov, 6
Nov, 6
Nov, 6
Oct, 6
Oct, 5/Nov. 5
Nov. 5
Nov. 5
Oct/Nov, 6
Nov. 6
Oct, 5
Nov, i
Nov, 5
Nov, 6
Nov, 6
Wallace (1985b)
Wallace (1985b)
Bothwell (1981)
\M~ll~s
et a1 (1985)
Babcock et al. (1986)
Bothwell (1981)
Willis et al. (19851, Wallace (1985b)
Babcock et al. (1986)
Wilhs et a1 (1985)
Babcock et al. (1986)
Willis et a1 (19851, Wallace (1985b)
Babcock et al. (1986)
Wallace (1985b)
Babcock et a1 (1986)
Babcock at a1 (1986)
Willis et a1 (1985)
Babcock et a1 (1986)
Bothwell (1981)
CVillis et al. (1985)
Babcock et a1 (1986)
Willls et a1.(1985), Babcock et a1 (1986)
Willis et al. (1985)
Wallace (1985b)
Babcock et a1 (1986)
Bothrvell (1981)
Bothwell (1981)
W~lliset al. (1985)
Babcock et al. (1986)
Bothwell (1981)
Babcock et a1 (1986)
Willis et al. (1985)
Willis et al. (1985)
Wallace (1985b)
Babcock et a1 (1986)
Babcock et a1 (1986)
Will~set al. (1985), Babcock et a1 (1986)
Willis et al. (1985)
Willis et al. (1985)
Babcock et a1 (1986)
Willis et a1 (1985)
'I,V~lliset a1 (1985). Wallace (1985b)
Babcock et a1 (1986)
Bothwell (1981)
Willis e t a1 (1985)
Babcock et a1 (1986)
Babcock et a1 (1986)
Will~set al. (1985)
Willis et al. (1985)
Babcock et al. (19861, Heyward & Collins (1985)
Babcock et al. (1986)
Willls et a1 (1985)
Babcock et al. (1986)
Willis et a1 (1985)
Babcock et al. (1986)
Babcock et a1 (1986)
Babcock et a1 (1986)
Babcock et al. (1986)
Nov, 5-6
Nov, 6
GV~lllset a1 (1985)
Babcock et a1 (1986)
Nov/Dec
Nov. 6
Nov, 6
Oct, 6
Nov, 5
sp-sr
Oct, 6/Nov, 5-6
Nov. 5-7
Nov, 6 7
Oct, 6
Oct/Nov
Nov. 5-6
X
R~chmond& Hunter. Reproduction and recruitment of corals
Table 1 ( c o n t ~ n u e d )
Species
Sex
Pacific
Great Barrier Reef
AGARICIIDAE
Pachyseris speciosa
Mode
T~ming
Source
S
Oct. 67/Nov. 5-6
Nov. C 7
Willis et al. (1985)
Babcock et al. (1986)
Marshall & Stephenson (1933)
Pavona cactus
CARYOPHYLLIDAE
CataIaphyUia jardinen
Euphyllia ancora
Euphyllia div~sa
S(sperm)
X
S
S
S
Physogyra lichtenstelni
S
Oct, 6
Oct. 6
Oct. 6
Nov, 6
Nov, 6
Willis et al. (1985)
Willis et al. (1985)
Willis et al. (1985)
Babcock et al. (1986)
Willis et al. (1985)
Babcock et al. (1986)
B
S
S
B
S
S
Dec. I
Jan-Jun
Apr-Jun
Nov. 6 7
Oct, 6
Nov. 7
Babcock et al. (1986)
Fisk (1981) in Hamott (1983a)
Fisk (1981) in Harriott (1983a)
Babcock et al. (1986)
Willis et al. (1985)
Willis et al. (1985)
Nov. 6
Nov. 5-7
Nov. 5
Oct, 6
Nov. 6
Oct, 6
Nov, 6
Oct, 6
Dec
OcWNov, 6
Nov. 6
Nov. 6
Nov. 6
Nov, 5-7
Nov. 6
Nov. 5-6
Nov, 5-7
Nov, 6
Nov, 6
Nov, 6
Nov, 6
Oct. 6
Nov. 6
Nov. 6
Oct. 5-7
Oct. 6-?/Nov, 6
Nov. 5-6
Nov. 6
Oct/Nov. 5-6
Willis et al. (1985), Babock et al. (1986)
Babcock et al. (1986)
Willis et al. (1985). Babcock et al. (1986)
Willis et al. (1985)
Babcock et al. (1986)
Willis et al, (1985)
Babcock et al. (1986)
Willis et al. (1985)
Harrison (1985)
Babcock et al. (1986)
Babcock et al. (1986)
Willis et al. (1985)
Babcock et al. (1986)
Harriott (1983a)
Babcock et al. (1986)
Babcock et al. (1986)
Willis et al. (1985)
Babcock et al. (1986)
Marshall & Stephenson (1933)
Willis et al. (1985)
Babcock et al. (1986)
Willis et al. (1985)
Babcock et al. (1986)
Babcock et al. (1986)
Kojis & (luinn (1982). Babcock et al. (1986)
Willis et al. (1985)
Babcock et al. (1986)
Willis et al. (1985)
Willis et al. (19851, Babcock et al. (1986)
Willis et al. (19851, Babcock et al. (1986)
Willis et al. (1985). Babcock et al. (1986)
Willis et al. (1985)
Babcock et al. (1986)
Babcock et al. (1986)
Babcock (1984)
Willis et al. (1985)
Babcock et al. (1986)
Babcock et al. (1986)
Kojis & Quinn (1981a). Kojis & Quinn (1982)
Oct, 6/Nov, 5-6
Nov, 5-6
Willis et al. (1985)
Babcock et al. (1986)
DENDROPHYLLIDAE
Dendrophyllja sp.
Heteropsammia cochlea
Heteropsarnrnia aequicostatus
Tubastrea faulkneri
Turbinaria frondens
Turbinaria reniformis
FAVIIDAE
Australogyra zelli
Barbattoia amicorurn
Caulastrea furcata
Cyphastrea chalcidium
Cyphastrea micropthalrna
Cyphastrea
Diploastrea
Echinopora
Echinopora
Ecliinopora
seraiha
heliopora
gemmacea
hornda
lamellosa
Favia favus
Favia lizardensis
Favia mathaii
Favia pallida (as F. doreyensis)
Favia pallida
Favia rotumana
Favia stelligera
Fa vla veronl
Favites abdita
Favites bennettae
Favites chinensls
Favites complanata
Favites nexuosa
Favites halicora
Favites pentagona
Fa vites russelli
Goniastrea aspera
Goniastrea edwardsi
Goruastrea favulus
(as G. australensis)
Goniastrea favulus
X
Nov, 6
Nov. 6
Oct, 6/Nov, 5
Nov. 6
Nov/Dec
Nov. 6
Nov. 6
Nov. 5-6
189
J T T
m
(Am
WV]
m
m
J
T T
T Z T
0000
00
X
X X T
3:T
T
J J T
J
T T
3:
V]
- - ( U - m - - - - 3 - m n r u r
r
-
r
w w - w
-
r
r
r
+
3
r
-
r
y
r
w w w w
w
w
w
mmzEzggKgzgzgzg
m m
- . m
m m 2
U
ru
n n
0 0
n o
(U
m
9
-
-
ru
n
0
n
(U
n
0
n
-
ru
n
0
n
X
22%
'D
U U U
h X
X
'D 'D
r" C"
m -
m
r n r n r n " ~ 9
U U
X
X
0
n
X
.E E
. E
.
---r".
w w w
w
-
c
+
m m m
m m m
U - -
L
m
m
m
9
hchrnond & Hunter: Reproduction a n d recruitment of corals
191
Table l (continued)
Species
Sex
Mode
Tirnlng
Source
Pacific
Great Barrier Reef
PECTINIDAE
Pectinia lactuca
H
S
Pectinia paeonia
H
S
Oct, 6
Nov, 6
Oct. 6-7/Nov, 6
Nov, 6
Willis et al. (1985)
Babcock et al. (1986)
Willis et al. (1985)
Babcock et al. (1986)
yr. 1-5
Marshall & Stephenson (1933)
-.
POClLLOPORlDAE
Pocillopora damicornis
(as P. bulbosa)
Pocillopora damicornis
Seria topora h ystrix
H
B
X
B
sp-sr
Harriott (198313)
Sammarco (1982)
PORITIDAE
Goniopora columna
Goniopora dijboutiensis
Goniopora lobata
G
G
G
S
S
S
G
G
G
G
G
G
S
S
S
S
S
S
Oct/Nov. 6
Nov. 6
Oct. 6
Nov. 6
Nov. 5-6
Oct, 6
Nov. 6
Nov. 5-7
Nov. 6
Oct-Jan
Willis et al. (1985)
Willis et al. (1985)
Willis et al. (1985)
Babcock et al. (1986)
Babcock et al. (1986)
Willis et al. (1985)
Willis et al. (1985)
Babcock et al. (1986)
Babcock et al. (1986)
Harriott (1983a)
Dec. 5-7
Kojis & Quinn (1981 b)
Goniopora minor
Goniopora norfolkensis
Goniopora tenuidens
Gonipora sp. 1
Gonipora sp.
Poriles australiensis
Porites cylindrica
(as P. andrews~)
W.S/sr,
G
(24
O/O
S
hermaphroditic)
1
Nov. 6
Dec, 1, 4-7, 8
Nov. 6
G
S
Jan, 4-8
Porites lu tea
Nov-Jan
Nov, 5-7
G
B
sp-sr-f
Porites m urrayensis
S
Nov. 5-7
Porites solids
G
B
sr-f
Porites stephensoni (as P,haddonj) X
34G-109H:2x 8B:136S.lx
Central Pacific (Gu_aam,MarGaL,-Islands,Palau)
ACROPORIDAE
X
B
Yr
Acropora bruggernanni
H
X
sr
Acropora cerealis
S
Jul. 4-5
X
B
Jun/Jul. 1 4
Acropora corym bosa a
Acropora delica tula
X
B
Jun/Jul. 1-3
Acropora humilis
H
S
Aug. 7
H
S
sr
Acropora h ystrixC
X
sr
Acropora irregularisd
H
Aug. 4-7
S
Acropora nasuta
H
S
Jul, 7
Acropora ocellata
H
S
Jul. 5-6
X
B
X
Acropora pala wensis
X
sr
Acropora smithie
H
S
JuVAug. 6 7
Pontes cylindnca
Porites lobata
G
"
S
Willis et al. (1985),Babcock et al. (1986)
Kojis & Quinn (1981b)
Babcock et al. (1986)
Kojis & Quinn (1981b)
Hamott (1983a)
Babcock et al. (1986)
Kojis & Quinn (1981b)
Babcock et al. (1986)
Marshal1 & Stephenson (1933)
Atoda (1951a)
Heyward (1989)
this study
Stimson (1978)
this study
Stirnson (1978)
this study
Heyward (1989)
Heyward (1989)
this study
this study
this study
Kawaguti (1940) in Fadlallah (1983)
Heyward (1989)
this study
Note: all taxonomic assignations for Central Pacific are according to Randall (1983, pers. comm.) and samples of each species are
deposited in the University of Guam Marine Laboratory reference collection
Synonymized with A. cytherea by Veron & Wallace (1984)
Synonymized with A. selago by Veron & Wallace (1984)
L Synonymized with A. cerealis by Veron & Wallace (1984)
%ynonymized with A. danae by Veron & Wallace (1984)
' Synonyrnized with A. robusta by Veron & Wallace (1984)
M a r Ecol. Prog. Ser. 60: 185-203. 1990
192
Table 1 (continued)
Specles
Sex
Pacific
Central Pacific (Cuam, Marshall, Islands. Palau)
ACROPORIDAE
Acropora squarrosa'
H
X
Acropora striata
Acropora surculosag
Acropora ten uis
Acropora valida
H
H
H
Acropora varia bilish
Astreopora randalli
Montipora foveola fa
Montipora verrucosa
H
H
H
H
CARYOPHYLLIDAE
Euphyllia glabrescens
DENDROPHYLLIDAE
Balanoph yllia sp.
FAVIIDAE
Fa via rna thaii
Favia steUigera
Favites abdita
Favites flexuosa
Goniastrea aspera
Goniastrea e d w a r d s ~
Goniastrea retiformis
Leptoria phrygia
Montastrea curta
Pla tygyra daedalea
Pla tygyra pini
FUNGIIDAE
Fungia fungites
Heliofungia actiniform~s
OCULINIDAE
Acrhelia horrescens
Galaxea fascicularis
(as G. aspera)
POCILLOPORIDAE
Pocillopora damicornis
(as P. caespitosa)
Pocillopora elegans
(P. meandrina?)
Pocillopora verrucosa
Stylophora pistjlla ta
Sen'atopora h ystrjx
PORITIDAE
Goniopora fruticosa
Porites cylindrica
X
X
X
Mode
Timing
Source
Jul. 2-3
Jun/Jul, 3-4
J a n . 1-8
Jul, 7-8
Jul. 7-8
sr
Jul. 6 7
sr
Jul. 8/Aug, 1
Jul. 2-3
Jun. 7-8
Sep. 2-?
this study
Stimson (1978)
this study
this study
Heyward (1989)
this study
Heyward (1989)
this study
this study
this study
X
Kawaguti (1941)
Yr
Abe (1937)
sr
JunIJul, 6 8
Jun/Jul. 5-7
sr
sr
Oct/Nov, l
sr
Jun/Jul, 7-8
Jul. 1-2/7-8
sr
Jul, 7-8
Aug, 6-7
Jul, 7-8
sr
Heyward (1989)
this study
this study
Heyward (1989)
Heyward (1989)
Abe (1937)
Heyward (1989)
thls study
this study
Heyward (1989)
this study
this study
this study
Heyward (1989)
sr
Sep-Apr, 1
Heyward (1989)
Abe (1937)
Yr
Jul/Aug, 1-3
Yr
Kawaguti (1941)
this study
Atoda (1951b)
Yr
yr. 1-3
Jun/Jul/Jan, 7-3
yr, 2-3
Jan, 3-4
Hada (1932). Kawaguti (1941)
Atoda (1947a)
Stlmson (1978)
Richmond & Jokiel (1984)
Stimson (1978)
Jun/Jul, 1-3
J a n , 1-4
yr, 5-7
sr/\v, 1-5
S t ~ m s o n(1978)
Atoda (1947b)
Kawaguti (1941),Atoda (1951c),Stlmson (1978)
this study
this study
Note. all taxonomic assignations for Central Paclf~care according to Randall (1983, pers. comm.) and samples of each species are
deposited In the University of Guam M a n n e Laboratory reference collection
Synonyrn~zedwith A. loripes by Veron & Wallace (1984)
g Synonymlzed with A. hyacinthus by Veron & Wallace (1984)
h Synonymlzed with A. valida by Veron & Wallace (1984)
'
R ~ c h m o n d& Hunter: Reproduction a n d recruitment of corals
193
Table l (continued)
Species
Sex
Mode
Tlm~ng
Pacific
Central Pacif~c(Guam. Llarsh&lI-Isla~,-~a~a_u],
PORITIDAE
G
S
Jul. 7-8
Porites lobata
G
X
sr
Pontes lutea
G
X
X
Por~tes(Synarea) rus
4G 28H-3-3'12x 14B.28S.3':2x
-H
ACROPORlDAE
Acropora cytherea
Acropora hurnilis
Acropora valjda
Monhpora dilatata
S
X
B
B
sr-f
sr-W
Edmondson (1929).Edmondson (1946)
Edmondson (1929). Edmondson (1946)
X
B
Yr
Edmondson (1929). Edrnondson (19461, Stlmson
(1978)
FUNGllDAE
Fungia scutarm
G
S
Jul-Sep, 5
Krupp (1983)
POCILLOPORIDAE
Poc~lloporadan~icomis
H
B
yr. 5-1
Jun/Jul/Aug, 1
1''. 5
yr. 6-8
yr. 3-5
Edrnondson (1946),Harrigan (1972)
Reed (1971)
St~mson(1978)
R~chmond& J o k ~ e (1984)
l
G
G
G
Jun-Aug. 5
Aug-Sep, 5-6
Aug, 7-8
sr
Hunter
Hunter
Hunter
Hunter
Jun. 5
Jun. 5
Jun, 5
Jun. 5-8
Jun. 5-8
Jun, 5
Jun. 5
Jun, 7
Jun. 7
Jun. 5
Jun. 5
Jun. 5
Jun, 5
Jun. 5-6
Jun. 5-6
Jul, 6/Aug, 6-7
R~chmondpers. obs.
Heyward e t al. (1987). Richmond pers obs.
Richmond pers. obs.
Heyward e t al. (1987). Richmond pers. obs.
Heyward et al (1987)
Heyward e t al (1987)
Richmond pers. obs.
Heyward et al (1987)
Heyward e t al. (1987)
kchrnond pers. obs.
k c h m o n d pers. obs.
K Sakai pers. comm.
K Sakai pers. comm.
Heyward e t al. (1987)
Heyward e t al. (1987)
Heyward e t al. (1987)
'Type Y'
'Type B'
PORITIDAE
Porites compressa
Porites e vern~anni
Pontes loba fa
Por~
tes brigharni
H
H
H
S
S
S
H
X
Grigg et al. (1981)
Gngg et a1 (1981)
Gngg et al. (1981)
Heyward (1985)
Heyward (1985)
Hunter (1989)
Heyward (1985)
Heyward (1985)
Heyward (1985)
Hunter (1989)
Heyward (1985)
DENDROPHYLLIDAE
Dendrophyllia rnanni
Tubastrea coccinea
(as T aurea)
FAVlIDAE
Cyphastrea ocellina
S
thls study
Heyward (1989)
thls study
Jul. 5-6
Jun/Jul/Aug
Jul/Aug, 5-7
Jul. 1
sr-f
Jul, 5-6
Jul, 1
Jun/Jul, 1
Jul, 5-6
Montipora flabella fa
Montipora s t u d e q i
A4on hpora verrucosa
(sensu Vaughan 1907)
Mon t~poraverrilli
H
Source
4G:6H:3' 4x
S
S
S
B
5B:9S:3'
Oklnawa
ACROPORIDAE
Acropora an thocercis
Acropora cytherea
Acropora dig1tifera
Acropora florida
Acropora formosa
Acropora g r a n d ~ s
Acropora hyacyn thus
Acropra la tistella
Acropora micropthalma
H
H
H
H
H
H
H
H
H
S
S
S
S
S
S
S
S
S
Acropora nob~lis
Acropora tenuls
Acropora vahda
A4ontipora aequ~tuberculata
Montipora digita fa
Montipora effusa
H
H
H
H
H
H
S
S
S
S
S
S
X
(1988). Hunter & Hodgson unpubl
& Hodgson unpubl.
& Hodgson unpubl.
& Hodgson unpubl.
Mar. Ecol. Frog. Ser. 60: 185-203, 1990
194
Table l (continued)
I Species
Sex
Pacific
Okinawa
ACROPORIDAE
Mon hpora turgescens
FAVIlDAE
Favia pallida
Favites chinensis
Goniastrea aspera
Platygyra pini
Platygyra ryukuensis
Mode
Timing
Source
Jun. 5
Heyward et al. (1987)
Jun. 5
Jun. &7/Jul, 7
Aug, 7
JunlJul, 5-6
Jun. 2
Jul, 7/Aug, 6-7
Heyward
Heyward
Heyward
Heyward
Heyward
Heyward
et
et
et
et
et
et
al. (1987)
al. (1987)
al. (1987)
al. (1987)
al. (1987)
al. (1987)
FUNGIIDAE
Fungia sp.
MUSSIDAE
Loboph yllia corym bosa
Symphyllia recta
Jul. 7
Heyward et al. (1987)
Jun, 6
Jun. 5
Heyward et al. (1987)
Richmond pers. obs.
OCULINIDAE
Galaxea fascicularis
JudJuYAug, 6-7
Heyward et al. (1987)
PORITIDAE
Gonipora queensland~ae
Yamazato et al. (1975)
Eastern Pacific - work in progress
After 2 yr, only immature ovaries observed (spring). No complete gametogenesis, spawrung or
Pocillopora damicornis
planulation observed (Richmond 1985)
Spermanes and ovaries observed near maturity during summer (A. Yedid pers. comm.)
Pocillopora elegans
Planulated Jun through Nov during both 1984 and 1985 (Richmond unpubl.)
Tubastrea aurea
Red Sea
ACROPORIDAE
Acropora eurystorna
Acropora hempn'chii
Acropora humilis
Acropora hyacinthus
Acropora scandens
Astreopora myriophthajma
May/Jun, 5
X
May/Jun. 7
Jul, 3
JunIJul, 5
JuVAug/Sep, 5
FAVIlDAE
Fa via fa vus
Favites abdita
Goniastrea retiforrnis
Pla lygyra lamellina
OCULINIDAE
Galaxea fascicularis
POCILLOPORIDAE
PociUopora verrucosa
Seriatopora caliendrurn
Stylophora p~stillata
PORITIDAE
Alveopora daedalea
H
OG:15H
Shlesinger & Loya (1985)
Rinkevich & Loya (1979a)
Shlesinger & Loya (1985)
Shlesinger & Loya (1985)
Shlesinger & Loya (1985)
Shlesinger & Loya (1985)
Jun/Jul, 6-7
Aug. 6
Shlesinger & Loya (1985)
X
Jul/Aug, 7
Jun/Jul, 1-2
Aug. l
Rinkevich & Loya (1979a)
Shlesinger & Loya (1985)
Rinkevich & Loya (1979a)
Shles~nger& Loya (1985)
JuVAug/Sep, 6-7
Shlesrnger & Loya (1985)
May. 1
Jul/Aug. 1
May-Dec, 8-1
Dec-Jul, 1-8
Fadlallah (1985)
Shlesinger & Loya (1985)
Rinkevich & Loya (1979a).Shlesmger & Loya (1985)
Loya (19761, Rinkev~ch& Loya (1979a, b),
Shles~nger& Loya (1985)
f-W
B
3B:llS:lx
base, spawners outnumber brooders 168:37 (Table 3).
Spawning is usually associated with higher fecundity,
while brooding produces fewer, larger larvae (Fadlallah 1983).
Shlesinger & Loya (1985)
Szmant-Froelich (1984) proposed that brooders
experience the greatest recruitment success in the
Caribbean, while spawners (particularly acroponds and
poritids) are the more successful recruiters in the Pacific.
Richmond & Hunter: Reproduction and recruitment of corals
195
Table 2. Global comparisons of reproductive characteristics in scleractinian corals from the Caribbean, Red Sea, and Pacific
regions. See individual entries under regional headings for literature citations
Caribbean
No, of species for which
reproductive data are
recorded
Gonochoric
Hermaphroditic
Not reported
Brood
Spawn
Not reported
No. of species in region
Hawaii
Central
Pacific
GBR
Okinawa
Red Sea
19
17
47
145
26
15
6
10
3
4
6
7
4
28
l5
34
109
2
2
24
0
0
15
0
12
7
0
62'
5
9
1
25
3
11
1
2446
3
452
14
28
5
3533
8
136
1
3564
0
242'
Sources: (1) Goreau & Wells (1967). (2) Jokiel (1987), (3) R. Randall, pers. comm., (4) Willis e t al. (1985). (5)Veron (1985). (6)
Sheppard (1987)
Table 3. Summary of reproductive mode for 210 species of
scleractinian corals for which data are available. Entries
reflect cumulative counts, omitting repeats for species found
In 2 or more regions
Hermaphroditic
Spawn
Brood
Unknown
Gonochoric
Unknown
390+
She also noted that Caribbean Porites, which are brooders, have small adult colony size, while the spawning
gonochoric Pacific Porites are large and long-lived.
Pacific Porites species which brood form small, encrusting colonies (P. stephensoni [as P. haddoni], Marshal1 &
Stephenson 1933; P. rnurrayensis, Kojis & Quinn 1981 b;
P. brighanii, Hunter & Hodgson unpubl.).
Timing, seasonality, synchrony, and periodicity
Sexual reproduction in corals may occur yearly (Willis et al. 1985, Babcock et al. 1986), seasonally (Rinkevich & Loya 1979a, b , Szmant 1986, Tomascik &
Sander 1987), monthly (Marshal1 & Stephenson 1933,
Kawaguti 1941, Atoda 1947a, Richmond & Jokiel
1984), or not at all (Grigg et al. 1981, Richmond 1985,
Richmond & Hunter unpubl.). Annual multispecies
synchronous spawning has been observed for over 140
species on the Great Barrier Reef (Harrison et al. 1984,
Willis et al. 1985, Babcock et al. 1986, Harrison pers.
comm.), while asynchrony is exhibited among coral
species in the Central Pacific. Hawaii, Okinawa, and
the Red Sea (Table 1).Synchronous development and
release of gametes among individuals in a population
are important to maximize the probability of successful
cross (and/or self) fertilization. Conversely, a presumed
advantage to multiple spawnings or planulations is to
minimize the effects of a single catastrophic event on
an individual's or population's reproductive success.
Temperature, photoperiod, and nocturnal illumination all appear to be important in providing temporal
cues which may allow synchrony within populations
(Kojis & Quinn 1981 a , Jokiel et al. 1985, Willis et al.
1985, Hunter 1989). The expanding database suggests
that the degree of multispecies synchrony may b e correlated with the annual temperature range experienced
by the corals (Shlesinger & Loya 1985, Babcock et al.
1986). Annual variation in seawater temperature is
2.2 "C in the Central Pacific (Guam; Emery 1962), 3.2 "C
in the Caribbean (Barbados; Tomascik & Sander 1987),
4.0 "C in Hawaii (Oahu; Jokiel 1985), 6.0 "C in the Red
Sea (Eilat; knkevich & Loya 1979 b), 9.8 "C in Okinawa
(Nakamura 1984),and 12.0 "C on the Great Barrier Reef
( M a g n e t ~ cIsland; Babcock et al. 1986).The percentage
of reported coral species spawning within the same
month and lunar phase for each of these regions is 18,
26, 29, 20, 65, and 88 '10,respectively. Oliver et al. (1989)
reported that reproductive seasonality and synchrony
among a n d within coral species distributed from the
southern Great Barrier Reef to Papua New Guinea
diminishes at lower latitudes. The trend appears to b e
one of tighter interspecific synchrony with increased
temperature range. A similar pattern of less restricted
spawning patterns with increasing proximity to the
equator was reported for echinoderm species by Pearse
(1968).
Differences in reproductive seasonality can occur
within a species over its distributional range (Table 4).
Most Great Barrier reef species spawn in the austral
spring, while spawnings in the Central Pacific, Hawaii,
Mar Ecol. Prog. Ser 60: 185-203, 1990
196
Table 4. Global comparisons of reproductive penodicity in coral species which have been reported from 2 or more regions.
Abbreviations are for month and lunar day. Month is divided into 8 phases: 1, new moon, 3, first quarter, 5, full moon, 7, last
quarter; 2, 4 , 6 and 8 indicate intermediate lunar phases (after Shlesinger & Loya 1985); W : winter, sp: spring, sr: summer, f: fall,
yr year-round; ' possibly sterile
GBR
ACROPORIDAE
Acropora cerealis
(as A. hystrixIa
Acropora cytherea
(as A. c o r y r n b ~ s a ) ~
Acropora digitifera
Acropora florida
Acropora formosa
Acropora grandis
A cropora h umilis
Acropora hyacinth~rs
(as A. surculosajc
Acropora latistella
Acropora loripes
(as A. ~ ~ u a r r o s a ) ~
Acropora rnicropthalma
Acropora nasuta
Acropora nobilis
Acropora robusta
(as A. smith.iIe
Acropora selago
(as A. de~icatula)'
A cropora ten uis
Acropora valida
(as A. variabi1is)g
Montipora aequituberculata
Montipora digitata
iMon tipora turgescens
FAVIIDAE
Fa via fa vus
Central
Pacific
Okinawa
Jun/Jul, 1-4
Jun, 5
Jun, 5
Oct, 6
Nov, 6
Oct, 6/Nov, 5-6
Nov. 6
Oct, 6/Nov, 5-7
(spawn)
Jun/Jul, 1-3
(brood)
Aug, 7
(spawn)
Jul, 7-8
-
Jul, 7-8
Aug, 6-7
Oct/Nov, b
Nov, 6
J u n , 5/7
Jun. 5
Nov. 6
Oct, 6
NOV.5-6
Oct, 6-7
NOV.5-6
sp-sr
Oct, 6
OcVNov, 5
Nov, 6
Jul. 6
Jun, 5
Jun, 5
Jun, 5-6
Jun, 5-6
Jun, 5
Nov. 6
Favites flexuosa
Goniastrea aspera
Nov. 6
Oct, 5-7
NOV,5-6
(spawns)
Nov, 6
Nov, 6
Nov. 6
sr
Oct/Nov, 1
(broods?)
Synonymized with A. cerealis by Veron & Wallace (1984)
"Synonymized w ~ t hA. cythcrea by Veron & Wallace (1984)
'Synonyrnized with A. hyacinthus by Veron & Wallace (1984)
Synonymized with A. loripes by Veron & Wallace (1984)
" Synonymized with A. robusta by Veron & Wallace (1984)
Synonymized with A. selago by Veron & WaIIace (1984)
g Synonymized with A. valida by Veron & Wallace (1984)
'
Jun, 3
Jul. 2-3
Oct/Nov, 6
Nov. 6-7
Jul. 7-8
Jul, 7-8
sr
-
"
J u n , 5?8
Jun, 5?8
Jun, 5
Jun, 7
NOV.5-6
Oct/Nov. 6
Nov, 6
NOV.5-6
Nov. 6
a
Red Sea
Nov, 6
Favia mathaii
Fa via pallida
Fa via stelligera
Favites abdita
Favites chinensis
Goniastrea edwardsi
Goniastrea retiformis
Leptoria p h r y g ~ a
Hawaii
Jun, 5
Jun, 6-7/Jul, 7
Aug, 7
Jun/Jul, 5-6
Jul, 3
Richmond & Hunter Reproduct~ona n d recruitment of corals
197
Table 4 (continued)
FAVIIDAE
Montastrea c ~ l r t a
Platygyra daedalea
Platygyl-a lamellina
Pla tygyra plni
FUNGIIDAE
Fungia fungl tes
Heliofungla actinifo~-mls
MUSSIDAE
Lobophyllia c o r y n ~ b o s a
Symphyllla recta
OCULINIDAE
G a l a s e a fascicularis
POCILLOPORIDAE
Poc~lloporadanllcornis
GBR
Central
Pac~f~c
Nov, 6
Oct/Nov, 6-7
Nov, 6-7
Aug. 6-7
Jul, 6-7
Nov, 6
sr
Oct/Nov, 6
Oct/Nov, 5
sr
Sep-Apr, 1
Por7tes lutea
Red S e a
-
Jul/Aug/Sep, 6-7
(spawns)
Jun. 6
Jun. 5
Oct/Nov, 6
(spawns)
Jul, 2-3
(spawns)
Y'
(broods)
yr, 1-5
yr, 1
yr, 3-8
Jun/Jul, 1 4
J a n , 1-4
(bloods)
SI-/W,1-8
y r , ., 7
sp-sr
Hawaii
Jun, 2
Nov. 6
Nov, 6
Pocillopora verrucosa
Seriatopora hystnx
St)~lophorapist~llafa
PORITIDAE
Poriles cylin dl-ica
Pal-~tes loba ta
Okinawa
Jun/Jul/Aug, 6-7
(spawns)
yr. 5-3
(spawns)
Dec-Jul. 1-8
Nov, 6
Nov, 6
Dec, 1, 4-7, 8
J a n , 4-8
Nov-Jan
NOV,5-7
Okinawa, and the Red Sea occur mostly during summer.
For many species, variations in t ~ m i n gand synchrony of
spawning have been observed within as well as
between the regions summanzed in thls paper (Table 1).
Seasonal (latitudinal) variability withln regions (e.g.
between Palau, Enewetak and Guam) may be of sufficient magnitude to cause differences in reproductive
timing. In addition, spawning may occul- at different
times for different sections of a single colony, or for
different colonies within a population (Willis et al. 1985,
Babcock et al. 1986, Hunter & Richmond unpubl.).
While temperature may be the seasonal cue, nocturnal illumination (lunar phase) may provide the 'fine
tuning' for the particular night or nights of gamete or
planula release. Both brooding and broadcasting
species have been shown to cue on night-time illumin a t ~ o n(Jokiel et al. 1985, Hunter 1989). It has also been
suggested that tidal regime and onset to darkness may
play roles as 'forcing functions', determining the actual
time of day when spawning will occur (Harriott 1983 a ,
Babcock et al. 1986, Hunter 1989).
A distinct lunar planulation cycle was reported for
Stylophora pistlllata in Palau (Atoda 1947 b ) , while Red
Sea populations of thls specles show no lunar
synchrony (Rinkevich & Loya 1979 b). Lunar periodicity
of planulation was found to d~fferbetween populations
of Pocillopora darnicojnis at Enewetak and Hawaii, and
within populations in Hawaii ( h c h m o n d & J o k ~ e l
1984). The 'Type B' morph of P. darnicornis planulated
consistently at first quarter moon, while 'Type Y' planulated at last quarter. Van Moorsel (1983) proposed
identification of a new species of Agaricia based partly
on its distinct planulation schedule.
Differences in timing among allopatric populations of
a species may represent adaptations to local environmental parameters and cues. Richmond & Joluel (1984)
suggested that asynchrony among sympatric populations of an identified 'species' may b e the result of the
immigration of planulae from one region into another.
Reproductive isolation, in this case via temporal
mechanisms, allows for divergence and eventual specia t ~ o n .Such reproductive differences within nominal
198
Mar Ecol. Prog Ser 60: 185-203, 1990
Fig. 1 Acropora tenuis.
Section through a ripe colony of a s~multaneoushermaphrodite, 1 wk prior to
spawning. Egg and sperm
are found within the same
polyp. Egg diameter is ca
0.5 mm
I
species raise questions concerning taxonomy based on
morphological characteristics alone.
Asexual reproduction in corals
Corals possess the ability to reproduce asexually,
which is displayed by a variety of mechanisms. Asexual
reproductive processes include formation of 'polypballs' (Rosen & Taylor 1969), polyp bail-out (Goreau &
Goreau 1959, Sammarco 1982, Richmond 1985), asexual production of planulae (Stoddart 1983), and fragmentation (Highsmith 1982).
Asexual reproduction via fragmentation appears to
be important for many coral species, and especially for
populations living at the extremes of their physiological
limits. Grigg et al. (1981) reported lack of mature
gonads in populations of 3 species of A c r o p o r a from the
Northwest Hawaiian Islands. Reproduction via fragmentation appeared to be the major means of population growth. Likewise, Pocillopora d a m i c o r n i s populations in the eastern Pacific had not produced mature
gonads nor planulae during a 2 yr study, yet were the
dominant reef species off the coast of Panama (Richmond 1985). Eastern Pacific P. d a m i c o r n i s exhibit
highc,r colony growth rates than Central Pacific populatiuns, which enhances population growth via fragmentation (Richmond 1985). High bioerosion rates on corals
in the eastern Pacific are proposed as making fragmen-
tation important for massive species such as P a v o n a
c a c t u s (Highsmith 1982).
Asexual reproduction of corals is found in all r e g o n s
covered in this review, but appears to dominate in
areas which are marginal for coral growth, including
the eastern Pacific (Richmond 1985), the northwest
Hawaiian Islands (Grigg et al. 1981), southwestern
Australia (Stoddart 1984), and possi.bly Bermuda
(Wyers 1985). Within regions with optimal conditions
for coral growth, asexual processes may dominate
specific habitats including areas of high wave energy
(Tunnicliffe 1981), soft or unconsolidated substrata
(Gilmore & Hall 1976), and stable, undisturbed sites
(Hunter 1988, unpubl.). In areas where sexual reproductive processes are prevalent, asexuaI reproduction
may augment recruitment at any time, especially during periods of environmental stress or disturbance
(Highsmith et al. 1980).
Enhanced colony growth rate and subsequ.ent fragmentation may result from allocation of energy away
from production of sexual products, notably in environments where sexual processes may be physiologically
constrained (Richmond 1987 a ) . The occurrence of
sterile populations has been described for other
invertebrate taxa as well (Mileikovsky 197 1). Asexual
reproduction has the advantages of not requiring a
partner, propagating locally adapted genotypes, and
providing a refuge-in-size from predation and burial by
sediments.
Richn~onti& Hunter Reproduct~onand recruitment of corals
PLANULA LARVAE
Coral planulae can result from elther internal fertilization and brooding, or external fertilization of
spawned gametes and subsequent development outside the parent colony (Harrigan 1972, Babcock & Heyward 1986). Stoddai-t (1983) suggested that planulae of
Poc~llopora damlcornls may also be produced asexually, based on similarities of parental a n d planular
multllocus genotypes and adult population structure.
Two other species, Tubastl-ea cocclnea and T.
dlaphana, were also found to have planulae with
lsozyme patterns identical to their broodparents, while
planulae of Acropora pallfera and Serjatopora hystnx
had genotypes consistent with sexual origin (Ayre &
Resing 1986).
The brooded planulae of Pocillopora damicornis contain symbiotic zooxanthellae upon release from the
parent, as well as a large quantlty of lipid (Fig. 2). With
the additional ability to feed whlle planktonic, these
planulae remain competent for over 100 d , a period
sufficient to allow dispersal over large distances (Richmond 1981, 1987 a ) . Planulae resulting from spawned
gametes (Fig. 3) may lack zooxanthellae upon fertilization (known exceptions being poritids and Montipora
spp., whose eggs contain maternal zooxanthellae), but
eventually acquire the algal cells from the environment, usually after settlement and metamorphosis
(Babcock 1989). Planulae of Fungia scutaria acquire
zooxanthellae after release from the parent colony but
before metamorphosis (Krupp 1983).
The spawned larvae of Acropora tenuis do not contain zooxanthellae, and have a shorter competency
period (ca 20 d ) than the brooded larvae of Poallopol-a
Fig. 2 Poc~llopol-a damlcornis. Brooded planula of a coral
Note the bands of zooxanthellae, and the oral opening. T h e
larva 1s ca l mm In length
199
dan~icornis(Richmond 1989). Larval competency (the
abllity of larvae to successfully settle and metamorphose) is a major factor affecting the distrlbution of
coral species, particularly for reefs in the eastern
Pacif~c.I t has been proposed that the present coral
fauna of the eastern Pacific is the result of long-distance dispersal of planulae from Central Paclflc stock
(Dana 1975, Rlchmond 1987 b). Some instances of
llnlited distrlbution patterns and endemlsln may b e the
result of abbreviated larval competency periods.
RECRUITMENT
Reproductive success may best b e measured by
recruitment. Recruitment of both sexual and asexual
propagules is medlated by biotic factors, such as predahon and competition, and by abiotic factors such as
environmental variability and disturbance. Sexual
recruitment of corals is a function of several parameters
including the timing of reproduction, competency
periods of planula larvae, current regimes, availability
of substrata, and densities of predators and competitors
(Bii-keland et al. 1981, FItzhardlnge 1985, Babcock
1989). Several studies have found evidence for 'open
populatlons' with non-localized sources of coral
planulae (Wallace 1985 a , Babcock 1989), but others
have suggested that reefs can be self-seeded (Baggett &
Bright 1985, Sammarco & Andrews 1988, 1989,
Andrews et al. 1989).In some areas, juvenile abundance
1s directly related to adult cover (Bak & Engel 1979,
Fig. 3 Acropol-a tenuls. Spawned clusters of e g g s sul-I-ounding
sperm from a coral. Each cluster contalns between 9 and 16
eggs around a single sperm packet Cluster diameters range
between 1.3 and 1 6 mm
200
Mar Ecol. Prog. Ser 60: 185-203, 1990
Rylaarsdam 1983), although this is not always the case
(Fitzhardinge 1985, Harriott 1985).Relative recruitment
rates have been shown to vary from year to year, and
among sites on the Great Barrier Reef (Wallace 1985 a ) .
There appears to b e a n inverse correlation between
success of larval recruitment and propagation by fragmentation in some coral species (Kojis & Quinn 1981 b,
Highsmith 1982, Wallace 1985b). Sammarco (1985,
1987) reported that, while planular recruitment of
Acropora spp. on the Great Barrier Reef is high,
juvenile recruits are rare in the Caribbean a n d populations there are structured primarily by asexual processes. Temporal and regional variations in biotic and
environmental factors can cause differences in reproductive a n d recruitment patterns of coral communities.
The pan-Pacific coral Pocillopora damicornis provides a n example of how life history characteristics
may vary with respect to local conditions (Richmond
1985, 1987 a). Enewetak Atoll a n d Hawaii arc characterized as having low rates of p r e d a t ~ o non P. damicornis, relatively low seasonal variability, yet relatively
high frequencies of mortality-causing disturbances
(typhoons and winter storms). P. damicornis colonies
planulate monthly throughout the year in Hawaii and
the Central Pacific, a n d the oldest colonies observed
are estimated to b e less than 10 yr old (branch lengths
< 20 cm). The eastern Pacific of Panama is characterized as having high rates of predation on P. damicornis, high levels of seasonal variability, and low frequencies of mortality-causing events. In this area, P.
damicornis has never been found to release planulae,
has a higher colony growth rate, a n d is the dominant
coral in terms of competitive interactions with other
corals, with some colonies estimated as over 70 yr old
(branch lengths > 240 cm).
Theories on the evolution of life history characteristics have been proposed which suggest that under
conditions of environmental instability, where lethal
disturbances occur at a relatively high rate, formation
of large numbers of motile propagules should occur,
while under stable conditions, selection will favor
clonal (asexual) growth (Williams 1975, MaynardSmith 1978).Likewise, under conditions of low juven~le
versus high adult mortality, a n d relatively low competitive ability versus competitive dominance, selection
will favor the sexual mode (Abrahamson 1980, Douglas
1981). Pocillopora damjcornis fits these predicted patterns over its distributional range.
RESOURCE MANAGEMENT IMPLICATIONS
As coral reefs throughout the world are showing
signs of degradation, management of reef resources is
becoming a growing concern. In the case of corals
which spawn during a very brief period each year, the
presence of contaminants such as petroleum products,
pesticides, herbicides and heavy metals from soils may
prevent successful fertilization of eggs by sperm, and
hence, severely limit coral recruitment (fichmond
unpubl.). It is suggested that pollution levels which
may not affect adult coral colonies could still be
responsible for the eventual loss of reefs if reproductive
processes are disturbed. In Guam and in Okinawa
(southern Japan), the peak coral spawning occurs during the rainy season, when levels of coastal contamination via runoff are at their highest (pers. obs.).
With reef degradation and destruction occurring on a
global scale, an application of the reproductive data is
in the area of reef recovery. Areas of reef which have
been destroyed may b e re-seeded, and the most efficient means will depend on local conditions. In areas
where sedimentation is high, corallivores are present in
large numbers, and/or disturbance rate is low, cementing larger numbers of smaller fragments may be more
effective based on reproduction and recruitment data.
In areas where environmental conditions support
sexual reproduction of corals, juvenile mortality is
expected to be relatively low, and suitable substratum
is available, transplantation of gravid adult colonies
into a n area may result in highest return of effort. Reef
re-seeding is an expensive process (Harriott & Fisk
1989) and management efforts might best be expended
in coral reef protection and conservation.
SUMMARY
Data on the reproductive biology of 200+ scleractinian corals indicate several apparent trends. Species
which broadcast spawn outnumber those which brood
planula larvae. Broadcast spawners typically have
limited, annual spawning periods, while most brooders
are iteroparous, releasing larvae over a large part of the
year. Most corals reported to date are hermaphroditic,
some of which have been found to self-fertilize readily
in the laboratory; the capacity for selfing is low to nonexistent in other hermaphroditic species. There are
reports of coral populations which are apparently
sterile, particularly those at the extreme limits of their
distributions.
There appear to be several geographical trends in
coral reproduction. The majority of species on the
Great Barrier Reef participate in a n annual synchronous spawning event following full moon in the austral
spring. In other areas of the Pacific, the Caribbean, and
the Red Sea, there is a greater partitioning of spawning
periods over more months, days, and lunar phases.
Degree of synchrony among species may be related to
seawater temperature ranges in each region.
kchmond & Hunter: Reproduction and recruitment of corals
M o d e of reproduction within a given taxa is g e n e r ally conservative, while timing m a y b e variable within
a n d a m o n g species. Differences i n reproductive patterns m a y represent adaptations to local environmental
conditions or m a y represent g r o u p s which h a v e not
b e e n sufficiently taxonomically differentiated. A point
w h i c h clearly e m e r g e s from t h e d a t a r e v i e w e d in this
p a p e r is t h e n e e d for examination of reproductive
characteristics i n conjunction with skeletal characters
to elucidate w h a t m a y b e real p r o b l e m s i n coral taxonomy. Do o b s e r v e d reproductive differences indicate
that speciation h a s occurred a m o n g populations? If w e
a c c e p t t h e biological species definition, it is evident
that morphological characters alone m a y fail to detect
valid biological differences. As more studies of reproduction a r e p u r s u e d , resulting information will e n a b l e
d e v e l o p m e n t of a m o r e a c c u r a t e u n d e r s t a n d i n g of t h e
ecology a n d evolutionary biology of t h e Scleractinia.
Acknowledgements. We gratefully acknowledge UNESCO
for sponsoring the meeting on Interoceanic Comparisons, held
at the University of the South Pacific, Suva, Fiji in March,
1986, for which this review was originally prepared. We thank
Drs C. E. Birkeland, P. L. Jokiel, R. A. Knzie, D. A. Krupp and
P. W. Sammarco for comments on manuscript drafts, R. Randall for taxonomic identifications, D. Hopper. F. Te, A. Kerr.
and A. Goodis for assistance with field and laboratory examination of Guam corals, and Dr K. Yamazato for access to the
corals of Ohnawa. This is contribution no. 277 of the University of Guam Marine Laboratory.
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This review was presented by Dr. P. W. Sammarco,
Townsville, Australia
Manuscript first received: J a n u a r y 8, 1988
Revised version accepted: October 18, 1989