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Tridacna Giant Clam
Tridacna Giant Clam
219-233
Printed in Great Britain
ABST RACT : Giant clams may be stimulated to spawn by the addition of macer-
ated gonads to the water. I ndividuals of Tridacna maxima collected at Anae Island,
Guam, spawned from N ovember to March. On Palau, Hippopus hippopus spawned
in June and Tridacna crocea, in July.
Tridacna crocea, T. maxima, and H. hippopus displayed a stereotype d deve lop ment
pattern in morphogenesis and rate of deve lopment . Fertilized eggs of T. crocea,
T . maxima, and H. hipp op us had mean diameters of 93.1, 104.5, an d 130.0 psx»,
respe ctively . The day-2 str aight -hinge veligers of T . crocea, T . maxima, and H.
hippopus had mean shell lengths of 155.0, 168.0, and 174.4 pm, respec tively.
Settlement occurred 12, 11, and 9 days after fertili zation at a mean shell length of
168.0, 195.0, and 202.0 pm for T . crocea, T . max ima, and H. bippopss, respectively.
Metamorphosis was basically complete about 1 day after settlement. Juveniles of
T . crocea, T. max ima, and H. hippopus first acqui re zooxanthellae after 19, 21, and
25 days, respectively. Growth rates increase sharply after the acquisition of zoo -
xanthellae. J uvenile shells show first signs of becoming opaque after 47 days for
T . maxima and after 50 days for H. hippopus.
G rANT CLAMS are protandric fun ctional herma- form ation, and initial organo genesis. T he larval
phrodites (Wada 1942, 1952). The male p hase of and postlarval develop ment of Tridacna maxima
th e gonad develops first and then the female and T. squamosa have been described by La-
phase. I n the adults, female and male follicles Barbera (1975) in his efforts to understand the
lie side by side throughout the extent of the mo de of transmiss ion of zooxanthellae from
gonad, and both eggs and sperm matur e generati on to generation and to elucidate the
simultaneously. mechanism of development and pos sible evo lu-
The giant clams, in contrast to the majority tion of the unique fo rm of tridacnids.
of func tional hermaphrodite biva lves, do not The motive behind investigating the early
discharge egg s and sperm simultaneously when life history of the giant clam is twofold . First,
spawning . Gia nt clams first spawn sperm and giant clams are being subjected to an ever-
th en egg s. The spawning reaction has been increasing fishing pressure because of their food
described by Wa da (1954). Stephenson (1934), and shell value. D ata abo ut the early life history
in her wo rk with Hippopus hippopus, provided of the clam might alleviate some of this pressure
useful information for jud ging the gonad condi- by providing conservationists with the informa-
tion of giant clams. tion they need to support efficient population
LaBarbera (1974) descr ibe d the transitional man agement. Second we must obtain a basic
stage between th e trochophore and the veliger knowledge of the early life history of the giant
larva of Tridacnasquamosa, prodissoconch I shell clam before we can determine the feasibility of
a large-scale aquaculture of the animal for food .
1 Supported by the Sea Grant Coherent Project On Guam, research was conducted on
(04-4-158-4) awarded to the University of Gu am Marine T . maxima at the University of G uam Marine
Labo ratory; contr ibution no. 72, Un iversity of Gu am Labo rato ry from September 1973 to March
Marine Labo ratory . Manuscript accepted 10 September
1975.
1975. Research was carried out on Tridacna
2 University of Guam Marine Lab or ator y, Post Office crocea, Tridacna derasa, Tridacna gigas, and
Box EK, Ag ana, Gu am 96910. H. hippopus in Palau at the Micronesian Mari-
219
220 PACIFIC SCIENCE, Volume 30, July 1976
culture D emons tration Center (MMD C) fr om of seawater containing sperm were adde d to the
7 June to 19 Aug ust 1974. con tainer to fertilize the eggs . A net system of a
210-,um nylon mesh over an 88-,um nylon mesh
was used to isolate th e eggs. Large detritus wa~
MATERIAL AND METHODS
caught on the top net , egg s were caught on th e
On Guam, specimens of Tridacnamaxima were bottom net, and sperm and water passed
collected exclusively at Anae Island at depths of th rough both nets. The eggs from th e bottom
3-1 0 meters. Because thi s species is usu ally net were gently washed into a 5-liter glass
found embedded in the substrate, I removed it container containing filtered seawater. T he
by chipping away the sur round ing substrate number of eggs was th en adjusted so that only
with a hammer and chisel, and th en by cutting one layer of eggs covered the bottom after
the byssus clean with a thin-bladed filet knife. settling had occurred . A settling and decanting
This method was preferred as it did not injure technique was used to wash th e egg s th ree times,
the ani mal. after which they were placed in several500-liter
Specimens of H . bippopus and T. derasa we re tanks at a concentration of about 10,000 eggs
collected in Palau at N garchabal Island at depths per tan k. A prolon ged washing pe riod decreases
of 6-1 0 meters. Sp ecimens of T. crocea we re survival, so fertilized eggs were washed only
collected from th e reef at th e MM D C dock at a thr ee time s before being placed in a large tank.
depth of 1-4 meters, with the same metho ds T he 500-liter h olding tan ks contained seawater
being used as described for T. maxima. Speci- th at had been filtered th rough an 88-,um nylon
mens of T . gigas were collected at a depth of mesh th at removed large detritus and potential
about 3 meters near the Malakal Passage light- predators. Larval and juvenile T. maxima were
h ous e; here a rope sling was attached to the raised in the 500-liter rearin g tanks. N o addi-
clam and th e animal was h oisted to th e bottom tional nu trients were added to th ose pres ent
ofth e boa t. T hen, with th e boat being run at slow natu rally in the seawater. The water in the
speed, the clams were towed underwater to rearin g tan ks for juv eniles was changed once
MMDC. a wee k.
F our to 12 individuals of T . maxima were Complete spawning was also stimulated when
collected monthly at Anae Island, Guam, from 22 T . maxima were transferred from a 500-liter
November 1973 to March 1975, except for the holding tank at 26.1° C to a lOOO-liter h olding
m onths of D ecember 1973, and Octob er 1974. tank that was being filled. Ini tially, th e holding
They were sub jected to spawning stimulus tan k had on ly 6 inch es of water in it. A s more
(macerated go nad) th e day after collection. If no water was added, a circul ar flow developed in
eggs were spawned, th e animals were dissected the tank. T he water being added was at 29.7° C.
and the go nads we re insp ected microsc opically. Once the tank had been filled, th e water cooled
In spawning experiments only clams greater than to 28.5° C and several clams spawned.
130 mm in shell length were used , as T. maxima To test substrate preference, I used various
does not attain full sexual maturity, i.e. , develop subs trat es (bare rock, ro cks covered with coral-
the female phase, until it attai ns a shell length of line algae, glass, shells, and dead coral) as spat
110-130 mm . G onad sp ecimens were preserved collector s duri ng the settl ement of T . maxima.
in 70 percent isop ropyl alcohol. Subs trates were suspended in the tan ks from a
The fecundity of T . maxima was estimated series of mon ofilament slings atta ched to sticks
from eight clams collected during F ebruary that had been placed across th e tops of the
1975. The number of eggs spawned by each of rearing tanks .
th e indiv iduals in separate containe rs was On Palau, susp end ed glass petri dishes, som e
counted by v olumetric estimate . Five to 10 ml with a sand bottom and others without, were
of macerated gonad was used to stimulate these used as spat collectors for H . hippopus. Sus-
clams to spawn. They were allowed to spawn pended glass petri dish es, some with a sand
until th e water was very dense with sperm". T he bottom and others with an assorted dead coral
clams were transferred repeatedly to new con- bottom , were used as spat collect ors for
tainers until they spawned eggs . Five to 10 ml T. crocea.
Early Life History of Giant Clams-JAMESON 221
T ABLE 1
M ONTHLY D ATA ON G ONAD CONDITION OF A POPULATION OF Tridacna max ima,
ANAE I SLAND, G UAM
R ipe 1 3 8 3 7
9/10 R ipe 3 1 2 2 1 3 1 1
7/10 Ripe 1 2 3 1 2 2 1 3
1/2 Ripe 3 3 3 2 4 3 1 1 1
Spent 4 2 5 3 1
NOTE: Ripe, sp aw ne d eggs after stimu lus; 9/10 R ipe, eg g size from 100-15 0 pm, g onad very larg e, white ;
7/10 Ripe, egg size from 80- 150 psn, g onad lar ge, w h ite ; 1/2 R ipe , egg size from 50-150 psu, gonad small, w hite;
Spe nt, very few to n o eggs p resen t, gona d small, b rown t o o live d rab.
7
U. 10
iJ
~
RESULTS
Q)
e Spawning and Gonad Condition
~
c After the macerated go nad had been added to
a.
VI the water , th e clams would usually spawn sperm
III
Cl
=.0 0 74 3 L 4 . 03 .
within 2 to 3 minutes. T hey continu ed to spawn
Cl sp erm for as long as 6 h ours or mor e, with indi-
w
-..
0
Q)
viduals exhibiting variation in duration . Aft er
the spawnin g of sperm, the spawning of eggs
will proceed if the go nad is full y rip e. If the
gonad is not full y rip e, only the spawning of
..c
E sperm will occur.
;:) The results fro m monitori ng the go nad condi-
Z tio n of T . maxima at Anae Island are given in
Table 1. Field-collected animals stimulated by
6 macerated go nad spawned eggs in N ovember
10 1973 ; in Ja nuary, February, and D ecember
1974 ; and in February 1975. During no other
months could T. maxima at Anae Island be
stimulated to spawn eggs. Spent go nads were
100 150 200 300 found in dissected specimens du ring the
months of November 1973 ; March, Septemb er,
Shell Length (mrn], L
and N ov ember 1974 ; and January 1975. From
FI GURE 1. Logarithmic p lo t o f num bers o f eggs April th rou gh Aug ust, specimens in va rying
spawned in rel ation to sh ell length of Tridacna maxima
at A n ae I sland, Guam .
degrees of rip eness were observe d.
The num ber of eggs spawned by T. maxima
15 H PS 3 0
222 PACIFIC SCIENCE, Volume 30, July 1976
TABLE 2
EARLY LIFE CHRONOLOGY OF THE GIANT CLAMS Tridacna crocea, Tridacna maxima,
AND Hippopushippopus
varied with the size of the individual (Figure 1) ary, and 22 February 1974. The early life
and can be expressed by the equation chronology is a composite of the above
cultures.
F = 0.00743U·03, Tridacna crocea, T. maxima, and H. hippopus
displayed a stereotyped development pattern
where F is the number of eggs spawned and L (Figures 2, 3, 4) that is considered to be typical
is the shell length. for bivalves in general. Development times and
Hippopus hippopus spawned sperm and eggs on sizes at a particular stage exhibit much individual
12 June in Palau. A clam 249 mm in length variation (Tables 2, 3, 4, 5). In general, the three
spawned approximately 25 million eggs. Later species observed exhibited similar development
in August a H. hippopus 280 mm in length was times, if the great range of individual variation
dissected and found to be in a halfripe condition. in these times is considered.
Tridacna crocea spawned sperm and eggs on
7 July. However, more than 30 clams were
Larval Development andBehavior
subjected to spawning stimulus before 2 finally
spawned eggs. After fertilization, typical bivalve spiral
Specimens of T. derasa 390 and 410 mm in cleavage results in a spherical blastula. The
shell length were subjected to spawning rotating ciliated gastrulae may be observed
stimulus on 3 July 1974 and no spawning 7 hours after fertilization for T. crocea and
occurred. The specimens were then dissected T. maxima and after 8 hours for H. hippopus.
and were found to be spent. Two T. gigas 530 Pelagic larvae exhibit typical bivalve develop-
and 560 mm in shell length were found on ment and behavior (D'Asaro 1967, Loosanoff
26 July 1976 to be still in the male phase. and Davis 1963, Raven 1958) with the following
specific variations. The trochophore stage
(Figures 2a, 4a) is reached for T. crocea, T.
Barty Life Chronology maxima, and H. hippopus 15, 16, and 17 hours
The early life chronologies of T. crocea, after fertilization, respectively. The transitional
T. maxima, and H. hippopus are presented in stage between the trochophore and the veliger
Table 2. Tridacna maxima was cultured on four larva (Figure 4b) has been described by La-
different occasions, with fertilization occurring Barbera (1974).
12 November 1973; and 31 January, 14 Febru- Straight-hinge veligers (Figures 2b, 3a, 4c)
223
FI G URE 2. Larval and juveni le T ridacna crocea. A , tr och ophore ; B, day-3 veligers; C, day-l0 ped iveliger ; D, day-25
juvenile.
A BBREVIATIONS : bg , byssus gland ; z, zooxanthellae.
224
A lOOIJ m
aa
vrl
v
5
vr2 e
vr3
tr
V'4
PJ4Jm~1~
B lOOIJm
aa- + -----;'{
li -----\r-'''''-~~ ; ..: .
SS ---T-~--"'--.",,--~
\,
==-.~-+-----+------,I-- ,e
·, :7---'"::'-"'-----/--- 'a a
ss -------=~------;--"'_- - -- ,---c7"=------ - - S
"- --- /-
"-
--- - - ~
FIGURE 3. Internal anatomy of larval and postiarval Tridacna maxim a. A, day-2 veliger; B, pediveliger; C, day-47
juvenile.
ABBRE VIATlONS : a, anus ; aa, anterior adductor ; bg, byssus gland ; dg, digestive gland ; e, esophagus ; f, foot;
g , gill ; i, in testine ; k, kidney ; pa , posterior adductor ; s, stomach ; ss, style sac ; t, telotroch; tr, telotrochal retractor;
v, velum; vr, velar retractors (numbered) ; z, zooxanthellae.
225
FI G UR E 4. Larva l and juvenile Hippopus hippopus. A, trochoph ore ; B, troc hophore-veliger transition stage;
C, day-4 veliger; D, day-l l ped iveliger; E, day-58 juvenile.
A BBREVIATIONS : me, mantle extensio n; vr , velar retractor.
226
TABLE 3
GROWTH IN LENGTH AND HEIGHT BY TIME FOR T ridacna crocea
T ABLE 4
G ROWTH IN LENGTH AND HEIGHT BY TIME FOR T ridacna maxima
TABLE 5
GROWTH IN LENGTH AND H EIGHT BY TI ME FOR Hippopus hippopus
NOTE: Numerals in parent heses follo wing standard deviation s represent ranges .
227
A lOOJ.lm
e
s
aa
vr 1
tr V2
B 100 JIm
j:- -'l-"'-T-
al - ----',--
....
. ..
c st 50J.lm
...........
.....
pa---'r-- ---'--+",/
a ---"c---- '
j ---\---~"",," :.~~
k----'.;--~, /
FIG URE 5. Inte rnal anatomy of larval and postlarval Tridacna crocea, A , day-2 veliger; B, pediveliger ; C, day-25
juveni le.
A BBREVIATIONS: a, anus; aa, anterior adductor; bg, byssus gland; dg, digestive gland; e, esop hagus; f, foot;
g, gill; i, intestine ; k, kidney ; pa, posterior adductor; 5, stomach ; 55, style sac; st, statocyst; t, telotroc h ; tr, telo-
trochal retractor ; v, velum; vr, velar ret ractors (numbe red).
228
A
v-----,~"'--
e---/-----'<,--""'"
s- t - - - - - - + - I,+ -+-- - a a
-3~ __V
bg,---~
+ - - -""""----!iit--e
dg --f----,f'----i-- '---'-
ss--+---+-------.........- t:+--+--aa
-+-,{-- - s
a------' "':'
c
SOlJ m
+7----------~---- z
..........
<.
-.
-';-+".-----'-- - -+- - -+-st
FIGURE 6. Internal anatomy of lar val and postl arval Hippopus bippopus. A, day-2 veliger (see Figure 4C for
arra ngement of retractor muscles); B, ped iveliger ; C, day-27 juvenile.
ABBREVIATIONS : a, anus ; aa, anterior adductor ; af, apical flagella; bg, byssus gland ; dg, digestive gland ;
e, esophagu s ; f, foot; g, gill ; i, int estine ; k, kidn ey ; pa, posterior adductor; s, sto mach ; ss, style sac; st, statoc yst ;
v , vellum ; z, zooxanthellae.
Earl y Life History of Giant Clams-JAMESON 229
rate in creases shar ply (6.8 pm/day for T. It is also interestin g th at H. hipp opus spawned
maxima, 13.9 pm/da y for H. hippopus). T his in the summer months in Australia an d Palau;
corresponds to th e time at which the majority whereas T. maxima spawn ed during the winter
of juveniles had acquired zooxant hellae. months in Guam and Fiji (LaBarbera 1975).
T he sam e species may spawn during the same
month of the year regardless of geog raphical
location. Further investigation is needed into
DISCUSSION
this question.
Obs ervati ons of th e gonad condition of a Many temperature-shifting expe riments simi-
p opulation of T. maxima at Anae Island indi- lar in style to those used in spawning temperate-
cated th at the winter months of N ovember to water bivalves (Galtsoff 1938) were carried out,
March are the most promising for spawning. with various temp eratures and different periods
The height of th e spawning period seems to of exposure being used . N one of these p roce-
occur in February. It appears that th e majority dures stimulated spawning; in some cases they
of the p opulation is in the process of recovering made the animal sick or even killed it. In the
from winter spawning between March and study of tropical bivalves one should consider
Au gust. Som e highl y mature individuals were the different enviro nmental conditions to which
observed at this tim e and some spawning may tropical coral-reef orga nisms are exposed and
occur but probably only to a small extent. not exp ect to find the same reproductive pro-
Spawning seems to increase in frequency cesses or str ategies th at are common to temper-
around Aug ust or September and to pea k again ate water species.
in F ebruary. Spawning in nature may be stimulated by
Step henson (1934) rep orted that H. hippopus water mo vement. Previous experiments de-
spawns in th e austra l summer months of signed to stimulate spawning by changing water
D ecemb er to March on th e reef flat at Low temperature failed, but spawning was stimu-
Isles, G reat Barr ier Reef. She suggested hig h lated when clams were sub jected to a current of
temp eratures du rin g two hot spells as being the circular flowin g water in a holding tan k that was
stimulus for spawning. The first hot spell being filled. In Guam, roug he r than average sea
occu rred from 7 December to th e end of th e states occur from Septemb er to January as a
month, with temperatures rangin g from 28.0°- result of large swells generated by the trade
30.7° C. The second hot spell occurred from winds, which are strongest at this time. At Anae
12-30 Janu ary, with temperatures ranging from Island continuous large swells that cause ve ry
29.0°-32. 0° C. Such hot spells may be a factor in strong currents underwater during one or
stimulatin g spawning among reef-flat animals several winter months are common. The
which are exp osed to temperature fluctuations . coincidence of strong underwater currents
It is doubtful that temperature plays any role during the spawning period at Anae Island may
in the spawning period of T . max ima at Anae further support this idea. LaBarbera (1975)
Island, because the annual temperature range noted the app arent effects of the chan gin g tid e
fo r oceanic surface water aro und Guam is in stimulating the spawning of clams in Fiji.
26.5°-29. 0° C. Deeper waters have insignificant Adding macerated gonad to seawater con-
annual temperature fluctuations ; giant clams taining giant clams seems to be the easiest and
living in deeper waters are p rob abl y influenced most efficient way to stimulate spawning. Wada
in th eir spawning activities by some other factor. (1954) contended th at the active pri nciple that
E vide nce suggests that temperature is n ot a stimulates spawning comes from both the eggs
factor in determining the spawning pe riod of and the primordial oogonia or other ovarian
sub tid al population s. P opulations in relativ ely tissue cells. The activ e principle seems to be
const ant temperature envi ronments may be species-speci ficin effect, as T. maxima would not
genetically disposed to synch ronize their react to stimulation ifT. squamosaeggs were used
spawning behavior to a characteristic envi ro n- and vice versa (LaBarb era 1975), nor would
ment al stimulus th at may be unique to each T. squamosa react to stimulation if H. hippop us or
population's location. T . derasa eggs were used, and T. derasa would
232 PACIFIC SCIENCE, Vo lume 30, July 1976
not react to stimula tion if T . squamosa or from day-2 to the acquisition of zoo xanthellae
H. bippop«: eggs were used (Wada 1954). because energy is being chann eled for various
When spawning giant clams for culture, one developmental changes, such as the change in
must ensur e that th e spawning of eggs occurs in form of th e p rodissoconch shell, leaving less
seawater with a low densit y of sp erm. Spawned available for gr owth in shell length. I ncr eased
eggs with an excess of sperm in th e surrounding growth rat es after the acquisition of zoo xanthel-
jellycoat do not develop normally. This lae are probably brought about in a large part
problem was also obs erved by LaBarbera by th e zooxanthellae th emselves, which, ac-
(1975). cording to G oreau, Goreau, and Yonge (1973),
The pelagic larval period of the three species supply T. maxima with energy fro m p hoto-
observ ed is short (12 days for T. crocea, 11 days synthetates and by digestion of older algal cells.
for T . maxima, 9 days for H. hippopus), and under The decrease in shell length of T . maxima at
better culture conditions could p robably be day 91 (Figure 7) occurs becaus e th e sampl e used
shortened further. LaBarbera (1975) also contained smaller sized clams. G rowth rates
observed similar short pelagic periods when should be understood in the light of th e limited
culturing T. maxima and T . squamosa in Fiji. nutrients available.
Larvae can be cultured adequately in a
stagn ant system but a food supplement and
bacteriostat (Da vis and Calabrese 1969) should ACKNOWLEDGMENTS
be added. Phytoplankton smaller th an the eso-
ph agus diameter of about 8 usi: should be con- My deepest th anks go to Pat Beeman, Frank
sidered. Monocbrysis lutheri or Isocbrysis galbana Cushing, Rich ard Dickinson , Bill Fitz Gerald,
might be suitable. Larval food preference needs J erry H eslinga, Storm Rideout, Ted T anzy,
inve stigatio n. T ob y Tobias, and D an Wo oster for th eir help
After settlement and metamorphosis, juve- with fieldwo rk on Guam .
niles seek a suitable permanent settling spot that I also wish to thank Jim McVey for allowing
will give them maximum protection . If juveniles me to work at the Micronesian Ma riculture
are to be maintained in a particular area after Demonstration Center, Theo Fuziwara and
settl ement, a suitable substrate must be available Ester Yaoca for their help with larval cultures,
to th em, otherwise th ey will leave thei r original and esp ecially Gerh art Pahl for his assistance
settli ng spot and be lost. For no nb oring species with fieldw ork in Palau .
a container with a sand bottom is ad equ ate. For Than ks also go to Clayton Carls on , Lucius
the boring species a spat collector that can give Eldredge, Mike LaBarbera, Lynn Raulerson ,
juveniles protection from as many angles as Masashi Yamaguchi, and C. M. Yonge for
po ssible is preferred. Some nontoxic material revi ewin g the manuscript.
with crevices or pits in it must be supplied for
the boring species.
LITERATURE CITED
After they have settled, juveniles attached
to spat collectors should be rem oved from the D'AsARO, C. N. 1967. The morphology of
stagnant rearing system, which contains much larval and p ostl arval Chione cancel/ata Linne
decaying matter and benthic bacteri a, and be (Eulamellibranchia: Veneridae) reared in the
transferred to a system th at has a current of lab oratory. Bull. Mar. Sci. 17(4): 949-972.
fresh seawater running th rough it. Juveniles DAVIS, H. c., and A. CALABRESE. 1969. Survival
sho uld be raised to about 3-4 ern in length in the and growth of larvae of the E uropean oyster
laboratory before they are introduced into the (Ostrea edulis L.) at different temp eratur es.
field to grow in natural ecosystems. Any size be- BioI. Bull. (Woo ds Hole, Mass.) 136(2):
low thi s is very susceptible to predatio n. T here is 193-199.
need for information on the survival of cultured G ALTSOFF, P. S. 1938. Physiology of repro-
juve niles th at have been introduced back into duc tion of Ostrea virginica. II. Stim ulation of
the field to gro w . spawning in the female oyster . Biol. Bull.
The animals probably show low growth rates (Woods Hole, Mass.) 75 : 286-307.
Early Life History of Giant Clams-JAMESON 233