Aquatic Botany, 8 (1980) 323--336 323

Elsevier Scientific Publishing Company, Amsterdam -- Printed in The Netherlands

FLOWERING UNDER CONTROLLED CONDITIONS BY CYMODOCEA

S E R R U L A T A , HALOPHILA STIPULACEA, SYRINGODIUM ISOETI-
FOLIUM, ZOSTERA CAPENSIS AND THALASSIA HEMPRICHII FROM
KENYA

CALVIN MCMILLAN
Plant Ecology Research Laboratory and Department of Botany, University of Texas at
Austin, Austin, TX 78712 (U.S.A.)
(Accepted 16 January 1980)

ABSTRACT

McMillan, C., 1980. Flowering under controlled conditions by Cymodocea serrulata, Halo..
phila stipulacea, Syringodium isoetifolium, Zostera capensis and Thalassia hemprichii
from Kenya. Aquat. Bot., 8: 323--336.

Flowers have been reported rarely for tropical seagrasses of the Indo--Pacific, but after
transplanting from Kenya to experimental cultures, flowering was observed in five out of
nine species. For Cymodocea serrulata (R. Br.) Aschers. & Magnus, pistillate flowers were
produced at 31/27°C (day/night) after eight months in culture, and flowering was conti-
nuous for three months under these conditions, but not at lower temperatures. In Halo-
phila stipulacea (Forsk.) Aschers., pistillate flowers were produced at 24°C, but were also
produced over a range of higher temperatures if the nutrient status of the artificial sea-
water medium was altered. Flowering spathes of Zostera capensis Setchell appeared in the
experimental cultures seven months after transplanting to a 16--18°C temperature regime.
In Syringodium isoetifolium (Aschers.) Dandy, flowers were not produced during nine and
a-half months at any of a range of temperatures, but were produced subsequently at 26
and 24°C in a specially prepared seawater medium. Thalassia hemprichii (Ehrenb.) Aschers.
did not flower during eleven months at a range of temperatures, but flowered after transfer
from a higher temperature regime (31/27 ° C, day/night) to a lower one of 26/24°C (day/
night). Four other species from Kenya did not flower under any of the experimental
conditions. It is concluded that a wide range of environmental conditions is involved in
the reproductive physiology o f members of this tropical seagrass flora.

INTRODUCTION

Flowering has been observed rarely for some seagrass species of the Indo-
Pacific. For Cymodocea serrulata (R.Br.) Aschers. & Magnus, den Hartog
{1970) reported pistillate flowers from New Caledonia (April 1869) and
fruits from Thursday Island, Queensland, Australia (November 1967). Isaac
(1968) described a single pistillate flower that was collected in beach drift
at Diani Beach, Kenya (January 1967) and Kay {1971) described further
details of pistillate flowers at Mida Creek, Kenya (August 1969). Kirkman

0304-3770/80/0000--0000/$ 02.25 © 1980 Elsevier Scientific Publishing Company

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(1975) reported staminate flowers of C. serrulata for the first time from col-
lections in Moreton Bay, Queensland, Australia, in February 1973 and 1974.
For Halophila stipulacea (Forsk.) Aschers., den Hartog (1970) cited flowering
collections from Kenya (August 1965) and Mauritius (October 1929), both
pistillate. Isaac (1968) noted only pistillate plants at Diani Beach, Kenya.
Den Hartog (1972) described staminate flowers of H. stipulacea from Malta,
and Lipkin (1975) reported staminate flowers in the Red Sea (July 1965,
August 1973 and several collections in 1975. For Zostera capensis Setchell,
Isaac (1968) reported no flowering collections from Kenya and den Hartog
(1970) noted that he had not seen flowering material from the Tropics. One
inflorescence fragment of Z. capensis was collected at the base of plants at
Shimo la Tewa, Kenya, during the investigation reported here, in September
1978. Because records of flowering are relatively rare for the above three spe-
cies, the possibility of floral induction under experimental conditions did not
seem likely.
Flowering records of other Indo-Pacific species are not common (den Hat-
tog, 1970), but flowering specimens have been recorded with slightly greater
frequency. For Syringodium isoetifolium (Aschers.) Dandy, den Hartog (1970)
cites eight records of flowering and/or fruiting plants in the N o , h e m Hemi-
sphere and seven records in the Southern Hemisphere. For Thalassia hempri-
ch ii (Ehrenb .) den Hartog ( 1970) cites ten records of flowering and/or fruiting.

MA TER I ALS AND METHODS

Vegetative axes were washed free of most sediment at the time of collec-
tion on the coast of Kenya. The plants of Cymodocea serrulata were in a tight
sod with Thalassia hemprichii on a coral shelf at Diani Beach. The collection
of Halophila stipulacea was made in sandy tide pools with H. ovalis (R. Br.)
Hook. f. Other seagrasses collected in various intermixtures at Diani Beach in-
cluded Thalassodendron ciliatum (Forsk.) den Hartog, Halodule uninervis
(Forsk.) Aschers, narrow-leaf clones in sites with greater exposure to air at
low tide; broad-leaf clones in tide pools) and Cymodocea rotundata Ehrenb.
& Hempr. ex Aschers. Syringodium isoetifolium beds tended to be more or
less isolated from the other seagrasses and occurred in deeper sites that were
least likely to be exposed at low tide. The plants of Zostera capensis were
collected at Shimo la Tewa, 12 km north of Mombasa, in sandy soils at the
mangrove fringe near the mouth of an inlet. The material collected in mid-
September 1978 was transported moist in plastic bags inside polystyrene
chests to Texas.
For experimental studies, vegetative axes were planted in standard seagrass
cultures (McMillan, 1979a). These cultures consisted both of 500-ml cups of
fine sandy loam from central Texas and synthetic seawater (Instant Ocean).
Triplicate plantings of each of the nine species were placed in each of three
 325

Instant Ocean refrigerated systems. In addition, the cultures of Zostera

capensis were placed in non-aerated aquaria with Z. marina L. collections
from North America. Some cultures of Halodule uninervis and Syringodium
isoetifolium involved collections t h a t were found flowering in Kenya.
The Instant Ocean systems were kept in growth chambers and the tem-
p e r a t u r e - d a y length regimes were varied during the study. The higher (31/27 °
C, day/night) and lower (26/24°C, day/night) temperature programs operated
continuously over the 12 m o n t h s study period. A third system was kept in
a fixed sequence of temperatures: 26°C for seven months, 24°C for three
months and 28°C for two months. The three systems had the same day
length sequence: three months at 13 h, three m o n t h s at 12 h and for each
succeeding m o n t h , 13, 12.5, 12, 13 and 13 h. Light was provided by cool,
white fluorescent bulbs only and the photon-flux density varied within an
aquarium between 70 and 200 ~mol m -2 s -~ . Two separate Instant Ocean
systems were also used for Syringodium isoetifolium.
The composition of the artificial seawater was altered at intervals through-
out the study. Approximately one-third of the water was changed in each
106 1 aquarium at three-monthly intervals. Additional chelated iron (EDTA
FeSO4) was given at intervals. The salinity was initially at 350/00 but was
altered at intervals to 25 and 30% 0. The salinity was monitored with a Gold-
berg refractometer and tap water was added to compensate for evaporation.
A special seawater was prepared w i t h o u t certain trace materials for S. isoeti-
folium.

RESULTS

C y m o d o c e a serrulata

Flowering in C. serrulata showed the strong controlling influence of tempe-

rature (Table I). Flowers, all pistillate (Fig. 1), were confined to the higher
temperature program (31/27 ° C, day/night). The first flower appeared eight
months after transplanting to the experimental cultures, but flowers were in
anthesis continuously for three months. A m a x i m u m of five reproductive
shoots was produced at any one time by each of two triplicates; two flower-
ing shoots appeared at one time on a third replicate. A total of 23 flowers
was recorded during the three m o n t h period but only a single flower appeared
on a short shoot at a time. As Kay (1971) has reported, the flowers are not
terminal. No flowers were produced by replicates that were kept at lower
temperatures. Flowers were produced over the day length range of 12--13
h. During the period that flowers were produced continuously, day lengths
varied at intervals from 12.5 to 12 and 13 h, b u t no disruption in the pro-
duction of flowers was observed. The day length had been changed from 13
to 12.5 h two weeks before the first flowers appeared.
Salinity and nutrients m a y play a role in the flowering response of C. ser-
rulata. During the three m o n t h s of flowering, salinities were varied periodical-
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TABLE I

Flowering in seagrasses o f Kenya under diverse temperatures ( fl. indicates flowering, reg.
vegetative)

Species Temperature (° C)

18 24 26 26/24 28 31/27
(day/night) (day/night)

Cymodocea serrulata I -- reg. veg. reg. reg. fl.

HalophzTa stipulacea 2 -- fl. fl. ft. ft. ft.
Syringodium isoetifolium 3 -- ft. fl. veg. -- veg.
Zostera capensis 4 fl. fl. --
Thalassia hemprichii s -- reg. veg. fl. veg. veg.

i C y m o d o c e a showed an increase in flower production with decreasing salinity to 25%0.

Flowers (a total of 23) were produced during a three month testing at 31/27°C (day/night)
with day lengths that varied from 13 to 12 h and salinity from 35 to 250/0o .
2 I-[alophila showed a strong correlation of flowering with the nutrient status o f the water

column. Flowering was most readily induced at the lower temperature range, 24, 26 and
26/24 ° C, b u t was most reliably induced at all temperatures after a partial change of the sea-
water or an addition of chelated iron (EDTA, FeSO 4 ).
3 S y r i n g o d i u m did not flower during a seven month period at 26 ° C in regular seawater.
It did flower subsequently (three out of five replicates) at 26°C in seawater that lacked
certain micronutrients (Cu, Zn, I, Co, V) that were provided in a liquid trace medium by
the manufacturers of Instant Ocean. One of the four replicates flowered at 24°C in the
nutrient-deficient seawater.
* Z o s t e r a failed to survive initial transplanting to the higher temperature regime, but surviv-
ed at 16 ° C. Flowers appeared after the temperature was raised to 18 ° C. After flowering,
the triplicates were transferred to 22 and then 24°C. Near the end of the 12-month study,
one o f the triplicates flowered at 24 ° C.
s T h a l a s s i a flowered after plants were transferred from the 31/27 ° C regime to lower tem-
peratures. The replicates were at the 31/27°C program for 11 months, then at 28/24°C
for two weeks and then at 26/24°C for two weeks. During the last two months of the
study, the plants were continuously at 13 h day lengths and 35°/o0 salinity.

l y f r o m 3 5 t o 3 0 a n d t h e n 25°/00. T h e n u m b e r o f f l o w e r i n g s h o o t s i n c r e a s e d
after changes towards lower salinity, but there was no indication that an
a b r u p t c h a n g e b a c k t o 35°/oo f r o m 250/00 a f f e c t e d f l o w e r s t h a t w e r e in an-
thesis. Stigmas that had elongated before the increase of salinity remained
white and fresh.

Halophila stipulacea

Flowering of H. stipulacea occurred over ranges of temperature and day

length but also showed a strong correlation with the nutrient condition of the
water column. Initially, it was suspected that temperature was the main flo-
wering control, but later, an interaction of temperature and day length
seemed a more likely explanation. By using a series of experimental mani-
 327

Fig. 1. Pistillate flower o f Cymodocea serrulata in experimental culture at temperatures of

3 1 / 2 7 ° C (day/night). The 27 m m long stigmas e x t e n d 15 m m above the sheath.

pulations, as described below, the primary control appeared to be linked to

the nutrient condition of the water column.
The influence of temperature on flowering was indicated b y responses re-
corded after transplanting to experimental cultures (Table I). After one
month, pistillate flowers appeared on plants under the t w o lower temperature
regimes (26°C and 26/24 ° C, day/night) b u t n o t in the higher temperature
program (31/27°C, day/night). After six months in culture, however, flowers
(Fig. 2) were produced at each of the temperatures, including the higher
temperature regime. As flower production occurred over the entire range of
temperature it was suggested that some other environmental factor might
play a crucial role in controlling flowering in H. stipulacea.
An interaction with p h o t o p e r i o d was suspected because the plants under
the higher temperatures had flowered under 12 h b u t n o t under 13 h day
lengths. The day length had constantly been 13 h during the first three
months of the study and then at 12 h during the second three m o n t h period.
Because the flowers appeared at the higher temperature only during exposure
to 12 h day length, a possible relation was suggested in which flowers would
appear at lower temperatures over a wide range o f photoperiods, b u t that
flowers would be induced at higher temperatures only if the day length was
shortened. Alterations to the composition of the seawater one m o n t h before
the flowering at the higher temperature suggested that nutrient status might
be involved in flowering control. The reduction in flower production in cul-
tures kept at various temperatures w i t h o u t seawater enhancement also sug-
328

iiii!!iiiiiiiii!iliiiiiiiiiiiii
i ~
i! iii!i

Fig. 2. Pistillate flower of Halophila stipulacea in experimental culture at temperatures of

26/24° C (day/night).

gested possible interactions of nutrients and flowering. Plants kept under

the 26/24°C regime stopped flower production after approximately t w o
months, b u t resumed flower production u p o n the addition of chelated iron.
After abrupt changes in salinity, flower production was also curtailed.
The possible interaction o f temperature, day length and nutrient status w a s
tested on six replicates. The six plants had been kept at 26°C and 2 5 % 0 salini-
ty and had stopped floral production before being placed at 24°C in new
seawater prepared with de-ionized water with salinity of 35%o. Flower
production resumed after approximately t w o weeks in the new condition and
continued for t w o months, although the day length was changed at intervals
from 13 h to 12.5 and 12 and then back to 13 h. All six replicates were
producing flowers as the temperature was changed to 28°C and the day
length was kept at 13 h. Chelated iron was added to the cultures at the time
the temperature was raised. Flower production resumed in t w o of the re-
plicates after t w o weeks at the higher temperature and four replicates were
 329

flowering after one m o n t h under this environmental regime. The flowering

data suggest the major influence of a nutrient, possibly iron, in the water
column.
Although no staminate flowers were observed on plants in the cultures,
fruit development was noted in the above series at 24°C. At the time the
temperature was raised to 28°C, fruits were enlarging and they continued to
do so at the higher temperature. Fruit that was dissected contained develop-
ing ovules, suggesting possible apomictic development of seed. Development
of fruit was n o t observed in other experimental cultures at various tempera-
tures and salinities.

Syringodium isoetifolium

Flowering of S. isoetifolium was primarily controlled b y temperature,

but also involved a salinity--nutrient interaction. Flowers were neither pro-
duced under temperature programs of 31/27 and 26/24°C (day/night), nor 26
and 24 ° C during the first nine and a half months of the study, b u t flowers ap-
peared five weeks after plants were transferred to a special seawater medium at
26°C. A few flowers appeared in the special seawater medium at 24°C b u t
not during the 12 months in the regular seawater at 26/24 ° C.
The special seawater had been effective in promoting flowering of the
Gulf--Caribbean species, Syringodium filiforme Kutz. (McMillan, 1980a)
and also was involved in flower production by S. isoetifolium. The exact
nutrient interaction is n o t known, b u t the seawater was deficient in the trace
elements, Rb, I, A1, Zn, V, Co and Cu. These trace elements occur in a liquid
trace medium that was included along with dry salts in two-part Instant
Ocean mixture that was commercially available before 1977. EDTA and Fe
also appear in the liquid trace, b u t these were added to the dry salts. The dry
salts have an ionic composition including C1, Na, SO4, Mg, Ca, K, HCO3, Br,
H3BO3, Sr, SIO3, PO4, Mn, F, MOO4, $203 and Li (listed in decreasing con-
centration, according to Aquarium Systems, Inc., Ohio). Because the plants
were growing in fine, sandy loam, the elemental composition may have in-
cluded c o m p o u n d s that were omitted from the liquid trace medium. The
special seawater was prepared with de-ionized water to avoid possible nutrient
addition from tap water.
Salinity may have played a role in the flowering of S. isoetifolium. The
salinity was lowered from 35 to 30 % 0 during the first three weeks, and then
to 25%0 for four days. The first flowers appeared t w o weeks after the
salinity was returned to 35 %0.

Zostera capensis

Zostera capensis reacted differently from the other Kenya seagrasses, in

that plantings o f Zostera did n o t survive in the higher temperatures after
initial transplanting to these conditions. Triplicate plantings in each of the
330

three Instant Ocean systems, under 31/27, 26/24 and 26°C temperature re-
gimes slowly lost vigor and none was living after the first m o n t h of the
study.
A triplicate planting of Z. capensis that was cultured with Z. marina was
the only surviving series. The planting was in a non-aerated aquarium under
16°C and 12 h day length conditions. These triplicates were transferred near
the end of the study to 22 and 24°C and grew vigorously.
T w o inflorescences appeared on one of the three triplicates after seven
and a half months in experimental culture. The flowering occurred t w o
weeks after the day length was changed to 13 h and the temperature raised
to 18°C from 16°C, b u t it seems likely that the floral induction occurred
under 12 h and 16°C conditions. Near the conclusion of the 12-month study,
a single inflorescence appeared on one of the triplicates at 24°C and 13 h
day length (Fig. 3).

Fig. 3. I n f l o r e s c e n c e o f Zostera capensis p r o d u c e d in e x p e r i m e n t a l culture at 24 ° C.

Thalassia hemprichii

Thalassia did n o t flower during 11 months of the study under any of the
environmental manipulations, b u t rapidly produced flower buds near the end
of the 12-month study period, after a temperature change. Plants that had
been at 31/27°C (day/light) continuously for 11 months were transferred to
a program of 28/24°C for t w o weeks and then to 26/24°C for t w o weeks.
 331

Fig. 4. T w o inflorescences o f Syringodium isoetifolium in an experimental culture at 26°C.

Buds appeared on four short shoots in two of the three replicates following
this temperature shift. Other replicates t h a t had been under a range of tem-
peratures during the study, at 26, 24 and 28°C successively, did n o t produce
flowers, and replicates t h a t were continuously at 26/24°C did n o t show
floral induction. The flowering of replicates moved from the higher to the
lower temperature programs, both with 13 h day length, suggests that the
major inductive influence involves a lowering of temperature.

Other seagrass species

The other four species survived in the experimental cultures, but no

flowers were observed under any of the environmental manipulations. All
were vigorous during 12 m o n t h s in culture. Clones of Halodule uninervis
that differed in leaf width at the time of transplanting continued to show
morphological differences. The wider-leaved clones that were collected in
tide pools and were least likely to be exposed to air at low tide continued
to produce wide leaves; the narrower-leaved clones that were collected in
areas exposed to air at the lowest tides continued to produce narrow leaves.
The environmental conditions that were involved in the flowering of Cymodo.
cea serrulata and Halophila stipulacea did not induce flowers in Cymodocea
rotundata, Halophila ovalis or Thalassodendron ciliatum. Although the sea-
grasses of Kenya are naturally intermixed on the coral shelf at Diani Beach, a
wide range of environmental conditions must be involved in the reproductive
physiology of members of this tropical seagrass flora.
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Fig. 5. Staminate flower of Thalassia hemprichii in experimental culture at temperatures of

26/24° C (day/night).

In September, 1978, at the time of collection in Kenya, some flowering

was observed among species at Diani Beach and Shimo la Tewa (Table II).
Plants of Halodule uninervis (narrow-leaved) which were completely exposed
to air at low tide were flowering vigorously. Both staminate and pistillate
clones were transplanted to the experimental cultures: staminate flowers
continued to be produced in the cultures for approximately t w o weeks b u t
immature fruits that had been produced in K e n y a failed to mature in experi-
mental cultures. Thalassia hemprichii grew mature fruits over wide areas of
the collection sites and seedlings germinated immediately from collected
fruits. Most of the fruits were still attached to the plants b u t some fruits
were found in beach drift. A transplanted Thalassia plant produced a flower
in the experimental cultures during the first m o n t h of the study, undoubted-
ly resulting from buds that had been induced under natural conditions in
Kenya. Pistillate and staminate flowers of Syringodium isoetifolium were ob-
served in Kenya; flowering plants that were transplanted to experimental
conditions did n o t continue to produce flowers. As indicated above, one
fragment o f an inflorescence was found in a collection of Zostera capensis,
b u t the plants were n o t producing flowers at the time of collection.

DISCUSSION

Among the seagrasses transplanted from Kenya, flowering occurred in five

species during 12 months of experimental culture. Flowering in each of the
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TABLE II

Flowering records in Indian and western Pacific Ocean seagrass beds (fl. indicates flowering,
ft., fruiting)

Species Diani Beach, Kenya I Thursday Island, Qld., Australia 2

Month Flowering stages Month Flowering stages

Cy modocea serrulata January 9 ft. November fr. (dH)

C. rotundata September None observed November gQ ft. (dH); ~fl. (Mc)
Halodule uninervis
(wide-leaved form) August 9 ft. November 69f1., fr. (dH)
(narrow-leaved form) September 69 fl., fr. November 69 ft., fr. (dH,Mc)
Halophila ovalis
(large-leaved form) July 69 fl., fr. November 69 fl., fr. (Mc)
(small-leaved form) Not present November 6q fl., fr. (Mc)
H. stipulacea August 9 fl. Not present
Syringodium isoetifolium September d 9fl.
December 9 ft. November 69 fl., ft. (dH)
Thalassia hemprichii August ~ 9 fl. November None Observed (dH,
September fr. Mc)
Thalassodendron ciliatum August d9 ft. November fr. (dH)
Zostera spp. September None observed 4 November None observed (dH)

i The September records at Diani Beach (latitude 4 ° S) were taken in 1978. Other records
are for Isaac's collections (Isaac, 1968; den Hartog, 1970). Two collections axe included
from other Kenyan sites, Thalassodendron (Blue Lagoon), Halodule, wide-leaved form
(Gazi). Isaac's records were from 1965. except C. serrulata, from 1967.
2 Den Hartog (dH) visited Thursday Island (latitude 10 ° S) in November 1967 ; McMillan (Mc)
visited in November 1979.
3Zostera capensis was observed at Shimo la Tewa, Kenya; Z. capricorni Aschers. at Thursday
Island, Australia.
4 A fragment of an inflorescence was recovered from the base of the plants at Shimo la Tewa
but the phenological pattern is not known for Kenya.

five s p e c i e s i n v o l v e d a d i f f e r e n t set o f e n v i r o n m e n t a l c o n d i t i o n s a n d s u g g e s t e d
t h a t t h e r e m a y b e diverse c o n d i t i o n s , in situ, a f f e c t i n g t h e r e p r o d u c t i v e p h y s i o -
l o g y o f t h e n i n e s p e c i e s o f this t r o p i c a l seagrass f l o r a .
D e s p i t e t h e n e a r - e q u a t o r i a l p o s i t i o n o f t h e seagrass b e d s o f K e n y a , t h e i r
h a b i t a t i n v o l v e s a w i d e r a n g e o f e n v i r o n m e n t a l c o n d i t i o n s , l a r g e l y as a c o n -
s e q u e n c e o f d a i l y t i d e s t h a t r e a c h a m a x i m u m o f ca. 4 m . T h e r e are t w o t i d e
c y c l e s f o r e v e r y p e r i o d o f s o m e w h a t o v e r 2 4 h a n d Isaac a n d Isaac ( 1 9 6 8 )
h a v e r e p o r t e d t h a t t h e l o w e s t t i d e s are d u r i n g d a y l i g h t h o u r s f r o m O c t o b e r
t o m i d - M a r c h , b u t are d u r i n g t h e h o u r s o f d a r k n e s s f r o m M a y t o A u g u s t .
D u r i n g t h e last h a l f o f M a r c h , t h e first h a l f o f A p r i l a n d t h e w h o l e o f Sep-
t e m b e r , t h e d i f f e r e n c e s in d a y a n d n i g h t - t i m e l o w w a t e r - l e v e l s are n e a r l y
n o n - e x i s t e n t . D u r i n g a 2 4 - h p e r i o d in m i d - S e p t e m b e r w h e n t h e c o l l e c t i o n s
w e r e m a d e , t h e seagrass b e d s w e r e s u c c e s s i v e l y u n d e r 4 m o f s u r g i n g w a t e r o f
t h e I n d i a n O c e a n a n d n e a r t h e i r g r e a t e s t e x p o s u r e t o air in t h e h e a t o f m i d -
334

day. As a result of the tidal range, the entire set of habitat conditions showed
wide diurnal variation, and it is not, therefore, surprising to find that the nine
species respond to a variety of environmental conditions in the p r o m o t i o n
of flowering.
Flowering has been reported b y Isaac (1968) in seven of the nine species
(Table II). For Cymodocea serrulata, one flower was collected in beach drift
in January 1967, as indicated above, at Diani Beach. Flowers were collected
in August 1965, from plants ofHalophila ovalis, H. stipulacea (cited as H.
balfourii Solered.), Thalassodendron ciliatum and Thalassia hemprichii. Sy-
ringodium flowers were collected in December and flowers of the wide-
leaved form of Halodule uninervis were collected in August. Only pistillate
flowers were observed on S. isoetifolium, H. uninervis and Halophila stipula-
cea. Isaac (1968) did not report flowers on the narrow-leaved form of H. uni-
nervis, which she referred to H. wrightii Aschers. b u t which McMillan (1980b)
has shown to have the same isozymes as the broad-leaved form. No flowers
were reported on C. rotundata or Z. capensis. The collections cited in den
Hartog (1970) indicate that only three species, Halophila stipulacea, H. ovalis
and T. hemprichii, were flowering at the same time (August) and in the same
place (Diani Beach).
Although phenological observations on seagrasses in Kenya are n o t com-
plete, those at Diani Beach in 1965 (cited in den Hartog, 1970) show some
correlations with the events observed in 1978 (Table II). In 1965, flowers
were recorded for Thalassia hemprichii in August; in 1978, the widespread
production of mature fruits in September suggests a recurrent reproductive
pattern. Collections of the narrow-leaved form of Halodule uninervis made at
various times during 1965 (March, June, July, August and December) had no
flowers, b u t the mid~qeptember 1978 collection had abundant flowers, both
pistillate and staminate. The time of flowering in Thalassia and Halodule sug-
gests a correlation with physical events which are seasonal. Some seasonality
does seem to be present in the coastal waters of Kenya. Isaac and Isaac
(1968) noted that the density of the intertidal vegetation and the variety of
species (including algae) were least when the lowest spring tides occurred
during daylight hours and temperatures were high, near the end of October
to mid-March. In contrast, the variety of species was highest when day tem-
peratures were lower and the lowest tides were at night. Isaac (1968) noted
that, on a m u d flat at Lamu, northeast Kenya, Halodule uninervis and Halo-
phila ovalis were plentiful in July b u t were almost absent in March. Zostera
seemed to be less affected, and was reported to be in abundance at both
times of the year. The highest average m o n t h l y sea temperature (29.7 ° C)
was recorded in March and the lowest average temperature (24.8°C) was re-
corded in September.
The phenology of each tropical seagrass c o m m u n i t y probably correlates
with local environmental conditions. At Thursday Island, Queensland (Table
II), den Hartog (1970) recorded flowering and/or fruiting in mid-November
1967, for five species; Thalassodendron ciliatum, Cymodocea serrulata, C.
 335

rotundata, Syringodium isoetifolium and Halodule uninervis, but no flowering

for Halophila ovalis and Thalassia hemprichii. McMillan visited Thursday
Island in November 1979(see Table II) and recorded flowering in C. rotunda-
ta, H. uninervis and H. ovalis. R.C. Phillips (personal communication, 1979)
in April 1979, observed a few fruits and I. Johnstone (personal communica-
tion, 1979) in May 1979 observed a few flowers of Thalassia, in drift material
at Thursday Island. The comparison of flowering records for Thursday Island
with those for Diani Beach suggests that the phenology results from control
by local habitat.
The flowering of each tropical seagrass probably involves a particular tem-
perature, salinity and/or nutrient relationship under day lengths of 12--13
h. The experimental flowering of Halophila stipulacea suggested that tempe-
ratures of 24--26°C were most likely to instigate floral induction, but that
certain nutrient conditions, possibly involving iron availability, permitted
flower production at higher temperatures. The flowering of C. serrulata sug-
gested a relationship involving higher temperatures, (31--27 ° C) and lowered
salinities to 25%0. The flowering of S. isoetifolium in a deficient seawater
medium at 26°C suggested an interaction of nutrients and lower temperatures
in its reproductive physiology. The flowering of Z. capensis at 18°C, a tem-
perature lower than any recorded in Kenyan waters (Isaac and Isaac, 1968),
and subsequently at 24°C suggests that conditions related to low temperatures
may initiate flowering. The flowering of T. hemprichii after temperatures
were lowered suggests that seasonal lowering of temperature is probably the
inductive feature. The individuality of flowering requirements is further em-
phasized by the lack of flower production by four of the tropical species
from Kenya, under any of the experimental conditions.
Flowering in other seagrass species in experimental cultures is primarily
a consequence of temperature conditions, but also involves day length and
nutrient interactions. In Halophila engelmanni Aschers., flowers were induced
at 24/22 ° C(day/night) under day lengths ranging from 14--24 h (McMillan,
1976). Studies of Syringodium filiforme (McMillan, 1980a) have shown that
floral induction may occur at temperatures from 22--25°C under day lengths
ranging from 12--24 h. Flowering in S. filiforme was most likely in seawater
prepared without certain trace elements, as indicated above. In both H. en-
gelmanni and S. filiforme, floral induction was inhibited by day lengths
shorter than those indicated above. Experimental cultures of Cymodocea
rotundata, from the Palau Islands of Micronesia, flowered under the same
temperatures that were involved in the flowering of C. serrulata from Kenya
(McMillan, 1979b). The cultures of C. rotundata, however, were kept under
constant salinities near 35%0, but those of C. serrulata in the present study
were subjected to salinities varying from 35 to 250/00. Because flowering in
the experimental cultures has involved artificial seawater and non-marine
planting media, it is suggested that the reproductive physiology of most sea-
grasses could be established through experimental manipulations, and that
details of the nutrient effects can be determined.
336

ACKNOWLEDGEMENTS

This r e s e a r c h w a s f u n d e d b y N a t i o n a l S c i e n c e F o u n d a t i o n G r a n t O C E 77-
2 6 3 9 9 . I t h a n k S. D o s s a j i f o r his a d v i c e a n d e n c o u r a g e m e n t in K e n y a a n d J . R
W i g i n t o n a n d S.C. W i l l i a m s f o r t h e i r t e c h n i c a l a s s i s t a n c e w i t h t h e e x p e r i m e n t -
al c u l t u r e s .

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