Taiwan J For Sci 27(1): 1-11, 2012 1

Research paper

Seed Germination and Storage of

Cycas taitungensis (Cycadaceae)
Ching-Te Chien,1,2) Shun-Ying Chen,1) Shu-Hwa Chang,1) Jeng-Der Chung1)

ȪSummaryȫ
Cycas taitungensis is endemic to an area of about 290 ha in Taitung County, eastern Taiwan.
However, this species has been attacked by a scale insect (Aulacaspis yasumatsui) and larvae of the
butterfly, cycad blue (Chilades pandara peripatria), and many plants have died in the last 20 yr. To
establish ex situ conservation of C. taitungensis, seeds were collected from a managed plantation
and the Cycas taitungensis Nature Reserve for germination. Some of the seeds from the plantation
were stored at 5ʨ for 8 and 16 mo and germinated at 30/20ʨ to test germinability. The embryo
length was measured during seed incubation at 25/15 and 30/20ʨ. Germination of seeds from
2007 from the plantation began in weeks 17~18, and final germination was about 58~60L at 30/20,
25/15, and 25ʨ incubation. Germination of seeds from 2008 from the same plantation began in
week 18, and final germination percentages were 13, 64, 75, and 84L at 20/10, 25/15, 30/20, and
25ʨ incubation temperatures, respectively. Thus, favorable germination temperatures were 30/20
and 25ʨ. Initial germination of seeds from 7 individual plants from the Nature Reserve extended
from weeks 3 to 18, and final germination ranged 30~88L. These differences may have been due
to the wide range in time of pollination in the Nature Reserve, thus resulting in different degrees of
seed maturity. Fresh seeds air-dried for 24 h and stored at 5ʨ for 16 mo retained their original ger-
mination, whereas seeds stored with moist moss at 5ʨ for 16 mo showed decreased germination
from 73L (fresh seeds) to 50L. Cold storage at 5ʨ shortened the time to begin seed germination
regardless of the storage conditions with or without moist moss. The underdeveloped embryo of C.
taitungensis seeds must increase in length by about 117L before seed germination can occur. We
concluded that seeds of C. taitungensis exhibit morphophysiological dormancy, a high temperature
at 30/20 or 25ʨ increases seed germination, and air-dried seeds can be stored at 5ʨ for at least 16
mo.
Key words: Cycas taitungensis, morphophysiological dormancy, seed germination, seed storage, un-
derdeveloped embryo.
Chien CT, Chen SY, Chang SH, Chung JD. 2012. Seed germination and storage of Cycas taitun-
gensis (Cycadaceae). Taiwan J For Sci 27(1):1-11.

1)
Silviculture Division, Taiwan Forestry Research Institute, 53 Nanhai Rd., Taipei 10066, Taiwan. ‫ݔ‬
ཿၑᡜ‫ݔيܛ‬ಣȂ10066ѯіҀࠓ੖ၰ53ဵȄ
2)
Corresponding author, e-mail:chien@tfri.gov.tw ೾ଊձ޲Ȅ
Received July 2011, Accepted November 2011. 2011Ԓ7Уଛቸ 2011Ԓ11У೾ႇȄ
2 Chien et al.˕Germination and storage of Cycas seeds

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២ӱ᎐‫ڨ‬ҪᎉࣾϮ෧ᙬ‫ݎڸ‬ହង២ϊԹፇ҃ᙬ࣐ড়ԬκȄ࣐࡛Ҵѯ‫ݎ‬ង២ୣѵߴ‫ي‬Ȃ௵Ս‫ݔ‬ଡ଼‫׌‬೪ဋ
‫ޠ‬ѯ‫ݎ‬ង២Սดߴ੽ୣ‫ڸ‬ѯ‫ݎ‬τ‫ݢ‬໑Κ೏‫ؾ‬΢စᕋ‫ޠ‬ѯ‫ݎ‬ង២਼ᆎୣ‫ޠ‬ᆎφȂϸրໍ՘ϛӤྤ࡚‫ޠ‬ᆎ
φึ߄ၑᡜȄѫѵȂ௵Ս‫ؾ‬΢਼ᆎୣ‫ޠ‬ᆎφഌӌܼ5ʨᓾᙡ8ঐУ‫ڸ‬16ঐУȂ‫ڦ‬яࡤӶ30/20ʨίึ߄Ȃ
пᕤ၍ѯ‫ݎ‬ង២ᆎφᓾᙡࡤϟึ߄૗ΩȄᆎφथߞ࡚໕ขҼӶᆎφክᆎࡤӤਣໍ՘Ȅ2007Ԓ௵Ս‫ؾ‬
΢਼ᆎୣᆎφ໡ۗึ߄ਣ໣࢑Ӷಒ17~18໋Ȃึ߄౦Ӷ30/20ȃ25/15‫ڸ‬25ʨίႁ58~60 L ȇ௵ՍӤӵୣ
2008Ԓᆎφ໡ۗึ߄ਣ໣࢑Ӷಒ18໋Ȃึ߄౦Ӷ25ʨίഷାႁ84 L ȂӶ30/20ȃ25/15‫ڸ‬20/10ʨίϸր
࣐75ȃ64‫ڸ‬13 L ȄӱԫȂᎍӬѯ‫ݎ‬ង២ᆎφึ߄ྤ࡚࣐30/20‫ڸ‬25ʨȄ௵Սѯ‫ݎ‬ង២Սดߴ੽ୣ7ੂѯ
‫ݎ‬ង២ᆎφȂ໡ۗึ߄ਣ໣௄ಒ3໋Վಒ18໋ϛ๊Ȃйึ߄౦௄30 L Վ88 L ϛ๊Ȃ೼‫ٳ‬ৰ౵Ѡ૗࢑ߴ੽
ୣҕੂ໣‫ڨ‬લਣ໣ϛΚȂᏳयᆎφԚዤ࡚ϛӤ‫ܛ‬Жକ‫ޠ‬Ȅѯ‫ݎ‬ង២ᆎφစࡊϲޫ੊ୂᕎ24ϊਣȂܼ5ʨ
ᓾᙡ16ঐУࡤϬ૗ߴࡼ঩ԥུᘁᆎφ‫߄ึޠ‬౦ȂկᆎφᇅᕇЬीెӬܼ5ʨᓾᙡ16ঐУȂึ߄౦௄ུᘁ
ᆎφ73 L ७Վ50 L ȂᡘұᆎφୂᙡЩᕇᙡ‫ٺ‬Ȅୂȃᕇᆎφᓾᙡܼմྤ5ʨࣲ૗ᕼ฼໡ۗึ߄‫ޠ‬ਣ໣Ȅѯ
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ԥ‫ם‬ᄙҢ౪‫ޠ‬ӅએȂାྤ30/20ʨ‫ڸ‬25ʨቩђᆎφึ߄౦Ȃᆎφജୂ24ϊਣࡤѠᓾᙡ5ʨՎЎ16ঐУȄ
ᜱᗥມȈѯ‫ݎ‬ង២ȃ‫ם‬ᄙҢ౪‫ޠ‬Ӆએȃᆎφึ߄ȃᆎφᓾᙡȃึ‫ي‬ϛ‫׈‬ӓथȄ
ᙐኋ኉ȃച๴ऽȃ஼ు๽ȃᗪਐ኉Ȅ2012Ȅѯ‫ݎ‬ង២ᆎφึ߄ᇅᓾᙡȄѯᢋ‫ݔ‬ཿऌᏱ27(1):1-11Ȅ

INTRODUCTION
Cycas taitungensis Shen, Hill, Tsou &
Chen, previously misidentified as C. taiwani-
ana, is endemic to a specific area of about
290 ha in Taitung County, eastern Taiwan
(22°51’30”~22°52’30”N, 120°57’30”~
121°01’00”E) (Shen et al. 1994). The Cycas
taitungensis Nature Reserve was established
by the Forestry Bureau, Council of Agricul-
ture, Executive Yuan, Taiwan in 1970 and
announced in accordance with the Cultural
Heritage Preservation Act of Taiwan in 1986.
The flora in the Cycas taitungensis Nature
Reserve was studied, and 293 species were
found with the 5 largest families being the Fa-
baceae, Asteraceae, Poaceae, Lauraceae, and
Euphorbiaceae (Tzeng et al. 2005). The plant
communities in the Nature Reserve were clas-
sified into 9 types by a matrix cluster analysis
and the dominant tree species were Cyclo-
balanopsis glauca, Zelkova serrata, Liq-
uidambar formosana, Fraxinus formosana,
Cinnamomum philippinense, Lagerstroemia
subcostata, etc. (Tzeng et al. 2005). Cycas
taitungensis has been attacked by cycad scale
insects (Aulacaspis yasumatsui Takagi) and
larvae of the butterfly, cycad blue (Chilades
pandara peripatria), and many Cycad plants
have died in the last 20 yr (Hsu 1989, Lan
1998, Bailey et al. 2010). Investigations in
the Nature Reserve showed that many scale
Taiwan J For Sci 27(1): 1-11, 2012
insects were found year-round on leaves,
twigs, and rachides, but larvae of cycad
blue were found in spring when new leaves
occurred.
Depending on the species, germination
of Cycas seeds can extend for 4~12 mo after
sowing, and there is no need to scarify the
hard seed coat (Broome 2011). Field seed
collection of several Australian Cycas spe-
cies germinated with no treatment, and they
can germinate only after the embryo is fully
developed (Dehgan and Schutzman 1989).
Thus, one objective of our study was to deter-
mine the best temperature to germinate seeds
of C. taitungensis.
Cycas seeds generally are large, the
weight per seed is 6.3~13.7 g (Zheng 2000),
and seeds cannot be dried when they are
stored, otherwise germination decreases (De-
hgan 1999, Broome 2011). The optimum stor-
age conditions to extend Cycas seed longevity
and reduce storage costs must be determined.
Thus, our second objective was to find the
best way to store the seeds for more than 1 yr
without loss of germinability.
A seed that does not have the capacity to
germinate in 30 d under suitable environmen-
tal conditions such as temperature, light, and
water is termed dormant (Baskin and Baskin
1998, 2004). Five classes of seed dormancy
are recognized, i.e., morphological dormancy,
physiological dormancy, morphophysiologi-
cal dormancy, physical dormancy, and com-
bined dormancy (physical + physiological
dormancy) (Baskin and Baskin 2004). Seeds
exhibiting both morphological and morpho-
physiological dormancy have underdeveloped
embryos that must grow inside the seeds
before the radicle emerges. Thus, our third
objective was to determine if the seeds of C.
taitungensis exhibit morphological or mor-
phophysiological dormancy.
Approximately 82L of Cycad species
3
in the world are threatened, and some of
them are in danger of extinction (Donaldson
2003). This paper provides information on
the storage of Cycad seeds and propagation
of the taxon from seeds for plantation use to
increase populations in the future.
MATERIALS AND METHODS
Seed collection and handling
Mature reddish-orange to vermilion
seeds of C. taitungensis were collected both
from the Cycas taitungensis Nature Reserve,
Taitung County on 19 December 2007 and
from a managed plantation in Tawu (22°22’N,
120°54’E), Taitung County, Taiwan on 8~10
December 2007 and 2008 (Fig. 1). Seeds from
the Nature Reserve were collected and indi-
vidually numbered for the germination test.
Seeds from > 20 plants of the the managed
plantation each year were mixed as a seed lot
for the germination test. The seed consists of
an embryo surrounded by a large megagame-
tophyte, a stony sclerotesta, and a fleshy outer
sarcotesta. Seeds without the fleshy sarcotesta
collected from managed plantation in 2008
were 49.79ɲ2.55 mm long, 32.32ɲ1.69 mm
wide, and 24.08ɲ1.17 mm thick (n = 20) (Fig.
2). There are 35 seeds L-1 and 52 seeds kg-1.
Due to scale insects and fungi present
on the seed surfaces, all seeds were treated
with 2000 ppm Benomyl (a fungicide and
pesticide) for 1 h and air-dried in the labora-
tory for 24 h before using them in subsequent
experiments.
Effect of temperature regimes on seed
germination
Fresh seeds with dried sarcotesta collect-
ed from the managed plantation in 2007 and
2008 and mixed with moist sphagnum moss
(cut into small pieces) were placed in sealable
polyethylene (PE) bags and incubated at 12
4 Chien et al.˕Germination and storage of Cycas seeds

A B

C D

Fig. 1. Cycas taitungensis in a managed plantation from which old leaves are pruned in
winter (A), in the Cycas taitungensis Nature Reserve (B) with many cycad scale insects on
the leaves, twigs, and rachides (C), and a cycad blue butterfly laying eggs on growing leaves
(D).

A B

C D

Fig. 2. Intact seed of Cycas taitungensis with a reddish-orange sarcotesta (A), longitudinal
section of a fresh seed with an underdeveloped embryo (B), seed with a fully grown embryo
before root emergence (C), and a seed after radicle emergence (D). The scale marks are
millimeters.
Taiwan J For Sci 27(1): 1-11, 2012
h/12 h of alternating temperature regimes of
30/20, 25/15, and 20/10ʨ and at a constant
temperature of 25ʨ. Likewise, fresh seeds
collected from the Nature Reserve in 2007
and mixed with moist sphagnum moss in
PE bags were tested at 30/20ʨ for germi-
nation. The water content of the sphagnum
moss inside the PE bags was about 400L
of the dry mass. The daily photoperiod of
12 h in the incubators (60~80 μmole m-2 s-1,
400~700 nm) was at the high temperature.
Each treatment consisted of 3 replications of
25 seeds each, except seeds from the Nature
Reserve which consisted of 2 replications of
10~28 seeds each depending on the collection
number. The outer sarcotesta was removed
after several weeks of incubation. Germina-
tion, i.e., a radicle at least 2 mm long, was
recorded weekly for 32~60 wk. Results are
expressed as a germination percentage (L).
Due to the coarse nature of the sphagnum,
most seeds received some light, but at any
given point in time a few may have been in
darkness. However, at weekly intervals the
contents of each bag were poured out on a
table to facilitate examination of seeds for
germination. After germination was moni-
tored, seeds and the sphagnum moss were
returned to the bag, resulting in a re-shuffling
of seeds with regard to their position in the
sphagnum and thus the light they received.
Consequently, all seeds were in light part
(or all) of the time the lights were on in the
incubator.
Embryo growth measurement
To monitor embryo growth, seeds of C.
taitungensis collected from the managed plan-
tation in 2007 with the sarcotesta removed
were mixed with moist sphagnum moss,
placed in sealable PE bags, and incubated
at 30/20 and 25/15ʨ for 8 mo. Incubation
conditions were the same as these described
5
above for the germination test. Growth of em-
bryos was measured at 4-wk intervals. Seeds
were sectioned longitudinally using a mini
electric saw, and lengths of 5 embryos from
the above 2 incubation temperatures were
measured.
Effect of 5ʨ storage on seed germination
Fresh seeds (with the sarcotesta intact)
collected from the managed plantation in
2008 were treated with Benomyl solution
and air-dried in the laboratory for 24 h (as
described above) and then separated into 2
portions: 1 portion of seeds mixed with moist
sphagnum moss was placed in sealable PE
bags and stored at 5ʨ in darkness for 8 and
16 mo. The moist moss-stored seeds at 5ʨ
were opened each month to allow the seeds to
breathe. The other portion of seeds in sealed
PE bags was directly stored at 5ʨ for 8 and
16 mo without moist sphagnum moss. These
stored seeds were incubated at 30/20ʨ for a
germination test. Water contents in the sclero-
testa and megagametophyte + embryo of the
fresh seeds were 12.9 and 43.5L on a fresh-
weight (FW) basis, respectively, as deter-
mined by oven drying for 17 h at 103ʨ (ISTA
1999).
Statistical analysis
Means (3 replicates) and standard errors
of the germination percentage were calculated
based on treated seeds. Means of the percent
germination were compared by an analysis of
variance (ANOVA) and the least significance
difference (LSD) test at the 0.05 level of sig-
nificance (SAS Institute, Cary, NC). Percent-
age data were arcsine square-root-transformed
before analysis, but only non-transformed
data are shown in the figures. Embryo length
data based on 5 seeds of an incubation time
at 30/20 and 25/15ʨ were measured, and
means and standard errors were calculated.
6 Chien et al.˕Germination and storage of Cycas seeds

RESULTS 5). For example, seeds from a plant began to

Effect of temperature regimes on seed
germination
Germination of seeds collected from the
managed plantation in 2007 began in weeks
17~18, and final germination was 58~60L
after incubation at 30/20, 25/15, and 25ʨ
for 56 wk (Fig. 3). However, the germination
rate was lower at 25/15ʨ than at 30/20 and
25ʨ. Seeds from the same plantation in 2008
incubated at 30/20, 25/15, 20/10, and 25ʨ
for 59 wk germinated to 75, 64, 13, and 84L,
respectively (Fig. 4). The optimum tempera-
tures for seed germination were 30/20 and
25ʨ.
Seeds collected from 7 plants of the
Cycas taitungensis Nature Reserve in 2007
germinated to 30~88L, and the time to begin-
ning of germination was in weeks 3~18 (Fig.
germinate in week 3, and seeds of other plants
began to germinate in weeks 7, 10, and 18.
Embryo growth measurement
The mean (ɲS.E.) length of embryos
of fresh seeds collected from the managed
plantation in 2008 was 8.50ɲ0.45 mm, and
the critical embryo length just before the seed
germinated was 18.42ɲ1.06 mm (Figs. 2,
6). Thus, embryo length increased 117L be-
fore seeds germinated. After 16 wk, embryo
growth in seeds was faster at 30/20ʨ than at
25/15ʨ (Fig. 6).
Effect of 5ʨ storage on seed germination
Germination of air-dried seeds stored at
5ʨ for 8 and 16 mo retained the same per-
centages compared to seeds without storage,
but those mixed with moist sphagnum at 5ʨ

Fig. 3. Effect of temperature on the germination of Cycas taitungensis seeds collected from
a managed plantation in 2007. Seeds mixed with sphagnum moss were incubated at 30/20,
25/15 and 25ʨ for germination. Vertical bars areɲthe standard error.
Taiwan J For Sci 27(1): 1-11, 2012 7

Fig. 4. Effect of temperature on the germination of Cycas taitungensis seeds collected from
a managed plantation in 2008. Seeds mixed with sphagnum moss were incubated at 30/20,
25/15, 20/10 and 25ʨ for germination. Vertical bars areɲthe standard error. Points with
different letters significantly differ at 59 wk of incubation (LSD, p = 0.05).

Fig. 5. Cumulative germination percentages of Cycas taitungensis seeds collected from the
Nature Reserve in 2007. Seeds mixed with sphagnum moss were incubated at 30/20ʨ for
germination. Each number represents a single female plant.
8 Chien et al.˕Germination and storage of Cycas seeds

Fig. 6. Effect of temperature on embryo growth of Cycas taitungensis seeds collected from a
managed plantation in 2007. Seeds mixed with sphagnum moss were incubated at 30/20 and
25/15ʨ, respectively, and embryo length was measured at 4-wk intervals. Vertical bars are
ɲthe standard error.

decreased by about 18L after 8 and 16 mo of DISCUSSION

storage (Fig. 7). Final germination percent-
ages between dry-stored seeds and moist-
sphagnum-stored seeds significantly differed
after 50 wk of incubation, but the final germi-
nation percentages of seeds stored for 8 and
16 mo within the same storage treatment did
not significantly differ. Storage shortened the
time to initiate seed germination from week
20 of fresh seeds to week 14 of 16-mo air-
dried stored seeds, or to week 10 of 16-mo
moist-sphagnum-stored seeds (Fig. 7). After
16 mo of storage, water contents in the sclero-
testa and megagametophyte + embryo of the
moist-sphagnum-stored seeds were 28.9 and
45.1L, respectively; water contents in the
sclerotesta and megagametophyte + embryo
of dry-stored seeds were 14.9 and 44.5L, re-
spectively.
The optimal temperatures for maximum
germination percentage and rate in seeds of
C. taitungensis were 30/20 and 25ʨ. Seeds
from the managed plantation began to germi-
nate at these 2 temperature regimes in weeks
17~18, and final germination ranged 58~60L
in 2007 and 75~84L in 2008, whereas seeds
from 7 individual plants at the Nature Reserve
began to germinate at 30/20ʨ from weeks 3
to 18, and final germination ranged 30~88L.
These differences in seed germination from
the Nature Reserve may have been due to the
wide range in times of pollination and thus to
different degrees of seed maturity and embryo
sizes (5~22 mm in length) of seeds. In the
managed Cycas plantation, old leaves are cut
off after seeds are harvested and before new
Taiwan J For Sci 27(1): 1-11, 2012 9

Fig. 7. Effect of storage at 5ʨ on germination of Cycas taitungensis seeds collected from a

managed plantation in 2008. Stored seeds mixed with sphagnum moss were incubated at
30/20ʨ for germination. Vertical bars areɲthe standard error. Points with different letters
significantly differ at 50 wk of incubation (LSD, p = 0.05).

buds emerge in January to reduce the amount
of scale insects and avoid cycad blue laying
eggs. Thus, C. taitungensis plants in the man-
aged plantation flower at the same time, and
seeds ripen in October through November and
are dispersed in December.
Fresh seeds of C. taitungensis have a dif-
ferentiated (root and shoot can be observed)
embryo, and the length of the small underde-
veloped embryo had increased about 1.2-fold
inside the mature seed before the root and
shoot emerge (Figs. 2, 6). Therefore, Cycas
seeds exhibit morphological dormancy. Since
the seeds also required > 30 d for germina-
tion, we concluded that seeds exhibit both
morphological and physiological dormancy,
i.e., morphophysiological dormancy. Howev-
er, Cycas seeds do not exhibit epicotyl mor-
phophysiological dormancy because shoots
emerged in 2 wk.
There are 9 levels of morphophysiologi-
cal dormancy, i.e., non-deep simple, interme-
diate simple, deep simple, non-deep simple
epicotyl, deep simple epicotyl, deep simple
double, non-deep complex, intermediate com-
plex, and deep complex (Baskin and Baskin
2004, Baskin et al. 2008, 2009). Which one of
the 9 levels of morphophysiological dorman-
cy do seeds of C. taitungensis exhibit? Based
on the temperature requirements for embryo
growth and breaking physiological dormancy,
seeds of C. taitungensis do not require cold
stratification for germination, and embryo
growth occurs at warm temperatures, leading
10
to the conclusion that the seeds exhibit non-
deep simple morphophysiological dormancy.
Seeds without moist sphagnum stored
dry at 5ʨ for 16 mo retained their high ger-
minability, and the final germination percent-
age was nearly the same as those of fresh
seeds. Meanwhile, dry-stored seeds at 5ʨ
germinated rapidly, and the time to initiate
germination was shortened. However, seeds
mixed with moist sphagnum stored at 5ʨ
showed a decreased germination percentage,
although the time to initiate seed germination
was also shortened, indicating that these seeds
may have become too wet during cold storage
with moist sphagnum. We actually measured
the water content of seeds after 16 mo of cold
storage and found that the water content in the
sclerotesta had increased from 12.9L of fresh
seeds to 28.9L of seeds mixed with moist
sphagnum, but the water contents of the mega-
gametophyte + embryo were similar. Thus, we
recommend that Cycas seeds be air-dried for
24 h after harvest and sterilized, placed in seal-
able PE bags, and stored at 5ʨ. If water drops
in inner PE bag are found during storage, these
seeds need to be air-dried again overnight.
Seeds are mainly divided into 2 catego-
ries on the basis of storage behavior: ortho-
dox and recalcitrant (Roberts 1973, Hong
and Ellis 1996). Orthodox seeds have a low
water content following maturation, and can
be dried to 5L and stored at subzero tempera-
tures for long periods of time without a loss
of viability, whereas recalcitrant seeds shed
their high water contents and lose viability
rapidly if they are dried to a water content of
< 20~30L. Cycas seeds were described as ex-
hibiting recalcitrant storage behavior because
seeds cannot be dehydrated to a water content
below a critical minimum (Dehgan 1999,
Broome 2011). Dehgan and Yuen (1983) also
indicated that a complete separation between
the megagametophyte and sclerotesta in
Chien et al.˕Germination and storage of Cycas seeds
seeds (which produces a rattling sound when
shaken) was recognized as indicating that the
seeds are inviable. In the present study, we
determined that wet storage of seeds of C.
taitungensis can extend seed longevity, but
further study is needed to find the optimal
seed storage conditions.
Seed dispersal of C. taitungensis occurs
in winter. However, newly dispersed seeds re-
quire higher temperatures, e.g., 30/20 or 25ʨ
for germination. We speculated that dispersed
seeds may germinate in summer or fall if suit-
able environmental factors such as water and
light are available. To propagate C. taitungen-
sis plants, seeds can be air-dried and stored
at 5ʨ for several months until the next warm
season. Under this air-dried storage condition,
seeds can be stored for at least 16 mo, which
not only extends the storage time but also
increases the germination rate (by decreasing
the incubation time).
ACKNOWLEDGEMENTS
The authors thank Yen-Wei Chang,
Chang-Yen Chen, Wen-Yu Hsu, Ta-Yuan
Chien, and Cheng-Yu Yeh, Taiwan Forestry
Research Institute, for technical assistance.
We also thank personnel from the Taiwan
Forestry Bureau, Council of Agriculture,
Executive Yuan, Taiwan for helping with
seed collection of Cycas taitungensis. This
research was supported by 2 grants (96AS-
11.2.3-F1-e4 and 97AS-11.2.4-F1-G1) from
the Council of Agriculture, Executive Yuan,
Taiwan. The paper is dedicated to our good
colleague the late Dr. Yu-Pin Cheng for
partaking in the C. taitungensis program. A
portion of these data was presented at Seed
Ecology III, the Third International Society
for Seed Science Meeting on Seeds and the
Environment, which was held at Salt Lake
City, UT on 20~24 June 2010.
Taiwan J For Sci 27(1): 1-11, 2012
LITERATURE CITED
Bailey R, Chang NT, Lai PY, Hsu TC. 2010.
Life table of cycad, Aulacaspis yasumatsui
(Hemiptera: Diaspididae), reared on Cycas in
Taiwan. J Asia-Pacific Entomol 13:183-7.
Baskin CC, Baskin JM. 1998. Seeds: ecol-
ogy, biogeography, and evolution of dormancy
and germination. San Diego, CA: Academic
Press. 666 p.
Baskin CC, Chien CT, Chen SY, Baskin JM.
2008. Germination of Viburnum odoratissi-
mum seeds: a new level of morphophysiologi-
cal dormancy. Seed Sci Res 18:179-84.
Baskin CC, Chien CT, Chen SY, Baskin JM.
2009. Epicotyl morphophysiological dormancy
in seeds of Daphniphyllum glaucescens, a
woody member of the Saxifragales. Int J Plant
Sci 170:174-81.
Baskin JM, Baskin CC. 2004. A classifica-
tion system for seed dormancy. Seed Sci Res
14:1-16.
Broome T. 2011. Optimizing cycad seed ger-
mination. Available at: http://cycadjungle.8m.
com. Accessed 12 March 2011.
Dehgan B. 1999. Propagation and culture of
cycads: a practical approach. Acta Hortic 486:
123-31.
Dehgan B, Schutzman B. 1989. Embryo de-
velopment and germination of Cycas seeds. J
Am Soc Hort Sci 114(1):125-9.
Dehgan B, Yuen CKKH. 1983. Seed mor-
phology in relation to dispersal, evolution, and
propagation of Cycas L. Bot Gaz 144:412-8.
Donaldson JS. 2003. Cycads: status survey
and conservation action plan. IUCN/SSC Cy-
11
cad Specialist Group. Gland, Switzerland and
Cambridge, UK: IUCN. 86 p.
Hong TD, Ellis RH. 1996. A protocol to de-
termine seed storage behaviour. Rome, Italy:
International Plant Genetic Resources Institute.
62 p.
Hsu YF. 1989. Systematic position and de-
scription of Chilades peripatria sp. nov. (Lepi-
doptera: Lycaenidae). Bull Inst Zool Acad Sin
28:55-62.
ISTA (International Seed Testing Associa-
tion). 1999. International rules for seed testing.
Seed Sci Technol 27 (Suppl):1-333.
Lan PL. 1998. The population biology of Chi-
lades pandava peripatria (Lepidoptera: Lycae-
nidae): the fluctuation of population. Master’s
thesis. Taipei, Taiwan: National Taiwan Nor-
mal Univ. 60 p.
Roberts EH. 1973. Predicting the storage life
of seeds. Seed Sci Technol 1:499-514.
Shen CF, Hill KD, Tsou CH, Chen CJ. 1994.
Cycas taitungensis C. F. Shen, K. D. Hill, C.
H. Tsou & C. J. Chen, sp. nov. (Cycadaceae),
a new name for the widely known cycad spe-
cies endemic in Taiwan. Bot Bull Acad Sinica
35:133-40.
Tzeng HY, Chiu CA, Sheu JK, Wang CC,
Ou CH, Lu KC. 2005. Study on the flora of
Cycas taitungensis Nature Reserve, Taiwan. Q
J For Res 27(4):1-22.
Zheng YS. 2000. Cycas L. (Cycadaceae). In:
the National Service Center for State-Owned
Forest Farms and Forest Seed and Seedling Af-
fairs of the Forestry Ministry, editor. Seeds of
woody plants in China. Beijing, China: China
Forestry Publishing House. p 1-4. [in Chinese].
 1

A Simple and Efficient Method of Germinating Cycad Seeds©

Bart Schutzman

Environmental Horticulture Department, University of Florida, Gainesville, Florida 32611-0670,

USA

Email: bart@ufl.edu

INTRODUCTION

Cycads, an endangered group of plants from the world’s tropics and subtropics, have been a

mysterious and intriguing plant group to botanists since they were first documented more than

200 years ago. The number of described species continues to grow as subtropical and tropical

regions are thoroughly explored; the latest count published in the World List of Cycads (q.v.) is

343. Interest in these plants has grown tremendously over the last 20 years, especially since

accurate information has become readily available on the internet. The World List of Cycads, the

Cycad Pages, the Cycad Society’s Web site, and a number of other groups readily share

information and photographs.

Many species of cycads are endangered, and both plants and seeds can be both difficult and

expensive to obtain. The seeds of several species can be difficult to germinate and keep alive.

The purpose of this paper is to explain and recommend the “baggie method” of germination, a

technique that already is well-known in palms. It is not a new method for cycads by any means,

but too many people are still unfamiliar with its ease and benefits. The method increases

germination percentage and survivability of scarce and expensive seed. The information is

especially useful to both the nursery industry and hobbyists; it will ultimately reduce the pressure

exerted by poaching on indigenous cycad populations by making plants of the species easier to
 2

obtain. Indirectly, greater availability and ease of germination will reduce cost per plant, making

cycad species readily available to those who wish to grow them.

Status of Wild Cycads. Unfortunately, at the same time as we continue to document new cycad

species, habitat destruction and poaching continue to exact a heavy toll on wild cycad

populations. Many species may become extinct in the wild. This is not new knowledge, with

notable figures such as the late Cynthia Giddy, working as tireless advocates for the protection of

cycad habitats in the 1960s. The IUCN Red List of Threatened Species can be accessed at

www.iucnredlist.org, and detailed information on each threatened to endangered species can be

found. Unfortunately, lack of knowledge about cycads has led to some imprudent regulations

prohibiting seed collection from the wild. Very few seed produced by cycads in the wild result in

mature, fertile offspring. Making allowances for collection of some seed from wild populations

would dramatically increase the number of living plants of a given species, and reduce pressure

on wild populations. Ironically, the prohibition of wild seed collection has increased the amount

of poaching and resulted in some species becoming more endangered, since it is almost

impossible to protect every endangered cycad population in the wild from poaching. In fact, the

IUCN Red List documents four Encephalartos species that are now extinct in the wild due to

poaching.

Seed Germination. While cultivating cycad species out of habitat is of limited use in preventing

extinction, it can be of great utility in making many species available to those who might

otherwise traffic in illegal collected plants. There are enough privately and publicly held

cultivated specimens of many species to make seed available. The cost of seed is still high

compared to many other groups of plants, but this cost is considerably less than the price of a

germinated seedling or a plant poached from the wild. The value of providing someone with a
 3

plant that is legally obtained is inestimable. The relative availability of seed alone is an invitation

to the horticulturally curious to attempt germinating their own seeds, with the great benefit of

making the cost per plant reasonable for most collectors. The Cycad Society has a seed bank

available to its members that routinely offers seed of fairly rare species at reasonable prices, and

many members have developed formidable plant collections just by obtaining and growing

Cycad Society seed bank offerings over the years.

The major problems in growing most cycads from seed (though there are exceptions to this

generalization) are: 1) the seeds of most cycads have a fleshy sarcotesta (outer seed coat) with

germination inhibitors that must be removed; 2) when the ripe female cones of many cycads

disintegrate, dropping their seeds, embryos are often underdeveloped, requiring time, sometimes

several months, for the embryos to reach maturity and germination to become possible; and 3)

hard sclerotestas (inner, stony seed coat) of many cycad seeds resist penetration by moisture,

thus slowing germination. The end result of these factors is that cycad seeds under normal

greenhouse or shade house conditions, when they survive, germinate slowly and over a long

period of time - a perplexing scenario to many nurserymen.

Three papers (Dehgan and Johnson, 1983; Dehgan and Schutzman, 1983, 1989) explain the

relative impenetrability of a cycad seed coat and immaturity of the seed of some species at cone

dehiscence. The drawbacks to the proposed method are twofold: the potentially dangerous and/or

expensive chemicals to improve germination, notably concentrated sulfuric acid and gibberellin,

and the fact that only three species, Cycas revoluta Thunb., Zamia integrifolia L., and Zamia

furfuracea L.f., were tested, and optimal times and concentrations would have to be determined

for other species. A skilled nurseryman, taking proper safety precautions, could use the acid and

gibberellin method satisfactorily, but it is not feasible for a hobbyist or collector that may only
 4

want to grow small quantities of each species, seed of which can cost upwards of $5 each. In

fact, anecdotal evidence suggests even nurserymen were not as successful with the chemical

methods and laboratory exactitude that were used in the published papers. Anyone wishing to

germinate species other than Zamia integrifolia and Cycas revoluta would have to determine

chemical concentrations and exposure times to produce optimal germination rates.

Having heard anecdotal evidence of great success growing cycad seeds with a simple method

requiring only readily available materials and simple procedures, I investigated the “baggie

method” and found it successful and gratifying. Two cycad species were available to test for a

report to this conference. Many hobbyists have been discouraged by low germination rates when

attempting to grow costly cycads from seed. Low percentage germination, first and foremost, can

be related to seed viability, but attempting to germinate cycad seed in greenhouses or shade

houses under mist can result in high attrition of the percentage of seed that are viable due to

insects, microorganisms, and seed pilfering by rodents. Because the method considered here

allows seed to be kept in protected locations until planting, a higher success rate can be achieved.

The method is equally attractive because of the amount of space, money, and expertise necessary

to establish a mist system and attempt chemical seed treatments. Success could be instrumental

in rekindling the desire of many people to germinate cycads.

MATERIALS AND METHODS

Seed of two cycads became available in time for this trial, Cycas bifida (Dyer) K.D. Hill, and

Cycas revoluta Thunb. (King Sago) × C. taitungensis (Emperor Sago). Cycas bifida (Fork-leaved

Cycad), from China and Vietnam (Figs. 1 and 2) is little known in cultivation and quite rare, but

a friend and I successfully pollinated a female plant and produced seed (Figs. 3,4,5 and 6). A few

seeds were sacrificed to look for embryos, and they were visible but very small, suggesting that a
 5

maturation period was most likely necessary. I also performed the pollination of a Cycas

revoluta plant with C. taitungensis pollen in late spring of 2014. Both parents of the hybrid are

known to have immature embryos in seeds at the time female cones either dehisce or the

abscission layer between seeds and the megasporophylls are fully developed and seeds may

easily detach.

The sarcotestas of all seeds were removed, and cleaned seeds mixed with slightly moistened

sphagnum peat moss, and then put into freezer bags (Fig. 7) and sealed. In the case of Cycas

bifida, seed coat removal was easy because the sarcotestas scrape off with very little effort. The

C. revoluta × C. taitungensis seed required repeated soaking and whisking in wet coarse sand

with a cordless drill fitted with a wire wheel, and washing. This process was repeated over the

course of approximately two weeks to completely remove the sarcotestas. As mentioned earlier,

removal of sarcotestas was done to completely eliminate 1) any germination inhibitors that might

be present as well as 2) fleshy seed coat material that could decompose, potentially infecting and

killing viable seeds. The amount of water required to moisten the sphagnum peat moss was

approximately equal to the weight of the unmoistened peat moss. Some 259 Cycas bifida seeds

were put into freezer bags on 20 November 2014, and kept at room temperature on a desk in my

office.

RESULTS AND DISCUSSION

The first signs of germination in the bags was noticed in mid-February (Fig. 8). Germinated

seeds were taken from the bags (Fig. 9) and planted nine times during the four month period

from February 28 to June 27, 2015 (Fig. 10). Each time, any ungerminated seeds were placed

back into the baggies, and planting was done again each time emerging roots were seen at the

extremities of the baggies. After the June 27th planting, the few remaining seeds were judged
 6

inviable and discarded. Cumulative germination of this seed batch was approximately 95%, and

no decomposing seeds were seen during plantings. No attrition due to insects, microorganisms or

rodents was experienced. It is also worth mentioning that the baggies were not routinely opened

throughout the length of any of these experiments, and this seems not to have stopped

germination.

Because this species is known to possess a strong taproot, germinating seeds were planted in

well-drained mix (2-1-1 Fafard 2P-fine pine bark-sand) in deep tree pots (Fig. 10). Germination

was rapid (Fig. 11) and seedling growth appeared brisk (Fig. 12).

The other Cycas experiment was begun much later and has not yet been concluded. For the

purposes of this paper, germinating seeds were deliberately left in the freezer bags to see if they

would be damaged by remaining unplanted (Fig. 13). The seedling roots of Cycas revoluta × C.

taitungensis were tangled, requiring patience and time to separate without damage (Fig. 13).

However, the unplanted, germinating seeds were in good health at the submission of this

manuscript.

CONCLUSIONS

The success of the “baggie method” in germinating cycad indicates that it is worth trying on any

available cycad seed. It seems to be a worthwhile way to optimize the percentage of viable seed

brought from cone abscission to successful establishment in individual containers, and should be

considered by nurserymen and hobbyists alike.

Literature Cited

Dehgan, B. and C.R. Johnson. 1983. Improved seed germination of Zamia floridana (sensu lato)

with H2SO4 and GA3. Scient. Hort.19(3-4):357-361.

 7

Dehgan, B. and B. Schutzman. 1983. Effect of H2SO4 and GA3 on seed germination of Zamia

furfuracea. HortScience 18(3):371-372

Dehgan, B. and B. Schutzman. 1989. Embryo development and germination of Cycas seeds. J.

Amer. Soc. Hort. Sci. 114:125-129.

The Cycad Pages (http://plantnet.rbgsyd.nsw.gov.au/PlantNet/cycad/)

The World List of Cycads (http://cycadlist.org/)

 8

Fig. 1. Generalized distribution of Cycas bifida in China and Vietnam.

 9

Fig. 2. Mature female plant of Cycas bifida.

 10

Fig. 3. Unpollinated female cone of Cycas bifida (forefront).

 11

Fig. 4. Female Cycas bifida cone a few weeks after pollination.

 12

Fig 5. Mature female cone of Cycas bifida prior to dehiscence

 13

Fig. 6. Cleaned Cycas bifida seeds.

Fig. 7. Cleaned Cycas bifida seeds in freezer bag

 14

Fig. 8. Germinating seeds of Cycas bifida in freezer bag

Fig. 9. Germinating seeds of Cycas bifida ready for planting

 15

Fig. 10. Cycas bifida germinating seeds planted in tree pots

 16

Fig. 11. Cycas bifida seedling producing its first leaf.

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Fig. 12. Cycas bifida seedlings several months after planting

Fig. 13. Cycas revoluta × C. taitungensis seeds germinating and becoming tangled in baggie.