Cytotaxonomic Study of 12 Species in the Polypodiaceae from Southern China

2011·5 MB·English

by  Ren-Xiang Wang

Cytotaxonomic Study
of 12 Species
in the
Polypodiaceae from Southern China
Wang
Ren-Xiang
School of Life Sciences, Guangxi Normal University, Guilin 541004, China
P. R.
Key
Laboratory of Ecology of Rare and Endangered Species and Environmental
Protection
Normal
(Guangxi
University), Ministry of Education, Guilin 541004, China
P. R.
Wen
Shao
Institute of Ecology and Geobotany, Yunnan University, Kunming 650091. China
P. R.
Shanghai Chenshan Academy
Plant Science Research Center, Chinese of Sciences, Chenshan
Botanical Garden, Shanghai, 201602, China
P. R.
Shu-Gang Lu
Institute of Ecology and Geobotany, Yunnan University, Kunming 650091, China
R.
P.
Shan-Yi Zhou and Shi-Chu Liang
School of Life Sciences, Guangxi Normal University, Guilin 541004, China
P. R.
Key
Laboratory of Ecology of Rare and Endangered Species and Environmental
Protection
(Guangxi Normal
University), Ministry of Education, Guilin 541004, China
P. R.
The
Polypodiaceae one most
is of the diverse groups of extant ferns (Schneider
They
et al, 2004). are not only a lineage of derived ferns that has a high number of
species, but they also display a vast range of morphological variation (Kreier
et
SW
al, 2008). Ching believed that China (especially China and the Himalayas)
was
the center of origin of Polypodiaceae, W\ih about half the number of the
total
species of Polypodiaceae occurring there (Ching, 1979; Lin, 2000).
and may
Cytological reproductive studies provide very useful information
phylogenetic and
for evolutionary studies of various groups of Data on
ferns.
chromosome
numbers, base numbers,
karyotypes, aneuploids, polyploidy
and modes
levels, reproductive are useful for understanding of the origin,
* Corresponding [email protected]
author: E-mail:
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evolution and speciation of the ferns (Manton, 1950; Wang, 1984; Wu, 1984;
Wang
et al, 2007). Data on chromosomes are also important discriminating
for
and
species species complexes.
Previous cytological studies have recorded chromosome numbers
in less
than 60 species of Polypodiaceae in South China (Yunnan, Zhejiang, Hainan,
and
Fujian) (Wang, 1984; Weng, 1990; Kato et al, 1992; Cheng, 1992; Kato and
Nakato, Nakato number
1999; et al., 1995; Lin et al., 1996, 2002). This accounts
23%
for only of the Chinese Polypodiaceous Data on chromosomes
ferns. of
some =
genera/species in Polypodiaceae have been reported, such as 2/7 72 (2x)
in Lepidogrammitis rostrata, Pbymatopteris oxyloba and Lepisorus macro-
= =
spbaerus; 2n 108 Pbymatopteris and 2n
(3x) in 70 Lepisorus
trisecta; (2x) in
bicolor and so on (Wang Wang and Lu and Wang,
et 2007; Lu, 2008;
2008).
al.,
Cytological research on these ferns has been severely lacking, because of
manipulating due
difficulties in root tips to their small size, inaccessibility of
and number
the species, the large of chromosomes. Cytological studies of the
southern China Polypodiaceous ferns can contribute our understanding
to of
and we
the origin evolution of the Polypodiaceae. Accordingly, chose 12
species in eight genera of Polypodiaceae as materials, following the alphabet
Chromosome
order. counts had already been performed
for eight of the species
=
used in this study: n 36 in Colysis elliptica (Tsai and Shieh, 1983; Weng,
= =
1990), 2/7 144 (4x) in C. bemionitidea (Mehra, 1961), 2/7 72 (2x) in C.
=
and
potbifolia (Tsai Shieh, 1983; Kato
et al., 1992), 2/7 50, 51, 75, 76, 100, 101,
and
102 103 in Lepisorus tbunbergianus (Takamiya, n=3G, 37
1996), in
=
Microsorium fortune! (Roy and Holttum, 1965; Mitui, 1973), 2/7 72 (2x) in
=
Pbymatopteris crenatopinnata and
(Panigrahi Patnaik, 1961), 2/7 74 (2x) in
=
Pbymatosorus cuspidatus (Kato and Nakato, 1999), and 2/7 72 (2x) in Pyrrosia
lingua (Takei, 1969, 1983a). The aim of study provide new
this to cytological
is
information on and
the classification phylogenetics Polypodiaceous
of
ferns.
Materials and Methods
Living plants (Table Fig. were collected from the field in Yunnan, Guangxi
1. 1)
and Hainan
provinces, southern China, and Voucher
cultivated in the
lab.
specimens were deposited Herbarium Yunnan
in the of University (PYU).
For the examination of mitotic chromosomes, were
root tips pretreated in
mol/L
0.002 8-hydroxyquinoline solution for 3-6h before being fixed in
Carnoy's solution (95% ethanol: glacial acetic acid=3:l) 12-24 then they
for h;
were hydrolyzed in 1 mol/L HCl at 60 C for 10-15 min. After washing 3-4
2%
times to eliminate residual hydrochloric acid, materials were stained in
aceto-orcein and squashed by the usual method (Manton, The "second
1950).
squashed" method (Wang and Zhang, 1981) was used when chromosomes
occurred in different planes. Mitotic were examined and photographed
cells
by using an Olympus BX51-DP70 photomicroscope. More than 30 chromo-
some
micrographs were observed each
in individual.
In the higher leptosporangiate ferns, there are two reproductive types: the
sexual reproductive type with 64 spores sporangium, and apogamous
in a the

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type with 32 spores (Manton, 1950; Lovis, 1977; Walker, 1979). To determine
number
the reproductive type, the of spores per sporangium was counted. At
unopened
sporangia were counted each
least five in individual.
Results and Discussion
The chromosome
photographs
of root tip materials of 12 species in eight genera
shown
Polypodiaceae and The chromosome numbers
of are in Figures 2 for four
3.
newly 2n=72
species are reported China: Lepidogrammitis drymoglos-
for (2x) in
2/7=72 2n=72
soides, (2x) in Neolepisorus ovatus, [2x] in Phymatopteris
and
rhynchophylla, 2/7=72 (2xj in Phymatosorus hainanensis. Four records are
new =
2/7=72
cytotypes: (2x) in Colysis hemionitidea, 2/7 72 [2x] in Lepisorus
thunbergianus, 2/7=108 in Phymatopteris crenatopinnata, and 2/7=108
{3x] {3x)
Phymatosorus
in cuspidatus. Lepisorus thunbergianus 2/7=72 has the base
[2x)
number x=36, which
different from the previous reports x=25, and
is 38, 50, 51.
We
confirmed the counts of 2/7=72 (2x) in Colysis elliptica, 2/7=72 (2x) in C.
2n=72
pothifoha, (2x) in Microsorium fortunei, and 2/7= 72 (2x) in Pyrrosia lingua.
The
reproductive type in P. cuspidatus and P. crenatopinnata was
apogamous, whereas was
in the other species of the sexual reproductive
it
The chromosome number mode
and
type. reproductive for each species are
and
described discussed below.
WANG
ET AL: CYTOTAXONOMIC STUDY OF POLYPODIACEAE CHINA
IN
^^^^
_
B
— —
D
—
H
G
*
Fig. Photographs of somatic chromosomes metaphase =
2. at A. Colvsis alliptica. 2/i 72. B. C.
hemionitidea, 2n = = =
72. C. C. pothifolia, 72. D. Lepidogrammitis drymoglossoides, 2n
2/2 72. E.
Lepisorw. timnbi rgianus An- Microsorium 2n = =
72. F. fortunei, 72. G. Neolepisorus ovatus, 2/3 72.
H. Phymatopteris crenatopinnata, 2n = =
108. P. rhvnchophylla, 2n Phvmatosorus
72.
I.
].
= =
cuspidatus, =
2/1 108. K. P. hainanensis, 2n 72. L. Pyrrosia lingua, 2n 72 (sca/e fcars=5nm).
Co7ysis elliptica (Thunb.) Ching
known
This species from across India, Vietnam, and southern
is Japan,
The
China. material from Mulun, Guangxi counted had
study
in this 72
chromosomes
The
in the mitotic 2A,
cells (Fig. 3A). species a diploid, with
is
64 spores per sporangium.
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J^Z-^^
B
A,
(I
.©J
= =
Fig. 3. Explanatory drawings to Fig. 2. A. Colysis elliptica, 2n 72. B. C. hemionitidea, 2n 72. C.
= = =
2n 72. F. Microsorium fortunei, 2n 72. G. Neolepisoms ovatus, 2n 72. H. Phymatopteris
= = =
crenatopinnata, 2n 108. L P. rhynchophylla, 2n 72. Phymatosorus cuspidatus, 2n 108. K. P.
J.
= =
bamanensis. 2n 72. L. Pyrrosia lingua. 2n 72.
Tsai and Shieh (1983) reported the material from Taiwan, China had 36
Weng
chromosomes
in the meiotic cells. {1990} suggested the material from
China had chromosomes and
Fujian, also 36 in the meiotic 64 spores per
cells,
sporangium, which suggests a diploid. Kato et (1992) believed the
it is al.
from Yunnan, had chromosomes The
material Dali, 72 in the mitotic
cells.
WANG
ET CYTOTAXONOMIC STUDY OF POLYPODIACEAE CHINA
AL: IN
materials from five Japanese populations had 36 chromosomes in the meiotic
cells
(Mitui, 1966, 1968; Kurita, 1968; Takei, 1983b), and 64 spores per sporangium,
which
suggested that these species are sexual diploids. These results indicated
stability of chromosome numbers in the different populations in China and Japan.
The made by Love chromosome
record
et (1977) shovi^ed that different
al.
numbers
occur genus
in six species of the Colysis, indicating that diploid
= = =
(2n 72), triploid (2n 108), tetraploid (2u=144), and hexaploid (2n 216)
hemionitidea ex
C. (wlll. Mett.) C. Presl
This species w^idespread in tropical and subtropical Asia. The material
is
from Guilin, Guangxi counted in this study had 72 chromosomes in the mitotic
cells (Fig. 2B, 3B). Sixty-four spores per sporangium were found, suggesting
it
new
a sexual diploid. This record a cytotype.
is is
Mehra shown
(1961) the material from Mt. Himalaya had 144 chromosomes in
and
the mitotic cells, 64 spores per sporangium, suggesting a sexual tetraploid.
it is
C. pothifolia (D. Don.) C. Presl
This species widespread in tropical and subtropical Asia. The material
is
from Guilin, Guangxi counted in this study had 72 chromosomes in the mitotic
We
on
cells (Fig. 2C, 3C). suggest this species a sexual diploid based the
is
observation of 64 spores per sporangium.
Tsai and Shieh (1983) found that the material from Taiwan, China had 36
chromosomes
in the meiotic cells. Kato et (1992) collected material from
al.
Yunnan
and
Dali, Lin et (2002) collected materials from Fujian, each of
al.
which had 72 chromosomes in the mitotic All materials above had 64
cells.
spores per sporangium, suggesting that these are sexual diploids.
Lepidogrammitis drymoglossoides Ching
(Bak.)
This species endemic to China, distributed widely in the southern region.
is
The material from Bama, Guangxi counted study had 72 chromosomes
in this
in the mitotic cells 2D, 3D). Sixty-four spores were present in each
(Fig.
chromosome
sporangium, suggesting that the species a sexual diploid. This
is
count
is reported for the first time.
Lepisorus thunbergianus Ching
(Kaulf.)
known
This species from across southern China, Japan, and the
is
Philippines. The material from Huanjiang, Guangxi study was found
in this
=
have 2n chromosomes The
to 72 in the mitotic cells (Fig. 2E, 3E). species a
is
new
sexual diploid, with 64 spores per sporangium. This a cytotype.
is
=
Lin et (2002) reported that the material from Fujian had 2n 100
al.
chromosomes and was
in the mitotic cells, suggested a sexual tetraploid.
it
Lepisorus thunbergianus a cytologically complicated species, chromo-
is its
=
some numbers have been variously reported as n 2/7=50,
25, 50, 51, 75, 76,
100, 101, 102 and 103, including examples of polyploidy and aneuploidy from
shown
Japan (Takamiya, 1996). has been that diploids, triploids, tetraploids,
It
and
polyploids thunbergianus (Takamiya,
exist in L. 1996).
Microsorium Moore) Ching
fortunei
(T.
known
This species from across southern China, Burma, Bhutan, and
is
Vietnam. Material from Guilin, Guangxi in this study had 72 chromosomes in
AMERICAN FERN VOLUME NUMBER
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the mitotic cells (Fig. 2F, 3F). Sixty-four spores were present each
in
sporangium,
suggesting
a sexual diploid.
is
it
Roy
and Holttum
M.
(1965) reported fortunei from southern China had
36
chromosomes
in the meiotic suggesting the same
cells, results as this paper. In
Mitui
contrast, (1973) suggested the material from Japan had 37 chromosomes
may
in the meiotic which be
cells, the result of aneuploidy.
Neolepisorus ovatus Ching
(Bedd.)
This species widespread
is in southern regions of China and The
India.
material from Guilin, Guangxi counted in this study had 72 chromosomes
in
the mitotic There were
cells (Fig. 20, 3G). 64 spores per sporangium. The
results suggest should be considered
it a sexual diploid species. This
chromosome
count
reported
is for the first time.
Phymatopteris
crenatopinnata
(C. B. Clarke) Pichi-Serm.
This NE
species occurs in southern China and The
also in India. material
from Weixi, Yunnan, had chromosomes
108
in the mitotic 2H,
cells 3H).
(Fig.
We
counted
32 spores per sporangium. The resuUs apogamous
suggest
is
it
=
with 2n 108 new
in this study. This a cytotype.
is
and
Panigrahi Fatnaik
(1961) reported the material from eastern India had 72
chromosomes
in the mitotic and 64 spores per sporangium,
cells, suggesting
it
sexual
a diploid.
is
rhynchophylla
P. (Hook.) Pichi-Serm.
This species widespread and The we
is in tropical subtropical Asia. material
collected from Pingbian, Yunnan, had chromosomes
72
in the mitotic
cells
There were
(Fig. 21, 31). 64 spores per sporangium, suggesting a sexual
it is
chromosome
diploid. This count
reported
is for the first time.
Phymatosorus
cuspidatus
Don.) Pichi-Serm.
(D.
known
This
species is from across southern China, India, Vietnam, Nepal,
and The
Thailand. material from Bama, Guangxi counted had
in this study 108
chromosomes
in the mitotic cells (Fig. Thirty-two spores were present
2
J, 3J).
per sporangium. The results indicate an apogamous with 2n=108
is triploid
it
new
and
represents a
cytotype.
it
Kato and Nakato
(1999) reported the material from Hainan had 74
chromosomes
in the mitotic and 64 spores per sporangium,
cells, suggesting
a sexual diploid.
it is
hainanensis Lu
P. (Noot.) G.
S.
This found
species is in Hainan, China and also in India and Vietnam. The
we
material collected from Mt. Wuzhishan, Hainan, had chromosomes
72
in
We
the mitotic cells (Fig. 2K, 3K). counted 64 spores per sporangium,
suggesting a sexual diploid. This chromosome count
it is is reported for the
time.
first
Pyrrosia lingua
(Thunb.) Farwell
known
This species from across southern China, Vietnam, and
is India,
The
Japan. material from Pingbian, Yunnan counted study had
in this 72
chromosomes
in the mitotic The
cells (Fig. 2L, 3L). species a sexual diploid,
is
with 64 spores per sporangium.
WANG
CYTOTAXONOMIC
ET AL: STUDY OF POLYPODIACEAE CHINA
IN
Takei
(1969, 1983a) believed the material from Japan also had 72
chromosomes
Our
in the mitotic
cells. result suggests that Pyrrosia lingua
is
in a stable
state.
This study was supported by
the National Natural Science Foundations of China, Grant No.
(31060030, 30970186), the Natural Science Foundation of Guangxi (2011GXNSFA018089), and
the
Foundation Key
of Laboratory of Ecology of Rare and Endangered Species and Environmental
Normal We
Protection (Guangxi University), Ministry of Education (1001Z017). sincerely thank
Wei Ming Chu Yunnan
Prof. of University for providing valuable references cited in this paper, and
Chao Deng
assistant Profs. Xi (Guangxi Hechi University), and Qiang Xie (Guangxi Normal
University) for providing experimental materials in this paper. The authors would also thank Prof.
Cindy Q. Tang of the Golden Gate National Parks Conservancy and Damian Shea North
Prof. of
Carolina State University for helpful suggestions and a critical reading of the manuscript.
AMERICAN FERN VOLUME NUMBER
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