全 文 ：Journal of Systematics and Evolution 46 (2): 205–212 (2008) doi: 10.3724/SP.J.1002.2008.07073
(formerly Acta Phytotaxonomica Sinica) http://www.plantsystematics.com
Observations of several cryptomonad flagellates from China Sea by
scanning electron microscopy
1Xiao-Li XING 2Xu-Yin LIN 1Chang-Ping CHEN 1Ya-Hui GAO* 1Jun-Rong LIANG
1Hong-Zhou HUANG 1Bing-Qian LI 3King-Chung HO 4Yu-Zao QI
1 (Key Laboratory of the Ministry of Education for Cell Biology and Tumor Cell Engineering, School of Life Sciences,
Xiamen University, Xiamen 361005, China)
2 (School of Life Sciences, Xiamen Ocean Vocational College, Xiamen 361012, China)
3 (School of Science & Technology, Open University of Hong Kong, Hong Kong SAR, China)
4(Research Center of Hydrobiology, Jinan University, Guangzhou 510630, China)
Abstract Three species and one variety of cryptomonad flagellates belonging to three genera were observed
with the scanning electron microscope in sea water samples from Tolo Harbor (Hong Kong), Changjiang River
Estuary and Xiamen Harbor. They are Hemiselmis sp. Novarino, Plagioselmis prolonga Butcher ex Novarino,
Lucas & Morrall, Plagioselmis prolonga var. nordica Novarino, Lucas & Morrall and Teleaulax acuta (Butcher)
Hill. The taxonomic characteristics, ecological habit and distribution of the above species are described and the
LM and SEM photographs of the species are also presented. This is the first record of the genus Hemiselmis Parke
in the China Sea, and the species Plagioselmis prolonga and Teleaulax acuta have records of producing blooms in
the China Sea.
Key words China, cryptomonad, Hemiselmis Parke, Hemiselmis sp., marine, new record, Plagioselmis pro-
longa, Plagioselmis prolonga var. nordica, Teleaulax acuta.
Cryptomonad flagellates are widespread and
abundant in the sea, especially in the pelagic envi-
ronment (Haigh et al., 1992; Novarino, 2005), where
their photosynthetic ability contributes significantly to
primary productivity (Robinson et al., 1999). In
addition, they are of great evolutionary importance
owing to the presence of the nucleomorph (Lucas,
1970), and are environmentally important because
some cryptomonads can form nuisance blooms in
coastal areas (Andreoli et al., 1986; Dame et al.,
2000). By virtue of their small size (mostly less than
20 µm), unambiguous cryptomonads identification
generally cannot be accomplished by light micros-
copy. The taxonomic importance of cryptomonads in
marine ecosystems and their contribution to marine
primary productivity were largely ignored until the
cryptomonads diversity in Adriatic Sea was investi-
gated for the first time (Butcher, 1967). From his
study, Butcher (1967) recognized 47 marine species
from British coastal waters by light microscopy.
Subsequently, with the recent applications of electron
microscopic techniques in the study of cryptomonad
flagellates, a large number of previously unknown
cellular features have been revealed, many of which
are important taxonomic characters within the cryp-
tomonads (Klaveness, 1985; Hill, 1991a, b; Novarino
& Lucas, 1993a; Clay et al., 1999; Novarino, 2003).
So far, the taxonomic studies of marine crypto-
monads have been carried out only in a few countries
(Butcher, 1967; Throndsen, 1976; Hill, 1991a; No-
varino et al., 1997; Barlow & Kugrens, 2002; No-
varino, 2003, 2005; Cerino & Zingone, 2006), and
there was a first attempt at investigating cryptomonads
diversity by Novarino (2005) using SEM. In China,
only Hu et al. (2002) reported six species of the genus
Cryptomonas Ehrenberg (Cryptophyceae) from China
Sea, and the genus Cryptomonas has long been recog-
nized as being exclusively freshwater. This situation
resulted in the taxonomy of cryptomonad flagellates in
China relatively understudied. This study reports
several taxa from China Sea, the genera Plagioselmis
Butcher emend. Novarino Lucas & Morrall,
Hemiselmis Parke and Teleaulax Hill, and the species
and/or varieties Plagioselmis prolonga Butcher ex
Novarino, Lucas & Morrall, P. prolonga var. nordica
Novarino, Lucas & Morrall, Hemiselmis sp. Novarino
and Teleaulax acuta (Butcher) Hill. This is the first
record of Hemiselmis Parke in Chinese water, and the
only other report of Plagioselmis prolonga and
Teleaulax acuta in China was in a marine environ-
mental monitoring report of Hong Kong waters (Ag-
riculture, Fisheries and Conservation Department of
Hong Kong, 2006).

———————————
Received: 16 May 2007 Accepted: 30 Janurary 2008
* Author for correspondence. E-mail: .
Journal of Systematics and Evolution Vol. 46 No. 2 2008 206
1 Material and methods
Water samples were collected by the Environ-
ment Protection Department on May 8 and June 5,
2006 from Tolo Harbor, Hong Kong, China, by Xiaoli
Xing on June 6, 2004, from Xiamen Harbor, Xiamen,
China, and on May 20, 2005, from Changjiang
(Yangtze) River Estuaries of East China Sea in China.
Samples were prepared for examination using scan-
ning electron microscopy (SEM). Fresh samples were
fixed by 1% Lugol’s solution and collected on 2 µm
pore size nucleopore filters and rinsed with pure
redistilled water for 10 minutes, then dehydrated in a
series of 30%, 50%, 70% and 95% ethanol. The cells
were submerged in each dilution of ethanol for 15
minutes. Each sample was then critical point dried
with liquid CO2. The dried samples were sput-
ter-coated with gold for 25 seconds and examined
with a LEO 1530 scanning electron microscope.
Cell count samples were condensed from 500 mL
to a final volume of 10 mL by settlement for more
than 48 h. Then 1 mL samples were used for cell
counting. The samples were counted under a light
microscope, Olympus BH-2, and the counting cham-
ber for phytoplankton was used.
2 Results
1. Plagioselmis Butcher emend. Novarino, Lucas &
Morrall in Cryptogamie Algologie 15: 90. 1994.
斜片藻属
Description: Cells with flagella inserted apically
or subapically from a shallow anterior depression, the
vestibulum; presence of the furrow; with a character-
istically acute cell posterior (tail); and the presence of
a non-plated periplast on the tail and the hexagonal
plated periplast on the main portion of the cell body;
marine and freshwaters.
Type: Plagioselmis prolonga Butcher ex No-
varino, Lucas et Morrall.
Plagioselmis prolonga Butcher ex Novarino, Lucas &
S. Morrall in Cryptogamie-Algol. 15: 90. 1994;
Butcher in Fish. Invest. London, Ser. 4: 18. 1967; D.
R. A. Hill in Ann. Bot. Fenn. 29: 165. 1992a; M.
Kuylenstierna & B. Karlson in Botanica Mar. 37: 22.
1994.
伸长斜片藻 Figs. 1–9
Cells are 5–7 µm long and 3–3.7 µm wide. The
cell anterior is rounded whereas the posterior end is
acute (Figs. 1–4). Two slightly subequal flagella are
inserted subapically into the right side of the vestibu-
lum (which is a shallow anterior depression), the
longer (dorsal) flagellum is approximately as long as
the cell (Fig. 6). A relatively acute cell posterior- tail
is present. A periplast composed of hexagonal plates
on the main portion of the cell body and a non-plated
periplast on the tail are present (Figs. 5–7). The tail is
1/5–1/3 of the cell length. Ventral furrow is present,
and extends to about 1/2 of the cell length (Fig. 7),
and a mid-ventral band extends from the tip of the tail
to the base of the furrow (Figs. 8, 9).
The abundance of Plagioselmis prolonga reached
4.4 × 106 cells/L in our samples from Tolo Harbor,
Hong Kong, on May 8, 2006, and decreased by 102
times, i.e. 3.6 × 104 cells/L on June 5, 2006.
Habitat: Marine.
Distribution: This species was observed from the
samples of Tolo Harbor and Changjiang River Estu-
ary. Previous records include the North Atlantic:
British coastal waters (Butcher, 1967), Gulf of
Saint-Laurent, Canada (Bérard-Therriault et al., 1999),
Irish Sea and oyster basins south of La Rochelle,
France (Novarino, 2005); Pacific Ocean: Salton Sea,
California (Barlow & Kugrens, 2002); Baltic Sea
(Hill, 1992a); Mediterranean Sea: Adriatic Sea (Po
river delta lagoon) (Andreoli et al., 1986), Sea of
Alboran and Barcelona coast (Novarino, 2005).
Plagioselmis prolonga var. nordica Novarino, Lucas
& S. Morrall in Cryptogamie-Algol. 15: 90. 1994; G.
Novarino et al. in J. Plankt. Res. 19: 1096. 1997; L.
Bérard-Therriault et al. in Can. J. Fish. Aquat. Sci.
128: 249. 1999; G. Novarino in Nord. J. Bot. 11: 602.
1991a.
伸长斜片藻诺尔变种 Figs. 10, 11
The variety nordica is distinguished from the
typical variety by the absence of a furrow on the

→
Figs. 1–11. 1–9. The cell of Plagioselmis prolonga (Figs. 1–4 are LM and others are SEM). 1. A cell from the field, dorsal view. 2. A cell from the
field, lateral view. 3, 4. A cell from the field, dorsal-lateral view, and the prominent hyaline tail is visible. 5. A cell from the field, lateral view,
showing the rounded anterior end. 6. A cell from the field, dorsal-lateral view, showing the unequal flagella. 7. A cell from the field, ventral-lateral
view. The furrow in ventral view is clearly visible, as is the hexagonal periplast plates on the main portion of the cell body, and the non-plated
periplast on the posterior tail. 8. A cell from the field, dorsal view. 9. A cell from the field, dorsal view, showing the presence of mid-ventral band
(MVB) (arrow) in the tail, extending to the base of the furrow. 10, 11. The cell of Plagioselmis prolonga var. nordica in SEM. 10. A cell from the
field, ventral view. Note the absence of furrow. 11. A cell from the field, dorsal view, showing the tail (arrow) and the hexagonal periplast plates in
the main body of the cell.
Scale bars: 1–4=5 µm in LM; 5–9=1 µm in SEM; 10, 11=1 µm.
XING et al.: SEM observations of cryptomonad flagellates from China Sea

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Journal of Systematics and Evolution Vol. 46 No. 2 2008 208

















































Figs. 12–19. 12–15. The cell of Hemiselmis sp. in SEM. 12. A cell from the field, dorsal view, showing the typical hexagonal plates. 13. A cell from
the field, dorsal view, showing the presence of the ridges between the periplast plates, probably the result of inadequate fixation. 14. A cell from the
field, dorsal view, showing the typical, larger and elongate hexagonal plates. 15. A cell from the field, ventral view, showing the flagella inserted in
the subapical position. 16–19. The cell of Teleaulax acuta. (Fig. 16 is in LM and others are in SEM). 16. A cell from the field, lateral-dorsal view,
showing the rostrate anterior and a pointed posterior. 17. A cell in dorsal view, showing the acute posterior end. 18. A cell from the field, lat-
eral-ventral view, showing the presence of rostrate anterior and the ventral furrow. 19. A cell in ventral-lateral view, showing the presence of a long,
deep furrow extending for about half of the cell length, and bearing numerous ejectosome pores (arrow).
Scale bars: 12–14, 17–19=1 µm; 15 = 2 µm; 16= 3 µm.

ventral cell surface (Figs. 10, 11).
Habitat: Marine.
Distribution: Our samples were collected from
Tolo Harbor and Changjiang River Estuary. This
species has been reported in the Mediterranean Sea:
Adriatic Sea (Andreoli et al., 1986) and the North
Atlantic: southern North Sea (Novarino et al., 1997)
and St Lawrence estuary and gulf (Bérard-Therriault
et al., 1999).
Note: Novarino (2005) raised the status of the
variety nordica to a new species, P. nordica, mainly
according to the features of cells with the point of the
flagella insertion and absence of the ventral furrow.
However, there is no clear difference between the
variety nordica described here and P. prolonga,
except absence of the ventral furrow. Hence, the
XING et al.: SEM observations of cryptomonad flagellates from China Sea

209
variety nordica is still used here.
2. Hemiselmis Parke in Journal of the Marine Bio-
logical Association of the United Kingdom 28: 279.
1949.
半片藻属
Description: Cells with the rounded anterior and
posterior end, dorsal-ventrally flattened, appearing
reniform in lateral view; a furrow absent; periplast
composed of the characteristic, larger hexagonal
plates, and the sizes of the plate difference between
species; marine and freshwaters.
Type: Hemiselmis rufescens Parke.
Hemiselmis sp. G. Novarino in Sci. Mar. 69 (1): 61.
2005.
半片藻 Figs. 12–15
Cells are 4.2–4.5 µm long and 3.2–3.4 µm wide,
ovoidal in dorsal view, and the posterior and anterior
ends of cells are rounded (Figs. 12, 14, 15). There are
two almost equal flagella inserted in a median or
subapical position (Figs. 12, 14, 15). It seems to
possess a very short ventral furrow, possibly attributed
to the absence of the hexagonal plates near the fla-
gella, and no posterior tail is present (Fig. 15). The
periplast is composed of the typical, larger and elon-
gate hexagonal plates about 0.8–1.5 µm long (Figs.
12–15).
Habitat: Marine.
Distribution: Our samples were collected from
Changjiang River Estuary.
Note: It could be asked if this cryptomonad
should be assigned to a different genus because the
feature of the periplast composed of hexagonal plates
is also found in members of the genera Plagioselmis
and Rhinomonas Hill, but the size of the hexagonal
plate is different within the three genera. The hexago-
nal periplast plates of Hemiselmis are larger than those
of Plagioselmis, and are elongate (Barlow & Kugrens,
2002). The cells of Plagioselmis have a characteristic
tail with non-plate, but the cells of Hemiselmis have
no tail and larger hexagonal plates are exhibited on all
surfaces of the cells. The plates of Rhinomonas were
also hexagonal but usually smaller than those of
Plagioselmis. So, the typical features of the larger
hexagonal periplast plates and no tail, as well as the
bean-like or reniform shape of the cells, made it
possible to distinguish Hemiselmis sp. from other
cryptomonads.
3. Teleaulax Hill in Phycologia 30: 177. 1991a.
全沟藻属
Description: Cells with an acute posterior and a
wide furrow on the ventral face reaching well into the
median region of the cell; marine.
Type: Teleaulax acuta (Butcher) Hill = Crypto-
monas acuta Butcher
Note: The genus Teleaulax was removed from
the genus Cryptomonas by Hill (1991a) because the
features of Teleaulax were different from the typical
characteristics of Cryptomonas. Cells of the former
have a long furrow and no plate-type periplast exists.
Teleaulax acuta (Butcher) D. R. A. Hill in Phycolo-
gia. 30 (2): 177. 1991a; D. R. A. Hill in Ann. Bot.
Fenn. 29: 173, 174. 1992b; L. Bérard-Therriault et al.
in Can. J. Fish. Aquat. Sci. 128: 250. 1999.
——Cryptomonas acuta G. Butcher in J. Mar. Biol.
Assoc. UK. 31 (1): 188 1952; G. Novarino in Nord. J.
Bot. 11: 602. 1991a; G. Novarino et al. in J. Plankt.
Res. 19: 1094. 1997. Non Cryptomonas acuta sensu F.
H. Chang in N. Z. J. Mar. Freshwater Res. 17: 291.
1983.
尖尾全沟藻 Figs. 16–19
Cells are 5.6–8.4 µm long and 2.8–3.7 µm wide,
with a strongly rostrate anterior and a pointed poste-
rior (Fig. 16). There are two equal flagella about 2/3
the cell length. They possess a characteristically acute
posterior end (Fig. 17), and a long furrow that extends
longitudinally from the vestibulum, extending roughly
halfway along the cell length and bearing numerous
ejectosome pores, usually four longitudinal rows of
large ejectosomes (Fig. 19). The periplast does not
have the contour of a plate-type periplast (Figs.
17–19). There are many small ejectosomes on the
entire cell surface, including the posterior end (Figs.
17–19).
Habitat: Marine.
Distribution: Our samples were collected from
Tolo Harbor and Changjiang River Estuary. This
species has been previously found in the Mediterra-
nean Sea: Adriatic Sea, port of Barcelona (Novarino,
2005) and the North Atlantic: Baltic Sea (Hill, 1992b),
the southern North Sea (Novarino et al., 1997), and
the St. Lawrence estuary and gulf (Bérard-Therriault
et al., 1999), Victoria, Australia (Hill, 1991a).
3 Discussion
Because the samples in this investigation were
collected in 2004, 2005, and 2006, and fixed with
Lugol’s solution in the field, the study of isolation and
culture of the living cells, and the identity of pigment
and the molecular phylogeny could not be done.
Therefore, the identification in this paper was based
mostly on the main features of the periplast revealed
by SEM.
The shape of the periplast plates has been used
Journal of Systematics and Evolution Vol. 46 No. 2 2008 210
extensively as a taxonomic feature at the generic level
(Santore, 1984; Kugrens & Lee, 1987; Hill & Weth-
erbee, 1988, 1989, 1990; Hill, 1991a, b; Novarino,
1991a, b; Novarino & Lucas, 1993a, b, 1995; No-
varino et al., 1994; Clay et al., 1999; Kugrens et al.,
1999; Novarino, 2005; Cerino & Zingone, 2006). The
particular feature of the genus Plagioselmis with the
presence of a non-plated periplast on the tail and the
hexagonal plated periplast on the main portion of the
cell body make it possible to distinguish all members
of this genus from other cryptomonads (Novarino,
2005; Cerino & Zingone, 2006). The diagnostic
features of the genus Plagioselmis are shown in
figures 5–11. The features of P. prolonga shown in
figures 5–9 are consistent with the descriptions and
features shown in other publications (Novarino et al.,
1994, 1997; Novarino, 2003, 2005; Cerino & Zingone,
2006). The variety nordica is differentiated from P.
prolonga with absence of the ventral furrow. Al-
though figure 11 shows the dorsal view of P. prolonga
var. nordica, the features in figure 11 are almost
identical to those in figure 15 of Novarino et al.
(1997).
The diagnostic features of the genus Hemiselmis,
with the bean-like or reniform shape, the larger hex-
agonal plated periplast, even elongate (Barlow &
Kugrens, 2002), and the presence of hexagonal perip-
last on the cell posterior, clearly set Hemiselmis apart
from other cryptomonads. Hemiselmis sp. here shares
the characteristics of strains by Novarino (2005), but
the size of the hexagonal plates is different. No-
varino’s Hemiselmis sp. (see Fig. 6) has scales
0.4–0.65 µm in size, while figure 14 here shows a cell
with larger size of hexagonal plates more than 1.0 µm.
The cells of the former were 4.2–4.5 µm long and
3.2–3.4 µm wide, while the cells of the latter were
about 4.1 µm long and 2.5 µm wide according to
figure 6 (C, D). Meanwhile, Hemiselmis sp. here also
had the similarities with Hemiselmis sp. by Cerino &
Zingone (2006), but the size of hexagonal plates
(about 0.8–1.5 µm, average more than 1.0 µm) of the
former were also larger than those of the latter (about
0.5 µm). However, Novarino’s Hemiselmis sp. (Fig 3:
C, D) also had larger and elongate size of hexagonal
plates (1.7–2.5 µm in size) (Novarino, 2005), and the
cell shape was very similar to that of Hemiselmis sp.
in this paper. Because our samples were fixed in field,
and the identity of the other features such as the
phycobilin pigment and presence of the refractive
body were unknown, all mentioned made it difficult to
identify them as the same species. In addition, the
features revealed by SEM here could not be compara-
ble with those eight new species of Hemiselmis in LM
reported by Butcher (1967), so Hemiselmis sp. de-
scribed here cannot be identified unambiguously as a
new species instead of belonging to any of those eight
by Butcher (1967).
The genus Teleaulax can be identified unambi-
guously due to its characteristically acute posterior
end where the non-plated periplast was covered and
the long furrow extended roughly halfway along the
cell length in these samples (Figs. 12–15). The cells of
Teleaulax superficially resemble those of Plagioselmis
by their acute posterior end. However, the periplast
covering the entire cell surface, including the posterior
end, helps to identify them clearly as Teleaulax. The
features shown in Figs. 12–15 were in accordance
with other publications such as Novarino et al. (1997),
Novarino (2005), and Cerino & Zingone (2006). The
cells possess many small ejectosomes on the entire
cell surface, including the posterior end. Kugrens et al.
(1994) suggested that these ejectosomes on the surface
are within the cell and are extruded once they are
stressed.
So far, the systematics of the cryptomonads is
based predominantly on ultrastructural features (Dean
et al., 2002). Several molecular phylogenetic studies
have been done by using nuclear SSU rDNA sequence
data (Marin et al., 1998; Clay & Kugrens, 1999; Dean
et al., 2002; Hoef-Emden et al., 2002). These studies
showed that cryptomonads with plastids represent a
monophyletic group. Plagioselmis prolonga was
closely related to Teleaulax species, and form a clade
together with Geminigera cryophila Taylor & Lee
(Dean et al., 2002). Hemiselmis species form a mo-
nophyletic group within the one clade (including
genus Hemiselmis, Chroomonas Hansgirg and Komma
Hill) (Marin et al., 1998; Dean et al., 2002). The
monophyletic evolution of cryptomonads showed that
the three genera described here, previously identified
mainly based on the ultrastructure features, are reli-
able. The molecular sequencing studies have provided
a number of new phylogenetic information (Marin et
al., 1998; Dean et al., 2002; Hoef-Emden et al., 2002).
However, the molecular sequence data that currently
exist in Genbank seem not to play a major role in
cryptomonad identification (Novarino, 2003).
Plagioselmis prolonga and Teleaulax acuta are
the most widespread and abundant cryptomonads in
several investigated sea areas (Novarino et al., 1997;
Barlow & Kugrens, 2002; Novarino, 2005; Cerino &
Zingone, 2006). The species Plagioselmis prolonga
was recorded throughout the year, and often reached
high population densities (e.g., Cerino & Zingone,
XING et al.: SEM observations of cryptomonad flagellates from China Sea

211
2006). Its abundance reached 106 cells/L in our sam-
ples. However, Hemiselmis sp. was recorded only at
certain periods of the year (Cerino & Zingone, 2006).
On the other hand, Plagioselmis prolonga and
Teleaulax acuta can cause blooms (Environmental
Protection Department, Hong Kong, 2005), although
they have not been found to produce toxins. The
taxonomic identifications are necessary to any studies
of such blooms, and the key taxonomic characteristics
as defined here using SEM will be helpful to accurate
taxonomy of future blooms.
Acknowledgements We are grateful to Hong Kong
Environmental Protection Department for supplying
the samples from Tolo Harbor, and to Professor
Hong-Jun HU (Wuhan Botanical Garden, the Chinese
Academy of Sciences) for his valuable comments and
for his supplying many references on the cryptomonad
flagellates studies. The authors also thank Ms. Ru
XUE and Mr. He-Sheng ZHAI in Electron Micro-
scope Lab, Xiamen University, for their assistance in
EM operation. This work is supported by the National
Natural Science Foundation of China, Grant No.
40627001, the Major State Basic Research Develop-
ment Program of China (973 Program), Grant No.
2005CB422305, and the Program for Innovative
Research Team in Science and Technology in Fujian
Province University.
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中国海区几种隐藻类鞭毛藻的扫描电镜观察
1邢小丽 2林旭吟 1陈长平 1高亚辉* 1梁君荣 1黄鸿洲
1李炳乾 3何建宗 4齐雨藻
1(厦门大学生命科学学院细胞生物学与肿瘤细胞工程教育部重点实验室 厦门 361005)
2(厦门海洋学院生命科学系 厦门 361012)
3(香港公开大学科技学院 香港)
4(暨南大学水生生物研究所 广州 510630)

摘要 报道了来自香港吐露港、中国长江口及厦门港的3个属的3种隐藻及1个变种, 即半片藻Hemiselmis sp. Novarino、伸长
斜片藻Plagioselmis prolonga Butcher ex Novarino, Lucas & Morrall、伸长斜片藻北方变种Plagioselmis prolonga var. nordica
Novarino, Lucas & Morrall、尖尾全沟藻Teleaulax acuta (Butcher) Hill, 并对每个种类的分类特征、生态分布进行描述, 同时提
供每个种的光镜和扫描电镜照片。其中, 半片藻属Hemiselmis Parke是中国海区首次记录的属, 而伸长斜片藻Plagioselmis
prolonga和尖尾全沟藻Teleaulax acuta可以引发赤潮。
关键词 中国; 隐藻; 半片藻属; 半片藻; 海洋; 新记录; 伸长斜片藻; 伸长斜片藻诺尔变种; 尖尾全沟藻