Cyclotella
species were collected at 51 sites from July 2010 to June 2013 in Korean coastal waters. A total of five
Cyclotella
species (
C. atomus
var.
marina, C. baltica, C. litoralis, C. meduanae
, and
C. meneghiniana
) were identified in this study. The diagnostic characteristics for five
Cyclotella
species are described, focusing on the spacing, position, number of satellite pores of the mantle fultoportula and valve face fultoportula. In addition, we put the salinity ranges of five species of
Cyclotella
together. Of the five
Cyclotella
species,
C. baltica, C. litoralis
and
C. meduanae
are newly recorded in Korea.
INTRODUCTION
The genus
Cyclotella
(Kützing) Brébisson is represented primarily in freshwater habitats, but many species, such as
C. atomus
Hustedt,
C. baltica
(Grunow) Håkansson,
C. caspia
Grunow,
C. choctawhatcheeana
Prasad,
C. cryptica
Reimann, Lewin
et
Guillard,
C. desikacharyi
Prasad,
C. litoralis
Lange
et
Syvertsen,
C. meneghiniana
Kützing,
C. scaldensis
Muylaert
et
Sabbe,
C. striata
(Kützing) Grunow and
C. stylorum
Brightwell occur in marine or brackish waters (Håkansson 2002, Prasad and Nienow 2006, Tanaka 2007).
Cyclotella
are characterized by having circular valves with different ornamentation between the central area and the marginal area of the valve face. Due to the morphological similarity among species and high intra-specific variation, taxonomic discriminations of the
Cyclotella
species are hard to unravel (Håkansson and Kling 1994, Meyer and Håkansson 1996). In addition, further taxonomic confusion is caused by the small size of the cells and numerous similarities among the species belonging to
Cyclotella
sensu lato. Recently Håkansson (2002) published the genus
Puncticulata
Håkansson from nine
Cyclotella
species based on the central area of valve faces, as well as the particular locations of fultoportulae and rimoportulae. Houk and Klee (2004) established the genus
Discostella
Houk
et
Klee from all stelligeroid
Cyclotella
taxa according to the position of the mantle fultoportulae and rimoportula on marginal striae.
In Korea, Lee and Lee (1988) firstly examined six
Cyclotella
species from southern and western coastal waters of Korea using scanning electron microscopy. Lee et al. (1994) studied the fine structure of
Cyclotella pseudostelligera
Hustedt. Lee et al. (1995a) reported 15
Cyclotella
species of diatoms in Korea. Lee et al. (1995b) originally described
Cyclotella orientalis
from the Nakdong River. Lee (1995) made a check-list and added two
Cyclotella
species in Korean waters. Cho (1996) founded nine
Cyclotella
species at Nakdong River,
C. woltereckii
Hustedt was firstly reported in Korea. Recently, Chung et al. (2010) first reported
Cyclotella atomus
var.
marina
from a diatom assemblage attached to an eelgrass sample at the Yeosu
Sampling information ofCylcotellain the Korean coastal waters1ES, East Sea,2SS, South Sea,3YS, Yellow Sea.
Sampling information of Cylcotella in the Korean coastal waters 1ES, East Sea, 2SS, South Sea, 3YS, Yellow Sea.
coast in Korea.
Although 21 species of
Cyclotella
diatom have been recorded in Korea (Lee et al. 1995a, Lee 1995, Cho 1996, Chung et al. 2010), several
Cyclotella
species have been transferred into the other cyclotelloid genera, such as
Discostella
and
Puncticulata
, following the recent complication of cyclotelloid diatoms (Håkansson 2002, Houk and Klee 2004): four
Cyclotella
species (
C. bodanica
Eulenstein,
C. comta
(Ehrenberg) Kützing,
C. comta
var.
glabriuscula
Grunow,
C. radiosa
(Grunow) Lemmermann) were transferred into the genus
Puncticulata
; five
Cyclotella
species (
C. comta
var.
glabriuscula
Grunow,
C. orientalis
Lee, Chung et Gotoh,
C. pseudostelligera
Hustedt,
C. stelligera
(Cleve et Grunow) Van Heurck,
C. woltereckii
Hustedt) were transferred into the genus
Discostella
.
We present the fine structure, description, distribution and taxonomic remarks of five
Cyclotella
species in Korean waters. Of the five species discussed in this paper, we include three species which are newly recorded in Korea.
MATERIALS AND METHODS
Sample collections were carried out at 51 sites from July 2010 to June 2013 in Korean coastal waters (
Table 1
). Phytoplankton were collected using a 20 μm mesh net by horizontal and/or vertical towing. Samples were immediately fixed with neutralized formalin (5% final concentration). Using a method presented by Hasle and Fryxell (1970), organic matter in the diatom cell was removed. Prepared diatom material was observed under a light microscope (Axioskop 40; Carl Zeiss, Oberkochen, Germany) and a scanning electron microscope (JSM-5600LV; JEOL, Tokyo, Japan), and was photographed with an MRc5 camera (Carl Zeiss). Diatom size analysis was completed with image calculation software (AxioVision AC version 4.5; Carl Zeiss). The general terminology was adopted from the proposals on Diatom Terminology (Anonymous 1975, Ross et al. 1979), and some special terms for
Cyclotella
followed Håkansson (2002) and Tanaka (2007).
RESULTS AND DISCUSSION
A total of five
Cyclotella
were identified in the present study, we present keys for five
Cyclotella
species based on morphological characteristics, such as the space, position, number of satellite pores of the mantle fultoportula and valve face fultoportula (
Table 2
).
- Key to 5 Cyclotella species in Korea
1a. Two satellite pores on mantle fultoportula ..................2
1b. Three satellite pores on mantle fultoportula ...............4
2a. No central fultoportula ..............
C. atomus
var.
marina
2b. Several central fultoportula ...........................................3
3a. Mantle fultoportula on recessed interstria ...
C. litoralis
3b. Mantle fultoportula on thickened interstria...
C. baltica
4a. Space of mantle fultoportula on every 2nd interstria ...........................................................................
C. meduanae
4b. Space of mantle fultoportula on every interstria ..............................................................................
C. meneghiniana
- Description of species
- Cyclotella atomus var. marina Tanimura, Nagumo et Kato 2004 (Fig. 1)
Description
: Valves are 2.8-5.0 µm in diameter. The central area is flat to slightly tangentially undulate without colliculate ornamentation (
Fig. 1
A and
1
B). The marginal striae density is 20-25 in 10 µm. Valve face fultoportula is absent. Mantle fultoportulae located on every 2nd to 3rd interstria (
Fig. 1
A-
1
D), internally the tubulus is surrounded by two satellite pores (
Fig. 1
C and
1
D). A single rimoportula located on the ring of mantle fultoportulae, internally the rimoportula has a narrow sessile labium (
Fig. 1
C and
1
D).
Distribution
:
Cyclotella atomus
var.
marina
was found from a diatom assemblage attached to an eelgrass sample taken from waters with a salinity of approximately 26, in the region of Yulimri, Yeosu City, Korea in July 1998 (Chung et al. 2010). However, the species occurred total 5 times as a planktonic in this study and reported once and four times from the South Sea and the Yellow Sea, respectively (
Table 3
).
Remarks
:
Cyclotella atomus
var.
marina
is distinguished from other variety of
C. atomus
by the absence of a valve face fultoportula (Chung et al. 2010).
- Cyclotella baltica (Grunow) Håkansson 2002 (Fig. 2)
Basionym
:
Cyclotella striata
var.
baltica
Grunow in Van Heurck 1882, pl. 92, figs 13-15, non
Cyclotella litoralis
Lange et Syvertsen sensu Håkansson 1996.
Description
: Valves are 13.9-37.0 µm in diameter. The central area tangentially undulate with a slightly colliculate ornamentation (
Fig. 2
A-
2
C and
2
E). The marginal striae density is 10-15 in 10 µm. Valve face fultoportula number two to nine (
Fig. 2
C-
2
F), internally surrounded by three satellite pores (
Fig. 2
D and
2
F). Mantle fultoportulae located on every 2nd to 3rd interstria (
Fig. 2
G and
2
H),
Morphological information of 5Cyclotellaspecies in the present studyNA: not applicable.
Morphological information of 5 Cyclotella species in the present study NA: not applicable.
Cyclotella atomus var. marina Tanimura, Nagumo et Kato. Arrow and arrowheads indicate the rimoportula and the fultoportula respectively. (A-B)External valve view, (C-D) Internal valve view. Scale bars,1 μm.
internally the tubulus is surrounded by two satellite pores (
Fig. 2
H). A single rimoportula located slightly above the ring of the mantle fultoportulae, internally the rimoportula has a sessile labium (
Fig. 2
H).
Distribution
: Grunow (1882) collected
C. baltica
from the Baltic. Tanaka (2007) collected this species from Hokkaido to Kyushu in Lake Abshiri, Mikawa Bay, and regarded as a brackish to marine water species.
Cyclotella baltica
occurred total 11 times in this study, and was observed as a planktonic from the East Sea (one time) and the Yellow Sea (10 times), there was no occurrence from the South Sea (
Table 3
). This species is firstly recorded in the present study.
Remarks
: Håkansson (2002) placed
C. striata
var.
baltica
in
C. baltica
according to the absence of the valve face fultoportula, and stated that the species were regarded as
C. striata
previously (Hustedt 1928, Cleve-Euler 1951, Helmcke and Krieger 1954, Helmcke et al. 1974, Takano 1976, Prasad et al. 1990) was in fact
C. baltica
. Tanaka (2007) described
C. baltica
from Japanese specimens. However, the Japanese material showed some differences from the lectotype of Håkansson (2002): a valve diameter
Cyclotella baltica (Grunow) Håkansson. Arrow and arrowheads indicate the rimoportula and the fultoportula, respectively. (A) External valve view, (B)Internal valve view, (C) Central area with 7 fultoportulae in the external valve view, (D) Seven fultoportulae with 3 satellite pores on the central area of valveface in the internal valve view, (E) Externally striated marginal area with mantle fultoportulae and rimoportula, (F) Internally alveolated marginal area withmantle fultoportulae and rimoportula. Scale bars represent: A, B, 10 μm; C, D, 5 μm; E, 2 μm; G, H, 1 μm.
Check-list of 5Cyclotellaspecies in Korean coastal waters
Check-list of 5 Cyclotella species in Korean coastal waters
and the position of external opening of rimoportula. The morphological characteristics in Korean specimens agree with the description of
C. baltica
in Håkansson (2002): a valve diameter, the external position of rimoportula, thestriae dense and the spacing of mantle fultoportulae. Although Korean specimens show well-developed alveoli in the internal valve view (
Fig. 2
B and
2
F), the variation of alveoli development within
Cyclotella
species have been reported (Beszteri et al. 2005). Therefore, we regarded the Korean specimens as
C. baltica
.
- Cyclotella litoralis Lange et Syvertsen 1989 (Fig. 3)
Description
: Valves are 23.0-63.0 µm in diameter. The central area tangentially undulates with colliculate ornamentation (
Fig. 3
A-
3
C and
3
E). The marginal striae density is 8-12 in 10 µm. Valve face fultoportula is eight to twelve
Fig. 3
C and
3
D), internally surrounded by three satellite pores (
Fig. 3
F). Mantle fultoportulae located on every 2nd
Fig. 3
G and
3
H), or sometimes, on a pair (
Fig. 3
G), internally the tubulus is surrounded by two satellite pores
Fig. 3
H). A single rimoportula is located on the fultoportulae below the mantle, internally the rimoportula has a sessile labium
Fig. 3
H).
Distribution
: Lange and Syvertsen (1989) originally described as
C. litoralis
from the south western Atlantic, and Tanaka (2007) described this species in Isahaya Bay, Nagasaki Prefecture, Japan. This species was observed 29 times from all of the Sea as a planktonic in this study (
Table 3
). Although
C. litoralis
is firstly recorded in this study, Lee and Lee (1988) already observed
Cyclotella
sp. A from Gwangyang Bay and Jinhae Bay. However, since then there was no additional study for identification of the species.
Remarks
: Cyclotella litoralis have been confused with C. striata complex (e.g., C. baltica, C. striata, C. stylorum): This species have been distinguished from
C. baltica
based on the spacing of the mantle fultoportulae (Håkansson 2002), as well as the number of central fultoportula (Prasad and Nienow 2006). In this study, the presence of the recessed costa is an additional characteristic for distinguishing between the two species (
Fig 2
B and
3
B).
Cyclotella striata
can be distinguished from
C. litoralis
based on the absence of a valve face fultoportula (Håkansson 2002). The spacing of mantle fultoportula can be used to distinguish between
C. litoralis
and
C. stylorum
(Lange and Syvertsen 1989).
- Cyclotella meduanae Germain 1981 (Fig. 4)
Description
: Valves are 6.2-7.8 µm in diameter. The central area is flat to slightly tangentially undulate, without colliculate ornamentation (
Fig. 4
A and
4
B). The marginal striae density is 12-15 in 10 µm. Valve face fultoportula is absent (
Fig. 4
A-
4
C). Marginal fultoportulae are located on every 2nd to 3rd interstria (
Fig. 4
A-
4
D), internally the tubulus is surrounded by three satellite pores (Fig. 4D). A single rimoportula is located on the ring of marginal fultoportulae, internally the rimoportula has a narrow sessile labium (
Fig. 4
D).
Distribution
:
Cyclotella meduanae
originally described from Mayenne, France (Germain 1981). Tanaka (2007) examined the Japanese specimen from Inba Pond, Chiba Prefecture and mentioned that C. meduanae has been found mainly in eutrophic water and freshwater. In the present study,
C. meduanae
is newly recorded and was found 3 times as a planktonic (
Table 3
).
Remarks
:
Cyclotella meduanae
has been noted to have morphological similarity with
C. meneghiniana
(Håkansson 2002). Håkansson (2002) mentioned that the only difference between both species is the absence or presence of the valve face fultoportula. However, we found an additional difference in the spacing of the mantle fultoportulae between
C. meduanae
and
C. meneghiniana
: in the first one, the mantle fultoportulae was located on every second interstria, while the latter on every interstria (compare
Fig. 4
C with
Fig. 5
B).
Cyclotella meneghiniana
Kützing 1844 (
Fig. 5
)
Synonyms
:
Surirella melosiroides
G.G.A. Meneghini ms. 1844;
Cyclotella kuetzingiana
var.
meneghiniana
(Kützing) Brun 1880;
Stephanocyclus meneghiniana
(Kützing) Skabichevskii 1975.
Description
: Valves are 8.0-30.3 µm in diameter. The central area is slightly tangentially undulate and without colliculate ornamentation (
Fig. 5
A-
5
E). The marginal striae density is 8-10 in 10 µm. Valve face fultoportula is one to seven (
Fig. 5
A-
5
F), internally surrounded by three satellite pores (
Fig. 5
F). Marginal fultoportulae are located on every interstria (
Fig. 5
G and
5
H), internally the tubulus is surrounded by three satellite pores (
Fig. 5
H). A single rimoportula located on the ring of marginal fultoportulae, internally the rimoportula has a stalked, and often bent, sessile labium (
Fig. 5
H).
Distribution
:
Cyclotella meneghiniana
have been found in varied habitats including brackish water, and both eutrophic and oligotrophic freshwater (Håkansson 2002, Tanaka 2007). In Korea, Lee and Lee (1988) firstly recorded
C. meneghiniana
in the Han River and Ulsan Bay near Taehwa River estuary. Cho (1996) described
C. meneghiniana
from the Nakdong River.
Cyclotella meneghiniana
was most frequently observed in this study. This species
Cyclotella litoralis Lange et Syvertsen. Arrow indicates the rimoportula. (A) External valve view, (B) Internal valve view, (C) Central area with 10fultoportulae (arrowheads) in the external valve view, (D) Twelve fultoportulae (arrowheads) with 3 satellite pores on the central area of valve face in theinternal valve view, (E) Externally striated marginal area with mantle fultoportulae (arrowheads) and rimoportula, (F) Internally marginal area with mantlefultoportulae (arrowheads) on the recessed interstria and a sessile rimoportula between two fultoportulae. Scale bars represent: A-D, 10 μm; E, 1 μm; F, 5μm; G, H, 1 μm.
Cyclotella meduanae Germain. Arrow indicates the rimoportula. (A-B) External valve view, (C) Internal valve view, (D) Internal mantle fultoportulaeon every interstria and sessile rimoportula between two mantle fultoportulae. Scale bars represent: A-C, 2 μm; D, 1 μm.
was reported total 30 times in this study, 8 times from the East Sea, 2 times from the South Sea, and 20 times from the Yellow Sea as a planktonic (
Table 3
).
Remarks
:
Cyclotella meneghiniana
has been known to have a wide ecological distribution (Håkansson 2002, Finlay et al. 2002), and highly variable frustule (Håkansson and Chepurnov 1999).
Therefore, this species has been confused with several
Cyclotella
species such as
C. ceylonica
Holsinger,
C. pratii
Toman,
C. kuetzingiana
Thwaites (Håkansson 2002). Recently, Beszteri et al. (2007) investigated the cryptic diversity of
C. meneghiniana
based on nuclear and plastid genes. Although this study did not analyze the molecular sequences, it is needed to understand the cryptic diversity of
C. meneghiniana
in Korea.
Occurrence of Cyclotella species according to the salinity
This study present the salinity range of five
Cyclotella
Cyclotella meneghiniana Kützing. Arrow indicates the rimoportula. (A) External valve view, (B) Internal valve view, (C) Central area with 6fultoportulae in the external valve view, (D) Two fultoportulae with 3 satellite pores on the central area of valve face in the internal valve view, (E) Externallystriated marginal area with mantle fultoportulae, (F) Internally marginal area with mantle fultoportulae on the interstria and a sessile rimoportula betweentwo fultoportulae. Scale bars represent: A, B, 10 μm; C, D, 5 μm; E-H, 1 μm.Cyclotella meneghiniana Kützing. Arrow indicates the rimoportula. (A) External valve view, (B) Internal valve view, (C) Central area with 6fultoportulae in the external valve view, (D) Two fultoportulae with 3 satellite pores on the central area of valve face in the internal valve view, (E) Externallystriated marginal area with mantle fultoportulae, (F) Internally marginal area with mantle fultoportulae on the interstria and a sessile rimoportula betweentwo fultoportulae. Scale bars represent: A, B, 10 μm; C, D, 5 μm; E-H, 1 μm.
Box plot of data of salinity range from five Cyclotella species inKorea.
species in the Korean coastal waters (
Fig. 6
).
Cyclotella atomus
var.
marina
occurred in a range of 10.17-28.01 psu from five sites;
C. baltica
was collected at 11 sites in a range of 10.17-31.73 psu;
C. litoralis
was reported in a range of 15.32-34.80 psu at 29 sites;
C. meduanae
have a salinity range of 10.17-22.70 psu from 3 sites;
C. meneghiniana
occurred in a range of 0.04-34.95 psu from 30 sites. Most of species occur a wide range of 10-30 psu, which indicate the leakage of
Cyclotella
species into the coastal waters may be more diverse than previous consideration. (Round and Sims 1980).
Acknowledgements
This work was supported by a grant from the National Institute of Biological Resources (NIBR), as funded by the Ministry of Environment (MOE) of Republic of Korea (NIBR No. 2013-01-001), and by a 2013 Research Grant from Sangmyung Universtiy.
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