Author's personal copy
ARTICLE IN PRESS
European Journal of
PROTISTOLOGY
European Journal of Protistology 45 (2009) 29–37
www.elsevier.de/ejop
Descriptions of two new marine scuticociliates, Pleuronema sinica n. sp.
and P. wilberti n. sp. (Ciliophora: Scuticociliatida), from the
Yellow Sea, China
Yangang Wanga, Weibo Songa,, Alan Warrenb, Khaled A.S. Al-Rasheidc,
Saleh A. Al-Quraishyc, Saleh A. Al-Farrajc, Xiaozhong Hua, Hongbo Pana
a
Laboratory of Protozoology, Ocean University of China, Qingdao 266003, PR China
Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK
c
Zoology Department, King Saud University, P.O. Box 2455, Riyadh 11451, Saudi Arabia
b
Received 10 March 2008; received in revised form 23 May 2008; accepted 17 June 2008
Abstract
Two new marine scuticociliates, Pleuronema sinica n. sp. and P. wilberti n. sp., collected from the sand beach of
Qingdao, China, were investigated in vivo and following protargol impregnation. Ciliates of the genus Pleuronema are
normally recognizable by their large sail-like paroral membrane although one species, P. grolierei, has shorter cilia in
the paroral membrane. Neither of the new forms has the conspicuous paroral membrane in vivo so in this respect they
are not typical members of this genus. Pleuronema sinica is characterized by its large, conspicuously flattened body, the
possession of only one preoral kinety, the irregular-shaped macronucleus and the rather unusual structure of the oral
apparatus. By contrast P. wilberti has a medium-size broad-oval body, six to eight preoral kineties and a highly
differentiated membranelle 3 that is five- or six-rowed. An identification key is supplied for the 15 species of
Pleuronema for which the infraciliature is known.
r 2008 Elsevier GmbH. All rights reserved.
Keywords: Identification key; Interstitial ciliates; Morphology; Pleuronematidae; Silverstain; Taxonomy
Introduction
Free-living scuticociliates are common inhabitants
of littoral and sandy habitats, especially in sediments,
where they may reach high abundances (Agamaliev 1968;
Al-Rasheid 2001; Borror 1963; Kahl 1926; Song 2000).
Pleuronema is a species-rich genus in the Scuticociliatida
with more than 20 morphotypes having been reported
from various geographical locations (Agatha et al. 1993;
Corliss and Snyder 1986; Czapik and Jordan 1977;
Corresponding author.
E-mail address: wsong@ouc.edu.cn (W. Song).
0932-4739/$ - see front matter r 2008 Elsevier GmbH. All rights reserved.
doi:10.1016/j.ejop.2008.06.001
Dragesco 1960, 1968; Dragesco and Dragesco-Kernéis
1986; Fernandez-Leborans and Novillo 1994; Grolière
and Detcheva 1974).
Since many marine forms of Pleuronema have similar
living morphologies, a combination of in vivo observation
and details of the infraciliature, especially with respect to
the oral apparatus, is needed for accurate species
identification and circumscription. However, such data
are only available for a few morphotypes, indicating that
the majority of species of Pleuronema need to be reinvestigated using modern methods (Wang et al. 2008a, b).
In the present paper, we describe two new species of
the genus Pleuronema, with emphasis on their living
Author's personal copy
ARTICLE IN PRESS
30
Y. Wang et al. / European Journal of Protistology 45 (2009) 29–37
morphology, infraciliature and silverline systems, and
supply a key for the identification of those forms for
which the infraciliature is known.
Materials and methods
Pleuronema sinica was collected from a sandy beach
quite near to a sewage outfall (water temperature
12.9 1C and salinity 27.9%) on 11 December 2005 in
Qingdao (361080 N; 1201430 E), China.
Pleuronema wilberti was also collected from a sandy
beach in Qingdao (361080 N; 1201430 E), China (water
temperature 11.2 1C and salinity 31%).
Samples were collected by digging a hole about 15 cm
deep in the sand, waiting for seawater to seep into the
hole, then taking the seawater with some sand back to
the laboratory. Samples were processed within 2 h of
collection. Isolations were carried out using a micropipette. Attempts to culture the two species failed, therefore
all studies were carried out on freshly isolated specimens. Individuals were observed in vivo using differential interference contrast microscopy. Protargol
(Wilbert 1975) and Chatton–Lwoff (Song and Wilbert
1995) methods were used to reveal the infraciliature and
silverline systems, respectively. Drawings of impregnated specimens were made with the help of a camera
lucida; measurements were performed under 100 to
1250 magnification. Systematics and terminology are
mainly according to Corliss (1979).
Results
Pleuronema sinica n. sp. (Figs 1–11, 22–33; Table 1)
Diagnosis: Body size in vivo 105–200 45–105 mm,
dorso-ventrally flattened about 4:3; several prolonged
caudal cilia; one preoral kinety and about 45 somatic
kineties; membranelle 1 (M1) about 15% of M2a in
Figs 1–11. Pleuronema sinica n. sp., from life (1–5), after protargol impregnation (6–8, 10, 11) and after Chatton–Lwoff
impregnation (9). 1. Ventral view of typical individual. 2. Swimming trace. 3. Ventral view of irregular cell. 4. Lateral view from left
side. 5. Dorsal view of typical individual. 6, 7. Structure of oral apparatus, arrows in 6 show the two-rowed sections of M2a, doublearrowhead in 7 points to the separated section of M2b, arrowhead points to the short row of kinetosomes in membranelle 1.
8. Different shapes of the irregular macronucleus. 9. Oral apparatus, arrow denotes the cytostome. 10, 11. Ventral and dorsal views
of the same specimen, showing the infraciliature. M1-3, membranelles 1–3; M2a, the upper part of membranelle 2; M2b, the lower
part of membranelle 2. Scale bars ¼ 50 mm.
Author's personal copy
ARTICLE IN PRESS
Y. Wang et al. / European Journal of Protistology 45 (2009) 29–37
Table 1.
31
Morphometric data for Pleuronema sinica n. sp. (upper line) and P. wilberti n. sp. (lower line)
Characters
Min
Max
Mean
SD
CV
n
Body length
101
89
199
138
143.5
110.0
27.5
13.6
19.1
12.4
20
24
Body width
42
36
102
78
67.1
54.1
15.1
9.8
22.5
18.0
20
24
Length of buccal field
66
39
162
90
93.3
71.6
20.1
10.6
21.5
14.8
19
24
Width of buccal field
15
19
29
33
20.0
26.2
4.5
3.5
22.6
13.2
13
23
Number of somatic kineties
41
40
52
49
45.5
45.0
2.5
2.6
5.4
5.7
21
26
Number of preoral kineties
1
6
1
8
1.0
6.9
0
0.5
0
7.0
17
26
Number of rows in M3
3
5
3
6
3.0
5.4
0
0.5
0
9.1
21
26
Measurements in mm.
Data based on protargol impregnated specimens. CV, coefficient of variation in %; Max, maximum; Mean, arithmetic mean; Min, minimum;
n, number of specimens measured; SD, standard deviation.
length, consisting of three rows of kinetosomes; M2a
extremely long with straight posterior end; M3 threerowed; contractile vacuole caudally positioned; one
irregular-shaped macronucleus; marine habitat.
Type location: A sandy beach in Qingdao (361080 N;
1201430 E), China.
Type specimens: One holotype slide with protargolimpregnated specimens is deposited in the Natural
History Museum, London, UK with registration no.
2007:5:16:4. One paratype slide (no. wyg-2005-12-11-01)
is deposited in the Laboratory of Protozoology, Ocean
University of China, China.
Etymology: The name sinica recalls the fact that this
species was first described from Chinese coastal waters.
Description: Cells in vivo about 105–200 45–105 mm,
elongate to slender elliptical in outline, broadly rounded
at both ends (Figs 1, 3–5, 22–24, 27). Dorso-ventrally
flattened about 4:3. In lateral view, body often slightly
narrowed at level of cytostome. Buccal field about 60%
of body length and 33% of body width (Fig. 10).
Cytoplasm often black at anterior part and slightly
greyish at posterior part, packed with many lemonshaped granules (mostly 6–8 4–5 mm) (Figs 1, 3–5,
22–24). Macronucleus irregular and variable in shape,
usually with many globular nucleoli (Figs 8, 30).
Contractile vacuole near posterior end, about 8 mm in
diameter (Figs 1, 5).
Cilia of membranelles 1 and 2 and anterior part of
paroral membrane undetectable while those of posterior
part of paroral membrane are long (about 20 mm) and
radiate conspicuously away from body (Figs 1, 3–5, 22,
24, 31). Somatic cilia usually about 10 mm long; ca. ten
prolonged caudal cilia, about 20 mm long, always
projecting radially from cell (Figs 1, 3–5, 27). Movement
moderately fast, rotating about main body axis, sometimes motionless for short periods (Fig. 2).
With 41–52 somatic kineties that terminate at the
anterior end of the cell forming a small, bald apical
plate. Somatic kineties composed of dikinetids in
anterior half of the body and monokinetids in posterior
part (Figs 10, 11). Invariably one preoral kinety to left
of buccal field (Figs 6, 7, 10, 26).
Within oral apparatus, membranelle 1 (M1) consists
of one short and two longer rows of basal bodies (Figs 6,
7, 10, 25); upper part of membranelle 2 (M2a) straight at
posterior end, anterior and posterior sections tworowed, mid-portion single-rowed (Figs 6, 7, 10, 32);
M2b irregularly V-shaped, in most specimens right
branch is fragmented and about twice length of left
branch (Figs 6, 9, 32, 33), although in others right
branch is more than 3 length of left branch (Figs 7,
26); membranelle 3 (M3) three-rowed; paroral membrane (PM) about 60% cell length (Fig. 10).
Silverline system typical of genus (Figs 28, 29, 33).
Comparison: Considering the general morphology,
habitat and especially the oral structure, four marine
congeners of the marinum-type (i.e. with posterior end of
M2a straight or only slightly curved) should be
compared with Pleuronema sinica (Wang et al. 2008a):
namely Pleuronema tardum Czapik and Jordan, 1977,
P. marinum Dujardin, 1841, P. oculata Dragesco, 1960
and P. grolierei Wang et al., 2008.
Author's personal copy
ARTICLE IN PRESS
32
Y. Wang et al. / European Journal of Protistology 45 (2009) 29–37
Considering the inconspicuous paroral membrane,
Pleuronema grolierei is similar to P. sinica, although the
former can be clearly distinguished from the latter by its
smaller body size (70–80 mm vs. 105–200 mm), fewer
somatic kineties (24–32 vs. 41–52), fewer rows of basal
bodies in M3 (2 vs. 3), longer two-rowed section of M2a
(entirely double-rowed vs. double-rowed only anteriorly
and posteriorly) and the different ratios of M2a/M3 (2:1
vs. 6:1) (Wang et al. 2008a).
Based on the description by Borror (1963), Pleuronema
marinum (Figs 50, 62) can be separated from
P. sinica by having a conspicuous sail-like paroral
membrane in vivo (vs. paroral membrane inconspicuous
and not sail-like in P. sinica), a different appearance of
the M2b, the right branch being about the same length
as the left branch (vs. right branch 2 length of
left branch in P. sinica), four preoral kineties (vs. one in
P. sinica) and a spherical macronucleus (vs. irregular in
P. sinica) (Borror 1963).
Pleuronema oculata (Figs 48, 56) differs from P. sinica
in having a conspicuous, sail-like paroral membrane in
vivo (vs. paroral membrane inconspicuous and not
sail-like in P. sinica), four preoral kineties (vs. one in
P. sinica), a spherical macronucleus (vs. irregular in
P. sinica), and the structure of the M2b (straight vs.
V-shaped in P. sinica) and the M3 (two-rowed vs. threerowed in P. sinica) (Dragesco 1960).
Although its living morphology has yet to be
described, Pleuronema tardum (Figs 54, 61) can be
distinguished from P. sinica by having more macronuclear nodules (six or seven vs. invariably one in
P. sinica), fewer kineties in M1 (two vs. three), dikinetids
along the entire length of the somatic kineties
(vs. dikinetids only in anterior half of somatic kineties
in P. sinica) and the appearance of M2b with the right
branch 3 to 4 the length of the left branch (vs. 2
in P. sinica) (Czapik and Jordan 1977).
Pleuronema wilberti n. sp. (Figs 12–21, 34–47; Table 1)
Diagnosis: Size in vivo 85–140 40–80 mm with
elongated oval body shape; about 15 prolonged caudal
cilia; six to eight preoral kineties and ca. 45 somatic
Figs 12–21. Pleuronema wilberti n. sp. in vivo (12, 15–17), after protargol impregnation (13, 19–21) and after Chatton–Lwoff
impregnation (14, 18). 12, 16, 17. Ventral view to show different body sizes and shapes. 13. Different appearances of the
macronuclear apparatus, arrow indicates the small spherical macronuclear segments. 14. Silverline system. 15. Swimming trace.
18. Structure of oral apparatus, arrow marks membranelle 3. 19. Infraciliature of oral apparatus, arrow points to the hook-like
structure of M2a. 20, 21. Ventral and dorsal views of the same specimen, showing infraciliature, arrowheads in 20 denote the preoral
kineties. Scale bars ¼ 50 mm.
Author's personal copy
ARTICLE IN PRESS
Y. Wang et al. / European Journal of Protistology 45 (2009) 29–37
33
Figs 22–33. Pleuronema sinica in vivo (22–24, 27, 28, 31), after protargol impregnation (25, 26, 30, 32) and after Chatton–Lwoff
impregnation (29, 33). 22. Right-ventral view of a typical individual, arrow marks a forming food vacuole. 23. Lateral view from left
side. 24, 27. To show different body shapes, arrowheads in 24 and 27 mark cilia of paroral membrane and caudal cilia respectively.
25. Anterior part of oral apparatus, arrow shows the anterior end of M2a, double-arrowhead marks membranelle 1. 26. Posterior
part of oral apparatus, arrow indicates M2b, double-arrowhead refers to the separate section of M2b, single arrowhead points to
preoral kinety. 28, 29. Surface of the cortex from life (28) and after Chatton–Lwoff impregnation (29), arrows in 28 show the
hexagonal honeycomb structure. 30. Shape of macronucleus. 31. Lateral view of the posterior part, arrow marks cilia of paroral
membrane. 32. Detailed view of oral apparatus, arrowhead marks membranelle 3, arrow shows the straight end of M2a, doublearrowhead points to the preoral kinety. 33. Silverline system of oral apparatus. Ma, macronucleus. Scale bars ¼ 100 mm.
kineties; M2a about the same length as M1, with
hook-like posterior end; M3 is five- or six-rowed;
contractile vacuole caudally positioned; one spherical
macronucleus; marine habitat.
Type location: A sandy beach in Qingdao (361080 N;
1201430 E), China.
Type specimens: One holotype slide with protargolimpregnated specimens is deposited in the Natural
History Museum, London, UK with registration no.
2007:5:16:3. One paratype (no. wyg-2006-05-01-02) is
deposited in the Laboratory of Protozoology, Ocean
University of China, PR China.
Dedication: The species is dedicated to Professor
Norbert Wilbert, Institut für Zoologie der Universität
Bonn, Germany, in recognition of his many contributions to our knowledge of this group of ciliates.
Description: Size range 85–140 40–80 mm in vivo,
elongate to slender oval in outline, broadly rounded at
both ends, gradually narrowing from level of cytostome
to posterior end of cell (Figs 12, 16, 17, 34–37). Dorsoventrally flattened about 3:2. Oral field large, extending
to ca. 65% of body length and 50% of body width
(Figs 12, 20). Cytoplasm slightly greyish, packed with
many greasily shining globules (mostly 1–3 mm across)
(Figs 34, 37). Food vacuoles usually large, filled with
diatoms (Figs 12, 17, 35). One spherical macronucleus,
usually with many globular nucleoli, although in some
specimens several small spherical macronuclear segments were observed (Fig. 13). Contractile vacuole
located sub-terminally in slightly dorsal position, about
10 mm in diameter (Figs 12, 17).
Cilia of both paroral membrane and membranelles
about 40 mm long, not forming a sail-like structure
(Figs 12, 17, 34), although sometimes cilia of M2a, M1
and anterior of PM extend radially away from body
somewhat (Figs 16, 36, 37). Somatic cilia usually about
Author's personal copy
ARTICLE IN PRESS
34
Y. Wang et al. / European Journal of Protistology 45 (2009) 29–37
Figs 34–47. Pleuronema wilberti from life (34–37), after protargol impregnation (38–45) and after Chatton–Lwoff impregnation
(46, 47). 34. Left-ventral view of a typical individual. 35–37. To show different body sizes and shapes, arrow in 35 marks an ingested
diatom, arrows in 36 show the prolonged caudal cilia, arrow in 37 indicates cilia of paroral membrane. 38. Infraciliature of anterior
part of cell, arrow refers to membranelle 1, arrowheads point to the preoral suture. 39, 40, 42–45. Ventral view of posterior part of
oral apparatus, arrow in 39 marks M2b, double-arrowhead in 39 shows the hook-like structure of M2a, arrows in 40 indicate
membranelle 3, arrow in 42 points to membranelle 3, arrowheads in 43 refer to the preoral kineties, arrowhead and arrow in 44 mark
membranelle 3 and M2b respectively, arrows and arrowheads in 45 point to membranelle 3 and micronuclei respectively. 41. Oral
apparatus, arrow refers to the hook-like structure of M2a. 46. Posterior part of oral apparatus, arrows show the cytostome,
arrowheads point to the oral ribs. 47. Silverline system, arrows mark preoral kineties, arrowheads indicate silverline system. Scale
bars ¼ 50 mm.
10 mm long; on average ca. 15 prolonged caudal cilia
each about 24 mm long, invariably extending radially
away from body (Figs 12, 16, 17, 37). Movement
moderately fast, rotating about main body axis,
motionless for short periods (Fig. 15).
With 40–49 somatic kineties that terminate at
anterior end of cell forming a small, bald apical plate
(Figs 20, 38). Small glabrous area posterior of buccal field
(Figs 20, 39). Somatic kineties composed of dikinetids in
anterior three quarters of body and monokinetids in
posterior quarter (Figs 20, 21). Six to eight preoral
kineties to left of buccal field (Figs 19, 20, 43, 47).
Oral apparatus generally genus-typical: M1 tworowed, long and prominent, about the same length as
M2a (Figs 13, 18, 19, 20, 41); M2a two-rowed with
hooked posterior end (Figs 19, 20, 41); M2b V-shaped,
with kinetosomes arranged in linear pattern, except for a
short zig-zag pattern in middle section (Figs 18, 19, 20,
39–44); M3 five- or six-rowed (Figs 19, 20, 40–42,
44–46); paroral membrane (PM) prominent, about 75%
of cell length with two rows of basal bodies in zig-zag
pattern (Fig. 20).
Silverline system with a hexagonal honeycomb
pattern (Figs 14, 47).
Comparison: In terms of the body shape and size,
macronucleus, oral apparatus and habitat, the following
species, all of which are of the coronatum-type (i.e.
posterior end of M2a is conspicuously hook-shaped)
(Wang et al. 2008a), should be compared with Pleuronema wilberti: P. coronatum Kent, 1881; P. salmastra
Author's personal copy
ARTICLE IN PRESS
Y. Wang et al. / European Journal of Protistology 45 (2009) 29–37
35
Figs 48–63. Some species of Pleuronema that are similar to P. wilberti and P. sinica. 48, 56. P. oculata (from Dragesco 1960).
49, 51. P. salmastra (from Dragesco and Dragesco-Kernéis 1986). 50, 62. P. marinum (from Borror 1963). 52. P. lynni
(from Fernandez-Leborans and Novillo 1994). 53. P. glaciale (from Corliss and Snyder 1986). 54, 61. P. tardum (from Czapik
and Jordan 1977). 55, 59. P. arctica (from Agatha et al. 1993). 57, 60, 63. P. coronatum (from Song 2000; Wang et al. 2008a).
58. P. puytoraci (from Grolière and Detcheva 1974).
Dragesco, 1968; P. arctica Agatha et al., 1993;
P. glaciale Corliss and Snyder, 1986; P. lynni Fernandez-Leborans and Novillo, 1994 and P. puytoraci
Grolière and Detcheva, 1974.
Pleuronema coronatum (Figs 57, 60, 63) differs from
P. wilberti in the following: having a conspicuously saillike paroral membrane in vivo (vs. paroral membrane
not sail-like in P. wilberti), a ratio of M1:M2a lengths of
around 1:4 (vs. 1:1 in P. wilberti), an M2a which is
double-rowed only in anterior and posterior regions (vs.
entirely double-rowed in P. wilberti), and an M3 which
is three-rowed (vs. five- or six-rowed in P. wilberti)
(Song 2000; Wang et al. 2008a).
Pleuronema salmastra (Figs 49, 51) can be easily
distinguished from P. wilberti by having a conspicuous
sail-like paroral membrane in vivo (vs. paroral membrane not sail-like in P. wilberti), the structure of M3
(three-rowed vs. five- or six-rowed in P. wilberti),
the structure of M2a (double-rowed only in anterior
and posterior sections vs. entirely double-rowed in
P. wilberti) and the ratio of M1:M2a lengths (1:5 vs. 1:1)
(Dragesco and Dragesco-Kernéis 1986).
Although P. arctica, P. glaciale, P. lynni and
P. puytoraci have not yet been described in vivo, they
can be separated from P. wilberti based on silverimpregnated specimens.
Pleuronema arctica (Figs 55, 59) differs from
P. wilberti by its larger body size after silver impregnation
(131–242 vs. 89–138 mm in P. wilberti), the ratio of
M1:M2a lengths (1:5 vs. 1:1 in P. wilberti), the structure
of M2a (double-rowed only in the anterior and posterior
regions vs. entirely double-rowed in P. wilberti) and the
structure of M3 (three-rowed vs. five- or six-rowed in
P. wilberti) (Agatha et al. 1993).
Pleuronema glaciale (Fig. 53) can be distinguished
from P. wilberti by the ratio of M1:M2a length (1:2 vs.
1:1 in P. wilberti), the structure of M3 (two-rowed vs.
five- or six-rowed in P. wilberti) and the oral field which
is conspicuously large and almost as long as the body
(vs. about 75% of the body length in P. wilberti) (Corliss
and Snyder 1986).
Pleuronema lynni (Fig. 52) and P. puytoraci (Fig. 58)
can be separated from P. wilberti by the number of
somatic kineties (34–40 and 28 respectively vs. 40–49 in
P. wilberti), the ratio of M1:M2a length (1:6 and 1:5
respectively vs. 1:1 in P. wilberti), and the structure of
M3 (three-rowed vs. five- or six-rowed in P. wilberti).
Furthermore, in P. lynni the M2b is curved (vs.
V-shaped in P. wilberti) (Grolière and Detcheva 1974;
Fernandez-Leborans and Novillo 1994).
Key to the identification of selected Pleuronema spp
There are 15 species of Pleuronema whose infraciliature is known (see: Agatha et al. 1993; Borror 1963;
Corliss and Snyder 1986; Czapik and Jordan 1977;
Dragesco 1960; Dragesco and Dragesco-Kernéis 1986;
Fernandez-Leborans and Novillo 1994; Grolière and
Detcheva 1974; Wang et al. 2008a, b). A key to their
identification is provided here:
Author's personal copy
ARTICLE IN PRESS
36
Y. Wang et al. / European Journal of Protistology 45 (2009) 29–37
1
10
100
2
M3 five- or six-rowed
M3 two-rowed
M3 three-rowed
PM conspicuous; contractile vacuole located dorsally; three preoral
kineties
PM inconspicuous; contractile vacuole located ventrally; one preoral
kinety
Several macronuclear segments
One macronucleus
Posterior end of M2a straight; 40–50 somatic kineties; one preoral
kinety
Posterior end of M2a hook-like; 25–39 somatic kineties; three to five
preoral kineties
Macronucleus sausage-shaped
Macronucleus irregular
Macronucleus spherical
M1 extremely long, about half length of M2a
M1 normally short, conspicuously less than half length of M2a
M2b straight or slightly curved
M2b V-shaped
Ratio of M1:M2a length about 1:6; 34–40 somatic kineties; ca. 7
preoral kineties; M2b slightly curved
Ratio of M1:M2a length about 1:3; 50 somatic kineties; ca. 4 preoral
kineties; M2b straight
Posterior end of M2a straight
Posterior end of M2a hook-shaped
M2a almost completely double-rowed
M2a double-rowed only in anterior and posterior regions
About 28 somatic kineties
X40 somatic kineties
About 40 somatic kineties
X53 somatic kineties
20
3
30
4
40
5
50
500
6
60
7
70
8
80
9
90
10
100
11
110
12
120
Acknowledgements
This work was supported by the National Science
Foundation of China (Project no. 40676076), the
Darwin Initiative Program (Project no. 14-015) which
is funded by the UK Department for Environment and
Food and Rural Affairs and a grant from the Center of
Excellence in Biodiversity, King Saud University.
Thanks are due to Jiamei Jiang, Hongan Long and
Xiangrui Chen (Ocean University of China) for their
help in sampling.
References
Agamaliev, F.G., 1968. Materials on morphology of some
psammophilic ciliates of the Caspian Sea. Acta Protozool.
6, 225–244 (in Russian with English summary).
Agatha, S., Spindler, M., Wilbert, N., 1993. Ciliated protozoa
(Ciliophora) from Arctic sea ice. Acta Protozool. 32,
261–268.
P. wilberti
2
3
P. arenicola
P. grolierei
4
5
P. tardum
P. czapikae
P.
P.
6
P.
7
8
9
P.
wiackowskii
sinica
glaciale
lynni
P. oculata
P. marinum
10
P. arctica
11
P. puytoraci
12
P. coronatum
P. salmastra
Al-Rasheid, K.A.S., 2001. New records of interstitial ciliates
(Protozoa, Ciliophora) from the Saudi coasts of the Red
Sea. Trop. Zool. 14, 133–156.
Borror, A.C., 1963. Morphology and ecology of the benthic
ciliated protozoa of Alligator Harbour, Florida. Arch.
Protistenkd. 106, 465–534.
Corliss, J.O., 1979. The Ciliated Protozoa: Characterization,
Classification and Guide to the Literature. Pergamon Press,
Oxford, New York, Toronto, Sydney, Paris, Frankfurt.
Corliss, J.O., Snyder, R.A., 1986. A preliminary description of
several new ciliates from the Antarctica, including Cohnilembus grassei n. sp. (1). Protistologica 22, 39–46.
Czapik, A., Jordan, A., 1977. Two new psammobiotic ciliates
Hippocomos loricatus n. gen. n. sp. and Pleuronema tardum
n. sp. (Hymenostomata, Pleuronematina). Acta Protozool.
16, 157–164.
Dragesco, J., 1960. Ciliés mésopsammiques littoraux. systématique, morphologie, écologie. Trav. Stat. Biol. Roscoff. 122,
1–356.
Dragesco, J., 1968. Les genres Pleuronema Dujardin, Schizocalyptra nov. gen. et Histiobalantium Stokes (Ciliés,
Holotriches, Hyménostomes). Protistologica 4, 85–106.
Author's personal copy
ARTICLE IN PRESS
Y. Wang et al. / European Journal of Protistology 45 (2009) 29–37
Dragesco, J., Dragesco-Kernéis, A., 1986. Ciliés libres de
l’Afrique intertropicale. Faune Trop. 26, 1–559.
Fernandez-Leborans, G., Novillo, A., 1994. Morphology and
taxonomic position of two marine pleuronematine species:
Pleuronema lynni and Schizocalyptra marina (Protozoa,
Ciliophora). J. Zool. London 233, 259–275.
Grolière, C.A., Detcheva, R., 1974. Description et stomatogenèse de Pleuronema puytoraci n. sp. (Ciliata, Holotricha).
Protistologica 10, 91–99.
Kahl, A., 1926. Neue und wenig bekannte Formen der
Holotrichen und Heterotrichen. Arch. Protistenkd. 55,
179–448.
Song, W., 2000. Morphology and taxonomical studies on some
marine scuticociliates from China sea, with description of
two new species, Philasterides armatalis sp. n. and
Cyclidium varibonneti sp. n. (Protozoa: Ciliophora: Scuticociliatida). Acta Protozool. 39, 295–322.
37
Song, W., Wilbert, N., 1995. Benthische Ciliaten des Süßwassers. In: Röttger, R. (Ed.), Praktikum der Protozoologie.
Gustav Fischer Verlag, New York, pp. 156–168.
Wang, Y., Hu, X., Long, H., Al-Rasheid, A.S., Al-Farraj,
S.A., Song, W., 2008a. Morphological studies indicate that
Pleuronema grolierei nov. spec. and P. coronatum Kent,
1881 represent different sections of the genus Pleuronema
(Ciliophora: Scuticociliatida). Eur. J. Protistol. 44,
131–140.
Wang, Y., Song, W., Warren, A., Al-Rasheid, A.S., Hu, X.,
2008b. Descriptions of two new marine species of Pleuronema, P. czapikae sp. n. and P. wiackowskii sp. n.
(Ciliophora: Scuticociliatida), from the Yellow Sea, North
China. Acta Protozool. 47, 35–45.
Wilbert, N., 1975. Eine verbesserte Technik der Protargolimprägnation für Ciliaten. Mikrokosmos 64, 171–179.