全 文 ：Received 26 Aug. 2003 Accepted 10 Feb. 2004
Supported by the National Natural Science Foundation of China (30270119，39899400).
* Author for correspondence. E-mail: .
http://www.chineseplantscience.com
Acta Botanica Sinica
植 物 学 报 2004, 46 (8): 883-888
Taxonomy of the Lemaneaceae (Batrachospermales, Rhodophyta) in China
XIE Shu-Lian1*, SHI Zhi-Xin2, WANG Ruo-Nan3
(1. School of Life Sciences and Technology, Shanxi University, Taiyuan 030006, China;
2. Institute of Hydrobiology, The Chinese Academy of Sciences, Wuhan 430072, China;
3. Department of Biology, Yunnan University, Kunming 650091, China)
Abstract: Six species, belonging to two genera of Lemaneaceae in China are described. They are
Lemanea Bory (including L. sinica Jao, L. crassa S. L. Xie et Z. X. Shi, L. ramosa S. L. Xie et Z. X. Shi and L.
simplex Jao) and Paralemanea (Silva) Vis et Sheath (including P. catenata (Kützing) Vis et Sheath and P.
parvula (Sirodot) S. L. Xie et Z. X. Shi). Among them L. crassa S. L. Xie et Z. X. Shi and L. ramosa S. L. Xie
et Z. X. Shi are new species. Moreover, Paralemanea (Silva) Vis et Sheath and P. catenata (Kützing) Vis et
Sheath, P. parvula (Sirodot) S. L. Xie et Z. X. Shi are newly recorded in China.
Key words: Lemaneaceae; Lemanea ; Paralemanea ; new species; new record; China
Lemaneaceae was established in 1828 by Agardh
(Agardh, 1928). The family, like other families of the order
Batrachospermales, has a heteromorphic triphasic life his-
tory consisting of chantransia, gametophyte and
carposporophyte phase. The plant is pseudoparenchyma-
tous with fascicles of cells that become progressively
smaller towards the exterior, with large vacuoles, occupy-
ing most of the cell volume in the cells towards the interior.
Medullar area with scattered ray cells (periaxial cells) is not
obviously continuous with the cortical filaments; ray cell
and inner fascicle cell are with small chloroplasts and con-
siderable vacuolation. Reproduction is similar to other
Batrachospermales but with carposporangia produced in
chains. Spermatangial sori is in discrete patches or rings.
Species of the family Lemaneaceae occur in clean
streams. They mainly distribute in the northern hemisphere
(Sirodot, 1872; Atkinson, 1890; Hamel, 1925; Israelson , 1942;
Jao, 1941; Palmer, 1941; Khan, 1973; Vis and Sheath, 1992)
and there are a few records from southern South America
(Necchi and Zucchi, 1995).
Type genus: Lemanea Bory.
There had been different views of taxonomical classifi-
cation on the family. In 1872, Sirodot divided Lemanea Bory
into two genus according to its morphological and ana-
tomical characteristics (Sirodot, 1872). He proposed that
the genus Sacheria includes those species that are devoid
of axial cortication and possess T- or L-shaped ray cells
closely adjacent to the outer cortex, and the genus Lemanea
consists of those species with axial cortication and simple
ray cells not abutting the outer cortex. Unfortunately, the
type of Sacheria Sirodot had assigned was L. corallina
Bory, as which Bory had already designated a type of
Lemanea (Bory, 1808). According to The International
Code Botanical Nomenclature, the name Sacheria is ille-
gitimate (Greuter, 1994). Later Ketel reduced Sacheria to
the rank of subgenus, and thereby recognized the co-ordi-
nate subgenus Lemanea (Ketel, 1887) and this idea was
taken supportedly (Atkinson, 1890). Hamel (1925) renamed
the latter as Eulemanea, and this arrangement had been
followed by subsequent researchers, however, it is also
illegitimate. Silva (1959) proposed that the subgenus
Lemanea includes those species that lack axial cortication
and the subgenus Paralemanea includes those species
that have axial cortication. Vis and Sheath (1992) changed
the two subgenera into genera. Sheath et al. (1996) con-
firmed this viewpoint after making a further research on the
morphology, ultrastructure and classification of genera in
Lemaneaceae.
In this paper, the genus and species found in China,
belonging to Lemaneaceae, are described.
1 Materials and Methods
Six species of Lemaneaceae were collected. They are
Lemanea sinica Jao, L. crassa S. L. Xie et Z. X. Shi, L.
rumosa S. L. Xie et Z. X. Shi, L. simplex Jao, Paralemanea
catenata (Kützing) Vis et Sheath and P. parvula (Sirdot) S.
L. Xie et Z. X. Shi.
Specimens were examined for the morphological char-
acteristics under an Olympus BH-2 microscope.
Acta Botanica Sinica 植物学报 Vol.46 No.8 2004884
2 Results and Discussion
Two genera Lemanea and Paralemanea of Lemaneaceae
in China are described.
2.1 Lemanea
Lemanea Bory, Ann. Mus. Hist. Paris 12: 178, 1808.
Apona Adanson, Fam. Pl. 2: 519, 1763. — Sacheria
Sirodot, Ann. Sci. Nat. Bot. Ser. 5, 16: 1, 1872 (nom.
illegitimate).— Subgen. Lemanea sensu. Silva, Taxon 8: 62,
1959. — Subgen. Sacheria Ketel, Anat. Unters. Lemanea,
p. 15, 1887 (nom. illegitimate).
Description: Outer cortex is 2 – 6-layered cells in
thickness. Each axial cell has four ray cells, T- or L-shaped,
closely abutting the outer cortex. Central axis has no corti-
cal filaments. Spermatangia are usually in patches.
Type species: L. fluviatilis (Linnaeus) Agardh (= L.
corallina Bory, nom. illegitimate).
The main taxonomic characteristics of Lemanea involve
the size and shape of plants, whether they are branched or
unbranched, stalked at the base or stalkless, with nodes
obvious or not; location and shape of spermatangia, and
morphology of carposporophytes and carposporangia.
More than ten species of this genus have been reported
(Kumano, 2002). Up to now, four species were found in
China among them, two were reported by Jao (1941).
Synoptic key of the Lemanea in China
1 Spermatangia in perfect rings⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯1. L. sinica
1 Spermatangia in patches or imperfect rings⋯⋯⋯⋯⋯
⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯2
2 Plants large, 9 – 13 cm long⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯2. L. crassa
2 Plants small, 1 – 3 cm long⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯3
3 Plants branched⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯3. L. ramosa
3 Plants unbranched⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯4. L. simplex
2.1.1 L. sinica L. sinica Jao, Sinensia 12: 270, pl. VII,
Figs. 49-57,1941.
The species is endemic to China and collected from
Zhongdian, Yunnan in Dec., 1936 by C. W. Wang (YN1).
The specimens of this alga were named as L. fucina Bory
by Li (1939). After being carefully examined, C. C. Jao found
that the carposporangia of the plant are large and single
and spermatangia are in rings, therefore, this alga differs
from L. fucina Bory.
In addition, L. fucina was reported from Yunnan in China
by Li (1940), but he made only very simple descriptions
and neither characteristics about carposporophytes and
spermatangia were described, nor figures were given. Be-
cause of not seeing the specimens, it is not determined.
2.1.2 L. crassa S. L. Xie et Z. X. Shi, sp. nov. L. crassa,
sp. nov. Figs. 1-3
Diagnosis: Species nova similis L. sinensi a qua fronde
haud ramosa, soris antheridiorum tubercularibus differt.
Frons rigida, attenuata versus apicem, angustata ver-
sus basem, 9-13 cm longa; ramis nullisi; basis pedicello
longo, 0.5-0.7 mm lato, 1.5-2.5 cm longo; nodi manifesto
inflati, 1.5-3 mm lati et 0.8-1.2 mm lati internodis.
Carposporangia magno cylindrico-obovoidea, ellipsoidea
vel subdumbbella, singulata, 50-80 mm lata, 90-120 mm
longa. Sori antheridiorum irregulariter tuberculares, 5-6
frustris omnino nodis, manifesto protuberantes.
Description: Plant is rigid, 9-13 cm long, attenuated
upwards and narrowed downwards at the base into a long
stalk, 1.5-2.5 cm long, 0.5-0.7 mm in diameter; unbranched;
nodes are inflated, 1.5-3.0 mm in diameter and internodes
are 0.8-1.2 mm in diameter. Carposporangia are large, cy-
lindrical-obovate, elliptical or dumbbell-shaped, single, 90-
120 mm long, 50-80 mm in diameter. Spermatangia are in
patches, 5- 6 per node, obviously prominent.
Habitat: Jinyutan, Tongliang, Chongqing, collected by
Zhu Haoran on Feb. 5, 1946, not numbered (type, HBI).
This species is similar to L. sinica, but it differs from the
latter in its unbranching and having patchy spermatangia.
2.1.3 L. ramosa S. L. Xie et Z. X. Shi, sp. nov. L. ramosa,
sp. nov. Figs. 4-7
Diagnosis: Species nova similis L. simplici a qua ramulis
densis differt.
Frons parva, dense caespitosa, attnuata versus apicem,
1-3 cm alta, ramis densis praesertim fronte superioribus;
basin haud manifesto pedicello; nodi 0.12-0.45 mm lati et
internodi 0.10-0.25 mm lati. Carposporangia magna,
ellipsoidea, singulata, 25-60 mm lata, 80-100 mm longa. Sori
antheridiiorum irregulariter tuberculares, 6-8 frustris omnion
nodis, interdum imperfecti continui annuli.
Description: Plant is small, densely caespitose, 1-3 cm
long, attenuated upwards, and with no obvious stalk at the
base; branches are dense, especially at the upper parts.
Nodes may be obvious, 0.12-0.45 mm in diameter and in-
ternodes are 0.10- 0.25 mm in diameter. Carposporangia
are large and elliptical being 80-100 mm in length, 25-60
mm in diameter and single. Spermatangia are in patches,
sometimes in imperfect rings with 6-8 per node, sometimes
in imperfect rings.
Habitat: Fuzhou, Fujian, collected on Dec. 10, 1964,
XIE Shu-Lian et al.: Taxonomy of the Lemaneaceae (Batrachospermales, Rhodophyta) in China 885
collector incognito, FK138 (type, HBI).
This species is similar to L. simplex Jao, but it differs
from the latter in its being densely branched.
2.1.4 L. simplex L. simplex Jao, Sinensia 12: 272, pl. VI,
Figs. 46, pl. VII, Figs. 47, 48, 1941.
The species is endemic to China and collected from
Beibei, Chongqing, on Jan. 18, 1940 and Mar. 10, 1941, by
C. C. Jao (SC1128 and SC1246).
2.2 Paralemanea
Paralemanea (Silva) Vis et Sheath, Phycologia 31: 177,
1992.
Lemanea Subgen. Paralemanea Silva, Taxon 8: 62, 1959.
Lemanea Subgen. Lemanea Sirodot, Ann. Sci. Nat. Ser. 5
Bot. 16: 1, 1872 (nom. illegitimate). Lemanea Subgen.
Eulemanea Hamel, Rev. Algol. 2: 59, 1925 (nom. illegitimate).
Description: Outer cortex is 4 – 6 cell-layers in thickness.
Ray cells are simple, not abutting the outer cortex. Central
axis is with slender cortical filaments. Spermatangia are usu-
ally in rings around the nodes.
Type species: Paralemanea catenata (Kützing) Vis et
Sheath.
The main taxonomic characteristics of Paralemanea are
Figs.1-7. 1-3. Lemanea crassa S. L. Xie et Z. X. Shi 1. Habit of plants. 2. Nodes and spermatangial sori. 3. Carposporangia. 4-
7. L. ramosa S. L. Xie et Z. X. Shi 4. Habit of plants. 5. Nodes and spermatangial zones. 6. Axis filament. 7. Carposporangia.
Acta Botanica Sinica 植物学报 Vol.46 No.8 2004886
the size of plants, whether it is branched or unbranched,
stalked at the base or stalkless, with obvious nodes or not,
as well as location and shape of spermatangia.
Four species of the genus were reported and only two
in China. This genus discovered in China is for the first
time reported.
Synoptic key of the Paralemanea in China
1 Spermatangia in rings⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯1. P. catenata
1 Spermatangia in patches⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯2. P. parvula
2.2.1 P. catenata P. catenata (Kützing) Vis et Sheath,
Phycologia 31: 177, Fig.11, 1992; Sheath et al., Phycol. Res.
44: 237, Figs. 45, 46, 52-55, 59, 1996. Figs. 8-11
L. catenata Ktzing, Phycol. German., p. 261, 1845.— L.
nodosa Kützing, Spec. Alg., p. 52.
Description: Plant is large and unbranched, 5-12 cm
long, attenuated upwards and narrowed downwards. There
Figs.8-14. 8-11. Paralemanea catenata (Kützing) Vis et Sheath 8. Habit of plants. 9. Nodes and spermatangial zones. 10.
Carposporangia. 11. Axis filament and cortical filaments. 12-14. P. parvula (Sirodot) S. L. Xie et Z. X. Shi 12. Habit of plants. 13.
Nodes and spermatangial sori. 14. Carposporangia.
XIE Shu-Lian et al.: Taxonomy of the Lemaneaceae (Batrachospermales, Rhodophyta) in China 887
is no obvious stalk at the base; nodes are inflated, 0.5-1.0
mm in diameter and internodes are 0.3-0.6 mm in diameter.
All cells of gonimoblasts transform into carposporangia in
chains, being branched. Carposporangia are small and
elliptical, 22-25 mm long and 15.0-17.5 mm in diameter.
Spermatangia are in rings being significantly prominent.
Habitat: Menghai, Yunnan, collected by Qian Chengyu
on Mar. 11, 1957 (YN5703).
Distribution: Germany, Sweden, Belgium, France and
America.
This species found in China and Asia is reported for the
first time.
2.2.2 P. parvula P. parvula (Sirodot) S. L. Xie et Z. X.
Shi, comb. nov. Figs. 12-14
L. parvula Sirodot, Ann. Sci. Nat. Bot. Ser. 5, 16: 85, pl. 5,
Figs. 35-39, 1872.
Description: Plant is 3-5 cm long, attenuated upwards,
short stalked at the base and unbranched; nodes are
inflated, 0.5- 0.8 mm in diameter and internodes are 0.3-
0.5 mm in diameter. All cells of gonimoblasts transform into
carposporangia in chains, branched; carposporangia are
small, elliptical to cylindrical-elliptical, (18-)27-30 mm long,
15-17 mm in diameter. Spermatangia are in patches and not
obviously prominent.
Habitat: Mengyang, Jinghong, Yunnan, collected by
Qian Chengyu in Mar., 1957 (YN5701).
Distribution: France and Britain.
This species found in China is reported for the first time.
So we can make a conclusion that two genus and six
species belonging to Lemaneaceae have been found in China
up to now.
References:
Agardh C. 1828. Species Algarum. Cryphiae. 189.
Atkinson G F. 1890. Monograph of the Lemaneaceae of the United
States. Ann Bot, 4: 177-229.
Bory de Saint-Vincent J B. 1808. Mémoire sur le genre Lemanea
de la famille des Conferves. Ann Mus Hist Nat, 12: 177-190.
Greuter W. 1994. International Code Botanical Nomenclature
(Tokyo Code). Königstein: Koeltz Scientific Books.
Hamel G. 1925. Floridées de France. Ⅲ, Ⅳ. Rev Algol, 2: 39-
67, 280-309.
Israelson G. 1942. The freshwater Florideae of Sweden: studies
on their taxonomy, ecology and distribution. Symb Bot Upsal,
6: 1-135.
Jao C C. 1941. Studies on the freshwater algae of China Ⅷ. A
preliminary account of the Chinese freshwater Rhodophyceae.
Sinensia, 12: 245-260.
Ketel K F. 1887. Anatomische Untersuchungen über die gattung
Lemanen. Drunk von F. W. Kunike. Greifswald, 39.
Khan M. 1973. On edible Lemanea Bory de St Vincent— resh
water red alga from India. Hydrobiologia, 43: 171-175.
Kumano S. 2002. Freshwater Red Algae of the World. Bristol:
Biopress Limited. 253-278.
Li L C. 1939. Freshwater algae of Yunnan expedition 1935-1937.
Bull Fan Mem Inst Biol Bot Ser, 9: 31-57.
Li L C. 1940. Additions to the freshwater algae Yunnan. Bull Fan
Mem Inst Biol Bot Ser, 10: 47-67.
Necchi O Jr, Zucchi M R. 1995. Record of Paralemanea
(Lemaneaceae, Rhodophyta) in South America. Arch Hydrobiol
Suppl, 109: 33-38.
Palmer C M. 1941. A study of Lemanea in Indiana with notes on
its distribution in North America. Butler Univ Bot Stud, 5: 1-
26.
Sheath R G, Müller K M, Vis M L, Entwisle T J. 1996. A re-
examination of the morphology, ultrastructure and classifica-
tion of genera in the Lemaneaceae (Batrachospermales,
Rhodophyta). Phycol Res, 44: 233-246.
Silva P C. 1959. Remark on algol nomenclature, II. Taxon, 8: 60-
64.
Sirodot M S. 1872. étude anatomique, organogénique et
physiologique sur les algues d’eau douce de la famille des
Lémanéacées. Ann Sci Nat Bot, 16: 5-95.
Vis M L, Sheath R G. 1992. Systematics of the freshwater red
algal family Lemaneaceae in North America. Phycologia, 31:
164-179.
(Managing editor: HAN Ya-Qin)