new red algal species, erythroglossum hyacinthinum ... · pdf filethis alga resembles...
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Algae 2014, 29(1): 1-13http://dx.doi.org/10.4490/algae.2014.29.1.001
Open Access
Research Article
Copyright © 2014 The Korean Society of Phycology 1 http://e-algae.kr pISSN: 1226-2617 eISSN: 2093-0860
New red algal species, Erythroglossum hyacinthinum (Delesseriaceae, Rhodophyta) from Korea
Jeong Chan Kang1 and Myung Sook Kim1,*1Department of Biology and Research Institute for Basic Sciences, Jeju National University, Jeju 690-756, Korea
The genus Erythroglossum is characterized by Phycodrys-type apical organization, Polyneura-type procarp, and the
presence of a midrib. We collected an unidentified Delesseriaceaen species from deep water off the southern coast of the
Korean Peninsula. This alga resembles Polyneura japonica in terms of having broadly flattened thalli with a cylindrical
stipe, the presence of a midrib and alternative lateral veins. To confirm the taxonomic status of this entity, we compared
the morphological features and rbcL sequences among other species of Erythroglossum and P. japonica. As a result, we
assigned the new species, Erythroglossum hyacinthinum, to the genus Erythroglossum because of the presence of a mid-
rib. This species is characterized by an elliptical to obovate blade with a short cylindrical stipe, a conspicuous midrib
and alternate veins, margins with numerous microscopic dentations, di-trichotomously branching, and bulish-violet iri-
descence. The phylogeny of rbcL sequences indicates that E. hyacinthinum is definitely a separate entity, but the genera
in the tribe Phycodryeae have inconsistent phylogenetic relationships. This is the first study comparing the molecular
phylogeny within the genus Erythroglossum.
Key Words: Delesseriaceae; Erythroglossum hyacinthinum sp. nov.; morphology; Phycodryeae; Polyneura; rbcL; Rhodophyta; taxonomy
INTRODUCTION
The genus Erythroglossum was established by Agardh
(1898) based on the vegetative characteristics such as
marginal ramifications and angular-rounded cell shape,
and he transferred five species of Delesseria (e.g., D.
schousboei J. Agardh, D. balearica J. J. Rodríguez, D. woodii
J. Agardh, D. bipinnatifida Montagne, and D. californica J.
Agardh) to the genus Erythroglossum. Later, Kylin (1924)
designated E. schousboei (J. Agardh) J. Agardh as the lec-
totype species of Erythroglossum, and he established an-
other genus Branchioglossum for E. woodii (J. Agardh)
J. Agardh (Wynne 1983). Mikami (1979) reported that E.
bipinnatifidum (Montagne) J. Agardh also has the same
characteristics as Branchioglossum. Wynne (1983) agreed
with Mikami and made the new combination, Branchio-
glossum bipinnatifidum (Montagne) M. J. Wynne. A total
of 12 species of the genus Erythroglossum are currently
accepted world-wide, including the original three spe-
cies, which were first transferred from Delesseria to Eryth-
roglossum by Agardh (1898) (Guiry and Guiry 2013).
Kylin (1924) divided the family Delesseriaceae into 11
groups, which were assigned to two subfamilies (e.g., De-
lesserioideae and Nitophylloideae) on the basis of veg-
etative and reproductive structures. He additionally as-
signed the genus Erythroglossum to his Phycodrys group
of the subfamily Nitophylloideae. Wynne (2001) proposed
23 tribes under two subfamilies, and he placed the genus
Received October 20, 2013, Accepted January 19, 2014
*Corresponding Author
E-mail: [email protected]: +82-64-754-3523, Fax: +82-64-756-3541
This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Com-
mercial License (http://creativecommons.org/licenses/by-nc/3.0/) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.
Algae 2014, 29(1): 1-13
http://dx.doi.org/10.4490/algae.2014.29.1.001 2
MATERIALS AND METHODS
Specimens were collected from 15 localities of the Ko-
rean coast and two localities of the coast of Japan, from
the intertidal and subtidal (Table 1). Field-collected
samples were put into an icebox with seawater and an
icepack, and transported to the laboratory. The samples
for DNA extraction were each assigned a voucher num-
ber, and a small part of the thallus was detached for
making silica-gel dried tissue samples. The major part of
the thallus was made into a pressed specimen and was
deposited in the Herbarium of Jeju National University
(JNUB), Jeju, and in the National Institute of Biological
Resources (NIBR), Incheon, Korea. Samples for morpho-
logical observations were fixed in 5% formalin / seawa-
ter. Sections for microscopic examination were made by
using a freezing microtome (NK-101-II; Nippon Optical
Works Co., Ltd., Tokyo, Japan). For staining, 1% aqueous
aniline blue acidified with a drop of 1% HCl was used.
The stained sections were mounted in 30% Karo corn
syrup. Photomicrographs were taken using a QImaging
1394 camera (QImaging, Surrey, BC, Canada) attached to
a BX50 microscope (Olympus, Tokyo, Japan). For compar-
ing molecular data, we used 25 Erythroglossum samples
containing the new species from Korea, four E. pinnatum
samples from Japan, five Polyneura samples from Korea,
and 27 rbcL sequence data from the tribe Phycodryeae
and Myriogrammeae (used as an outgroup) from previ-
ous studies.
For the extraction of total DNA from the silica-gel
dried specimens, we used the same methods and prim-
ers as Kang and Kim (2013). The polymerase chain reac-
tion (PCR) products were purified using the AccuPrep
PCR Purification Kit (Bioneer, Daejeon, Korea) and were
sequenced commercially (Macrogen, Seoul, Korea). Elec-
tropherogram outputs from each sample were edited us-
ing Chromas version 1.45 (McCarthy 1996). The total rbcL
sequence was organized using the multiple-sequence
editing program BioEdit (Hall 1999) and aligned visually
(Kang and Kim 2013). None of the alignments posed a
problem, as no gaps were observed.
Eleven rbcL sequences of the new species were aligned
with 43 sequences of other species belonging to the tribe
Phycodryeae, and seven species of Myriogrammeae were
used as an outgroup. Maximum likelihood (ML) analyses
were produced using RAxML (Stamatakis 2006) with the
GTR + Г evolutionary model. We used 200 independent
tree inferences by the -# option with default –I (automati-
cally optimized SPR rearrangement) and –c (25 distinct
rate categories) options of the program to identify the
Erythroglossum within the tribe Phycodryeae (Lin et al.
2001a). Recently, Lin et al. (2001a) established the third
subfamily as Phycodryoideae, based on molecular analy-
sis including large subunit ribosomal DNA and rbcL se-
quence data. They assigned the four tribes Phycodryeae,
Myriogrammeae, Schizoserideae, and Cryptopleureae to
the subfamily Phycodryoideae: however, they did not test
any species of Erythroglossum.
In the tribe Phycodryeae (Wynne 2001), there are two
groups with different procarp development: 1) Phyco-
drys-type having one carpogonial branch with two sterile
groups, and 2) Polyneura-type having two carpogonial
branches with one sterile group. Five genera including
Erythroglossum, Polyneura Kylin, Sorella Hollenberg,
Sorellocolax Yoshida & Mikami, and Womersleya Papen-
fuss are known to have the Polyneura-type procarp (Yo-
shida and Mikami 1991, 1996, 1997, Maggs and Hom-
mersand 1993, Kim and Nam 1994, Lin and Kraft 1996,
Womersley 2003, Díaz-Tapia et al. 2009). Erythroglossum
is distinguished from Sorella by tetrasporangial posi-
tion (Hollenberg 1943, Yoshida and Mikami 1991), from
Womersleya by the presence of a midrib or vein (Lin and
Kraft 1996), and Polyneura by the presence of a midrib
(Maggs and Hommersand 1993). The genus Sorellocolax
is not comparable to Erythroglossum in terms of size or
their host (Yoshida and Mikami 1996). In Erythroglos-
sum, morphological features such as thallus habit, blade
shape, structure and branching, and veins have been
used for separating species (Maggs and Hommersand
1993, Yoshida and Mikami 1997, Díaz-Tapia et al. 2009).
Recently, rbcL sequence data have been used for compar-
ing the taxonomic position among the species of the fam-
ily Delesseriaceae (Lin et al. 2001b, Lin and Nelson 2010,
Kim and Kang 2011).
Three species of Erythroglossum, namely E. minimum
Okamura, E. pinnatum Okamura, and E. latum Yoshida
& Mikami, are currently known to occur in the north-
western Pacific region, and two of these species (E. mini-
mum and E. pinnatum) are in the Korean coast (Guiry
and Guiry 2013). The aims of this study are to compare
the morphological features of the new species and other
Erythroglossum species and to investigate the phyloge-
netic relationships in the tribe Phycodryeae based on the
analysis of rbcL gene. To compare the molecular data, we
collected two species (E. minimum and E. pinnatum) of
Erythroglossum and Polyneura japonica from Korea and
Japan. Unfortunately, we could not collect E. latum. This
is the first study comparing molecular data within the ge-
nus Erythroglossum.
Kang & Kim Erythroglossum hyacinthinum sp. nov.
3 http://e-algae.kr
Daesambudo, Yeosu on Jul 25, 2012 (JN120725-1, female
gametophyte; JN120725-2, tetrasporophyte); Jakdo, Yeo-
su on Jul 25, 2012 (JN120726-54, tetrasporophyte); Geo-
mundo, Yeosu on Jul 25, 2012 (JN120725-42, female ga-
metophyte).
Morphology. Thalli are erect, broadly lobed, shortly
stipitate, growing up to 3-6 cm high, 2.5 cm wide in the
broadest portion of the lobes, and attached by a discoid
holdfast (Fig. 1A-C & E). The blades are at first simple
and elliptical to obovate in outline, and later divided di-
trichotomously. Each lobe has an elliptical to obovate
outline with a round apex (Fig. 1A-C). The margins of the
blades are beset with microscopic dentations (Fig. 1D).
The blades are mostly monostromatic, except for the
midrib and lateral veins. The midrib is connected with a
stipe, which is conspicuous except near the apical portion
of the blade. Lateral veins divide alternately or dichoto-
mously from the midrib (Fig. 1A, B, E & F). Microscopic
veins or anastomosing nerves are absent. The basis of
stipe produces creeping cylindrical branches, from which
additional thalli are generated. The stipe is cylindrical in
the basal part and is compressed near the starting point
of the blade wing. Young blades irregularly initiate from
the stipes (Fig. 1E-G). In the cross-section views, the veins
are composed of large roundish cells, which are arranged
side by side, and are located in the center of the branch
with several cortical layers. The numbers of cortical cell
layers decrease toward the apical part of veins (Fig. 1H-J).
Fig. 1K shows the apical organization: the growth of thalli
is initiated by an apical cell, which divides transversely
generating the primary cell row. The cells produced by the
transverse division of an apical cell are divided longitudi-
nally to form second-order cell rows laterally. After pro-
ducing second-order cell rows, certain cells of the prima-
ry cell row divide transversely, a process which is termed
intercalary division, and also divide longitudinally. The
second-cell rows reach the thallus margin, and some
continue growth to develop microscopic dentations from
the blade margin. The third-order cell rows are mostly cut
off abaxially from the second-order cell rows, occasion-
ally adaxially. Transverse and longitudinal intercalary cell
divisions are frequent in the primary cell row and higher
order cell rows.
Gametophytes are dioecious. Procarps are scattered
between the midrib and the blade margin in median to
upper parts of blades, consisting of a supporting cell, a
7-8 celled sterile group, and two 4-celled carpogonial
branches (Fig. 2A-C). Mature cystocarps are hemispheri-
cal and 500-700 µm in diameter, more swollen on the os-
tiolate side of blade, and composed of a large branched
best tree. To generate bootstrap values we used the same
program with the same settings for 1,000 replications
(Kang and Kim 2013).
RESULTS
Erythroglossum hyacinthinum J. C. Kang & M. S. Kim sp. nov.
Description. Thalli erect, 3-6 cm high and 1.5-2.5 cm
wide, lobed and membranous with a short cylindrical
stipe, attached to the substratum by discoid holdfast,
simple or divided di- or trichotomously; blades ellipti-
cal to obovate with rounded apex; blade margins beset
with microscopic dentations; midribs conspicuous and
faint near apical part of lobes, lateral veins divided from
midrib alternately to dichotomously; microscopic veins
absent; bright red with bulish-violet iridescence in liv-
ing condition; cystocarps hemispherical with an ostiole,
scattered over the monostromatic areas of the middle to
upper blades; procarps consist of a supporting cell, a 7-8
celled sterile group, and two groups of 4-celled carpogo-
nial branches; spermatangial sori produced on the lamel-
lae between midrib and blade margin in the upper parts
of blade; tetrasporangial sori round to irregular shape,
scattered on monostromatic portions of middle to upper
blades, composed of two layers of tetrasporangia; mature
tetrasporangia spherical and divided tetrahedrally.
Holotype. JN130604-1 (tetrasporophyte), collected
from 12 m depth of Chujado, Jeju province, Korea
(33°58′04.28″ N, 126°17′08.64″ E) on Jun 4, 2013, and de-
posited in the Herbarium of Department of Biology, Jeju
National University, Korea (JNUB).
Isotypes. JNUB (JN130604-3, JN130604-5 to -9) and KB
(NIBRAL0000137942-3).
Etymology. The specific epithet (hyacinthinum) was
chosen to represent the color of this species when alive,
having a bulish-violet iridescence. Latin: hyacinthinus,
-a, -um. adj. A.
Korean name. 푸른빛붉은혀
Habitat. Erythroglossum hyacinthinum was collected
at 10-30 m depth where they were growing on bedrock.
Other specimens examined. Sasudo, Jeju on Jun 5,
2013 (JN130605-1, tetrasporophyte; JN130605-2, female
gametophyte; JN130605-3, male gametophyte); Gwideok,
Jeju on May 31, 2011 (H011 and H015, female gameto-
phyte); Jocheon, Jeju on May 20, 2011 (H010, tetrasporo-
phyte); Dueokdo, Wando on Jul 26, 2012 (JN120726-23,
tetrasporophyte; JN120726-26, female gametophyte);
Algae 2014, 29(1): 1-13
http://dx.doi.org/10.4490/algae.2014.29.1.001 4
Fig. 1. Erythroglossum hyacinthinum J. C. Kang and M. S. Kim sp. nov. (A) Holotype specimen from Chujado (JN130604-1), Jeju Island on Jun 4, 2013. (B) Habit of young thalli showing simple and elliptical to obovoid outline. (C) In the living condition, the thalli having bulish-violet iridescence. (D) Blade margin with microscopic dentations. (E & F) Basal part of thalli composed of discoid holdfasts (arrowheads) with creeping cylindrical branches (arrows) and short stipes. (G-J) Cross section views of stipe (G), the lower (H), the middle (I), and the upper part (J) of main branch: the blade composed of monostromatic lamellae and a polystromatic midrib. (K) Apical organization of thallus is showing primary (1) and higher order cell-rows (2-4), and cells resulting from intercalary divisions (i). Scale bars represent: A, 2 cm; B & D, 500 µm; E & F, 1,000 µm; G-J, 200 µm; K, 20 µm.
A C
D
B
E GF
HI
J
K
Kang & Kim Erythroglossum hyacinthinum sp. nov.
5 http://e-algae.kr
Fig. 2. Erythroglossum hyacinthinum J. C. Kang and M. S. Kim sp. nov. (A) Cystocarps (cp) scattered on the monostromatic lamellae between midrib (md) and blade margin. (B & C) Procarp composed of a supporting cell (sc), two groups of four-celled carpogonial branches (cb) with a trichogyne (tr), and a group of sterile cells (st), which are connected by pit-connections (arrows). (D) A mature cystocarp composed of a large branched fusion cell (fu), short-chained carposporangia (ca) on the terminal of gonimoblast filaments (gb), several cells layer pericarp, and an prominent ostiole on apical portion. (E) Spermatangial sori (ss) produced between midrib (md) and the blade margin. (F) Surface view of spermatangial sorus showing numerous spermatangia (sp). (G) Cross-section view of spermatangial sorus showing spermatangia (sp) produced by spermatangial mother cells (smc), which occur on opposite sides of central cells (cc). (H) Tetrasporangial sori (ts) produced between midrib (md) and blade margin. (I) Surface view of tetrasporangial sorus. (J) Cross-section view of tetrasporangial sorus showing two layers of tetrasporangia. Scale bars represent: A & H, 1,000 µm; B, C, F & G, 20 µm; D, 100 µm; E, 500 µm; I & J, 50 µm.
A C
D
B
E
GF
H
I J
Algae 2014, 29(1): 1-13
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Fig. 3. Maximum likelihood phylogenetic tree (log likelihood = -7,719.609264) for the genus Erythroglossum and putative relatives derived from plastid-encoded rbcL sequence data. The bootstrap values (1,000 replicates) are shown above branches. Scale bar represents: substitution per site.
Kang & Kim Erythroglossum hyacinthinum sp. nov.
7 http://e-algae.kr
Erythroglossum, Polyneura, Sorella, Sorellocolax, and
Womersleya are known to have procarp consisting of a
pair of carpogonial branches and one sterile-cell group
(Lin and Kraft 1996, Yoshida and Mikami 1996). The diag-
nostic morphological characteristics among the five gen-
era are as follows: 1) the presence of midrib or midveins,
and the marginal position of tetrasporangial sori for
Erythroglossum; 2) presence of macroscopic midveins,
and median position of tetrasporangial sori for Sorella; 3)
midrib absent, presence of macroscopic and microscopic
anastomosing nerves, and inter nerves position of tetra-
sporangial sori for Polyneura; 4) lack of microscopic and
macroscopic veins, and growth by transversely dividing
marginal apical cells, with the later formation of a con-
tinuous marginal meristem for Womersleya; 5) very small
stellate and hemiparasitic thalli on the genus Sorella for
Sorellocolax (Hollenberg 1943, Maggs and Hommersand
1993, Yoshida and Mikami 1996, Womersley 2003). The
diagnostic characters of the genus Erythroglossum are
entirely satisfied by morphology of our new species.
In the genus Erythroglossum, the following diagnostic
characters for identification at the species level have been
used: habit of thallus, branching patterns, and veins and
blade structures (Maggs and Hommersand 1993, Yoshida
and Mikami 1997, Díaz-Tapia et al. 2009). In observations
of the morphological features, E. hyacinthinum is clearly
separated from other species of Erythroglossum from the
northwestern Pacific Ocean (Table 1): in terms of thallus
size from E. minimum (3-6 cm high and 1.5-2.5 cm width
vs. 1-2.5 cm high and 1-2 mm width), branching patterns
from E. pinnatum (di-trichotomous vs. 2-3 times pinnate
manner), and blade structures from E. latum (mostly
monostromatic vs. mostly polystromatic). This new spe-
cies more closely resembles Polyneura japonica rather
than other Erythroglossum species in terms of discoid
holdfast, cylindrical stipe, di-trichotomous branching,
presence of midrib and lateral veins, and blade margins
with fine teeth, but differs in blade structure (mostly
monostromatic vs. mostly polystromatic), iridescence
(bulish-violet vs. none), and shapes (elliptical to obovate
vs. broadly linear) (Table 2). Upon first observation of E.
hyacinthinum at 30 m depth, we were very confused as
to its taxonomic status due to the extensive morphologi-
cal similarities with P. japonica. However, we assigned the
new species to the genus Erythroglossum because the
morphological features (i.e., presence of midrib and lat-
eral veins, marginal position of tetrasporangial sori) are
more in agreement with the genus Erythroglossum than
Polyneura (Maggs and Hommersand 1993).
P. japonica was originally described by Yamada as Het-
fusion cell generating numerous radiate gonimoblast
filaments, 2-4 chains of ovoid to pyriform carposporan-
gia terminating each gonimoblast filament, and 5-7 cells
thick pericarp (Fig. 2D). Spermatangial sori are produced
between the midrib and blade margin in the upper parts
of the blades, roundish to elliptical at first and later ex-
pand and become irregular in shape, consisting of a layer
of central cells, two layers (each layer on opposite sides
of central cells) of quadrangular spermatangial mother
cells, which bearing elongate spermatangia (Fig. 2H-G).
Tetrasporangial sori are scattered between veins and
blade margin in median to upper parts of blades, circular
at first and adjacent sori becoming coalescent as they ex-
pand, consisting of two layers of tetrasporangia. Mature
tetrasporangia are spherical, 400-500 µm in diameter and
divide tetrahedrally (Fig. 2I & J).
Molecular analysis. We determined a total of 61 rbcL
sequence data: 25 Erythroglossum samples containing
the new species from Korea, four E. pinnatum samples
from Japan, five Polyneura samples from Korea, and 27
rbcL from the tribe Phycodryeae as putative relatives, and
the tribe Myriogrammeae as an outgroup. We aligned
1187 nucleotide base pairs of rbcL gene, and of all sites,
383 (32.2%) were variable and 270 (22.7%) were phyloge-
netically informative. Eleven specimens of E. hyacinthi-
num from seven sites in Korea formed a clade with 0-0.2%
divergence within the clade. E. hyacinthinum showed 2.7-3.3% divergence from other Erythroglossum species, and
1.8-2.0% divergence from Polyneura japonica.
In the phylogenetic tree (Fig. 3), the clade contain-
ing Erythroglossum spp., P. japonica, and P. latissima
was clearly separated from the other species of the tribe
Phycodryeae, and strongly supported monophyly by the
100% ML bootstrap value. The clade of E. hyacinthinum
formed a sister clade with P. japonica in a 70% boot-
strap value. However, the Polyneura-type procarpic spe-
cies (i.e., Erythroglossum spp., P. japonica, P. latissima, P.
bonnemaisonii, and Womersleya monanthos) were not
monophyletic. In contrast, the Phycodrys-type procarpic
species (i.e., Phycodrys, Nienburgia, Heterodoxia, and Hy-
menenopsis) were monophyletic with low bootstrap sup-
port (69%).
DISCUSSION
Our data clearly indicates that we have collected a new
species of Erythroglossum in Korea, E. hyacinthinum.
This species does not match any other species present-
ly described. In the tribe Phycodryeae, the five genera
Algae 2014, 29(1): 1-13
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Tabl
e 1.
Col
lect
ion
info
rmat
ion
of s
ampl
es u
sed
in th
is s
tudy
Taxa
C
oll
ecti
on
info
rmat
ion
Hab
itat
(dep
th /
su
bst
ratu
m)
V
ou
cher
Gen
Ban
k ac
cess
ion
No.
Ref
eren
ces
Ery
thro
glos
sum
hya
cin
thi-
nu
m s
p. n
ov.
Ch
uja
do,
Jeju
, Ko
rea;
Jun
4, 2
013
12-1
5 m
dep
th /
bed
ro
ckJN
1306
04-1
()
KF
3052
99T
his
stu
dy
Ch
uja
do,
Jeju
, Ko
rea;
Jun
4, 2
013
12-1
5 m
dep
th /
bed
ro
ckN
IBR
AL0
0001
3794
2 (♂
)K
F30
5300
Th
is s
tud
y
Dae
sam
bu
do,
Yeo
su, K
ore
a; Ju
l 25,
201
225
-30
m d
epth
/ b
ed r
ock
JN12
0725
-2 (
)K
F30
5294
Th
is s
tud
y
Du
eokd
o, W
and
o, K
ore
a; Ju
l 27,
201
210
m d
epth
/ b
ed r
ock
JN12
0727
-23
()
KF
3052
95T
his
stu
dy
Gw
ideo
k, Je
ju, K
ore
a; M
ay 3
1, 2
011
25 m
dep
th /
bed
ro
ckH
011
(♀)
KF
3052
97T
his
stu
dy
Gw
ideo
k, Je
ju, K
ore
a; M
ay 3
1, 2
011
25 m
dep
th /
bed
ro
ckH
015
(♀)
KF
3052
96T
his
stu
dy
Jakd
o, Y
eosu
, Ko
rea;
Jul 2
6, 2
012
20-2
5 m
dep
th /
bed
ro
ckJN
1207
26-5
4 (
)K
F30
5293
Th
is s
tud
y
Joch
eon
, Jej
u, K
ore
a; M
ay 2
0, 2
011
30 m
dep
th /
bed
ro
ckH
010
()
KF
3052
98T
his
stu
dy
Sasu
do,
Jeju
, Ko
rea;
Jun
5, 2
013
12-1
5 m
dep
th /
bed
ro
ckJN
1306
05-1
()
KF
3053
01T
his
stu
dy
Sasu
do,
Jeju
, Ko
rea;
Jun
5, 2
013
12-1
5 m
dep
th /
bed
ro
ckJN
1306
05-2
(♀
)K
F30
5302
Th
is s
tud
y
Sasu
do,
Jeju
, Ko
rea;
Jun
5, 2
013
12-1
5 m
dep
th /
bed
ro
ckJN
1306
05-3
(♂
)K
F30
5303
Th
is s
tud
y
Ery
thro
glos
sum
min
imu
m
Oka
mu
raB
iyan
gdo,
Jeju
, Ko
rea;
Nov
22,
201
210
m d
epth
/ C
lad
oph
ora
wri
ghti
ana
JN12
1122
-4 (
)K
F30
5292
Th
is s
tud
y
Biy
angd
o, Je
ju, K
ore
a; N
ov 2
2, 2
012
10 m
dep
th /
C. w
righ
tian
aJN
1211
22-6
(♀
)K
F30
5291
Th
is s
tud
y
Biy
angd
o, Je
ju, K
ore
a; F
eb 2
6, 2
013
Low
inte
rtid
al /
bed
ro
ckJN
1302
26-1
1 (
)K
F30
5287
Th
is s
tud
y
Biy
angd
o, Je
ju, K
ore
a; N
ov 2
2, 2
012
Low
inte
rtid
al /
bed
ro
ckJN
1302
26-8
()
KF
3052
82T
his
stu
dy
Do
kdo,
Ule
un
g, K
ore
a; A
pr
22, 2
013
5-10
m d
epth
/ b
ed r
ock
JN13
0422
-6 (
)K
F30
5284
Th
is s
tud
y
Do
kdo,
Ule
un
g, K
ore
a; A
pr
22, 2
013
5-10
m d
epth
/ b
ed r
ock
JN13
0422
-9 (
)K
F30
5283
Th
is s
tud
y
Gap
ado,
Jeju
, Ko
rea;
Mar
26,
201
38-
12 m
dep
th /
C. w
righ
tian
aJN
1303
26-1
3 (♀
)K
F30
5285
Th
is s
tud
y
Gap
ado,
Jeju
, Ko
rea;
Mar
26,
201
38-
12 m
dep
th /
C. w
righ
tian
aJN
1303
26-1
4 (♂
)K
F30
5286
Th
is s
tud
y
Gap
ado,
Jeju
, Ko
rea;
Mar
26,
201
38-
12 m
dep
th /
C. w
righ
tian
aJN
1303
26-2
1 (
)K
F30
5288
Th
is s
tud
y
Jeo
ngd
ori
, Wan
do,
Ko
rea;
Jun
9, 2
012
Low
inte
rtid
al /
bed
ro
ckJN
1206
09-2
9K
F30
5289
Th
is s
tud
y
Jeo
ngd
ori
, Wan
do,
Ko
rea;
Jan
16,
201
3Lo
w in
tert
idal
/ b
ed r
ock
JN13
0116
-9 (
)K
F30
5290
Th
is s
tud
y
Ery
thro
glos
sum
pin
nat
um
O
kam
ura
Mu
nse
om
, Jej
u, K
ore
a; Ja
n 3
0, 2
013
4 m
dep
th /
bed
ro
ckJN
1301
30-1
5 (♀
)K
F30
5275
Th
is s
tud
y
Mu
nse
om
, Jej
u, K
ore
a; Ja
n 3
0, 2
013
4 m
dep
th /
bed
ro
ckJN
1301
30-1
9 (♂
)K
F30
5277
Th
is s
tud
y
Mu
nse
om
, Jej
u, K
ore
a; Ja
n 3
0, 2
013
4 m
dep
th /
bed
ro
ckJN
1301
30-2
0 (
)K
F30
5276
Th
is s
tud
y
Mis
aki,
Jap
an; A
pr
10, 2
013
Low
inte
rtid
al /
bed
ro
ckJN
1304
10-7
9K
F30
5281
Th
is s
tud
y
Shim
od
a, Ja
pan
; Ap
r 12
, 201
3Lo
w in
tert
idal
/ b
ed r
ock
JN13
0412
-1 (
)K
F30
5279
Th
is s
tud
y
Shim
od
a, Ja
pan
; Ap
r 12
, 201
3Lo
w in
tert
idal
/ b
ed r
ock
JN13
0412
-2 (
)K
F30
5280
Th
is s
tud
y
Shim
od
a, Ja
pan
; Ap
r 12
, 201
3Lo
w in
tert
idal
/ b
ed r
ock
JN13
0412
-3 (
)K
F30
5278
Th
is s
tud
y
Pol
yneu
ra ja
pon
ica
(Y
amad
a) M
ikam
iC
hu
ja, J
eju
, Ko
rea;
Jun
4, 2
013
12 m
dep
th /
bed
ro
ckJN
1306
04-1
0K
F30
5274
Th
is s
tud
y
Gan
jeo
lgo
t, U
lsan
, Ko
rea;
Jul 2
1, 2
012
Was
hed
up
on
the
sho
reJN
1207
21-1
3K
F30
5271
Th
is s
tud
y
Gw
anga
nri
, Bu
san
, Ko
rea;
Dec
20,
201
25-
10 m
dep
th /
bri
dge
po
stJN
1212
20-4
8 (♀
)K
F30
5273
Th
is s
tud
y
Jeo
ngd
ori
, Wan
do,
Ko
rea;
Jun
9, 2
012
Was
hed
up
on
the
sho
reJN
1206
09-2
7K
F30
5270
Th
is s
tud
y
Son
gjeo
ng,
Bu
san
, Ko
rea;
Dec
20,
201
28-
12 m
dep
th /
bed
ro
ck
JN12
1220
-06
KF
3052
72T
his
stu
dy
Kang & Kim Erythroglossum hyacinthinum sp. nov.
9 http://e-algae.kr
Tabl
e 1.
Con
tinue
d
Taxa
C
oll
ecti
on
info
rmat
ion
Hab
itat
(dep
th /
sub
stra
tum
)Vo
uch
er
Gen
Ban
k ac
cess
ion
No.
R
efer
ence
s
Cla
dod
onta
lyal
lii
(J. D
. Ho
oke
r &
Har
vey)
Sk
ott
sber
g
Ro
oke
ry B
ay, S
tan
ley,
E
. Fal
klan
d Is
lan
ds;
Ja
n 4
, 199
8
--
AF
2541
69Li
n e
t al.
(200
1a)
Har
ald
iop
hyl
lum
bon
nem
aiso
nii
(K
ylin
) A
. D. Z
inov
aN
ear
Fan
ad H
ead
, Co.
Do
neg
al,
UK
; May
21,
200
0-
-A
F31
2311
Lin
et a
l. (2
001a
)
Har
ald
iop
hyl
lum
cri
spat
um
(J
. D. H
oo
ker
& H
arve
y) L
in,
Ho
mm
ersa
nd
& N
elso
n
Mill
Cre
ek E
stu
ary,
Ob
an, S
tew
art I
slan
d,
New
Zea
lan
d; O
ct 3
1, 2
004
--
DQ
9163
05Li
n e
t al.
(200
7)
Har
ald
iop
hyl
lum
mir
abil
e
(Kyl
in)
A. D
. Zin
ova
Can
ove
Isla
nd
, San
Juan
Isla
nd
, W
ash
ingt
on
, USA
; Ju
n 2
9, 1
998
--
AF
2541
85Li
n e
t al.
(200
1a)
Har
ald
iop
hyl
lum
sp.
La H
erra
du
ra, C
oq
uim
bo,
C
hile
; Jan
19,
199
5-
-A
F25
4188
Lin
et a
l. (2
001a
)
Har
ald
iop
hyl
lum
ud
oen
sis
M
. S. K
im &
J. C
. Kan
gH
aum
okd
on
g, U
do,
Jeju
, K
ore
a; Ju
n 1
4, 2
009
12 m
dep
th /
sto
ne
JNU
-MSK
3060
1HU
()
JN56
1293
Kim
an
d K
ang
(2
011)
Het
erod
oxia
den
ticu
lata
(K
un
tze)
J.
Aga
rdh
War
rnam
bo
ol,
Vic
tori
a,
Au
stra
lia; J
ul 1
3, 1
995
--
AF
2541
90Li
n e
t al.
(200
1a)
Hym
enen
opsi
s h
eter
oph
ylla
S.
-M
. Lin
, W. A
. Nel
son
& M
. H.
Ho
mm
ersa
nd
Mar
fells
Bea
ch, M
arb
oro
ugh
, So
uth
Isla
nd
, New
Zea
lan
d;
Nov
5, 2
010
-
W
ELT
A03
0892
JF49
5097
Lin
et a
l. (2
012)
Myr
iogr
amm
e li
vid
a
(J. D
. Ho
oke
r &
Har
vey)
Kyl
inSe
alio
n Is
lan
d,
Falk
lan
d Is
lan
ds;
Jan
7, 1
998
--
AF
2573
91Li
n e
t al.
(200
1a)
Myr
iogr
amm
e m
angi
nii
(G
ain
) Sk
ott
sber
gB
ahia
Ele
fan
te, B
ase
Frei
, Kin
g G
eorg
e I.
, A
nta
rcti
c Pe
nin
sula
; Feb
5, 1
994
--
AF
2573
92Li
n e
t al.
(200
1a)
Nie
nbu
rgia
an
der
son
ian
a
(J. A
gard
h)
Kyl
inH
ors
esh
oe
Cov
e, B
od
ega
Bay
, CA
, U
SA; J
an 1
9, 1
993
--
AF
2573
96Li
n e
t al.
(200
1a)
Nie
nbu
rgia
bor
eali
s
(Kyl
in)
Kyl
inM
osq
uit
o B
ay, W
ash
ingt
on
, U
SA; J
ul 2
, 199
8-
-A
F25
7398
Lin
et a
l. (2
001a
)
Ph
ycod
rys
adam
siae
S.
-M
. Lin
& W
. A. N
elso
nB
lan
d B
ay, N
ort
h Is
lan
d,
New
Zea
lan
d; D
ec 3
0, 2
007
-
W
ELT
A02
8764
GQ
4799
40Li
n a
nd
Nel
son
(2
010)
Ph
ycod
rys
anta
rcti
ca
(Sko
ttsb
erg)
Sko
ttsb
erg
Wen
dy'
s R
ock
, Bo
rrad
ile I
., B
alle
ny
Is
lan
ds,
An
tarc
tica
; Feb
22,
200
6-
-G
Q47
9932
Lin
an
d N
elso
n
(201
0)
Ph
ycod
rys
aust
roge
orgi
ca
Sko
ttsb
erg
Her
o In
let,
Ave
rs I
.,
An
tarc
tica
; Ap
r 19
, 200
3-
-G
Q47
9930
Lin
an
d N
elso
n
(201
0)
Ph
ycod
rys
fim
bria
ta (
Ku
ntz
e)
Kyl
inC
ape
No
sap
pu
, Ho
kkai
do,
Ja
pan
; Ju
l 26,
200
2-
-G
Q47
9929
Lin
an
d N
elso
n
(201
0)
Algae 2014, 29(1): 1-13
http://dx.doi.org/10.4490/algae.2014.29.1.001 10
Tabl
e 1.
Con
tinue
d
Ta
xa
C
oll
ecti
on
info
rmat
ion
Hab
itat
(dep
th /
sub
stra
tum
)Vo
uch
er
Gen
Ban
k ac
cess
ion
No.
R
efer
ence
s
Ph
ycod
rys
fran
iae
S.
-M
. Lin
& W
. A. N
elso
nM
arfe
lls B
each
, Mar
lbo
rou
gh,
Sou
th I
., N
ew Z
eala
nd
; Dec
7, 2
007
-W
ELT
A02
3691
GQ
4799
41Li
n a
nd
Nel
son
(2
010)
Ph
ycod
rys
nov
ae-z
elan
dia
e
S. -
M. L
in &
W. A
. Nel
son
Mar
fells
Bea
ch, M
arlb
oro
ugh
, So
uth
I.,
New
Zea
lan
d; D
ec 7
, 200
7-
WE
LT A
0235
03G
Q47
9934
Lin
an
d N
elso
n
(201
0)
Ph
ycod
rys
ovif
olia
(K
ütz
ing)
M
. J. W
ynn
eIs
la M
ance
rra,
Bah
ia c
ora
l,
Pro
v. V
ald
ivia
, Ch
ile; J
an 1
1, 1
998
--
AF
2574
23Li
n e
t al.
(200
1a)
Ph
ycod
rys
quer
cifo
lia
(B
ory
de
Sain
t-V
ince
nt)
Sk
ott
sber
g
Ro
oke
ry B
ay, S
tan
ley,
E
. Fal
klan
d Is
lan
ds;
Jan
4, 1
998
--
AF
2574
24Li
n e
t al.
(200
1a)
Ph
ycod
rys
rad
icos
a (O
kam
ura
)
Yam
ada
& In
agak
iG
ingd
ao, S
hu
ntu
ng
Pen
insu
la,
Ch
ina;
Jun
23,
199
4-
-A
F25
7427
Lin
et a
l. (2
001a
)
Ph
ycod
rys
rigg
ii N
. L. G
ard
ner
Kit
tiln
goo
k B
ay, S
t. L
awre
nce
Isla
nd
, A
lask
a, U
SA; J
ul 5
, 199
6-
-A
F25
7430
Lin
et a
l. (2
001a
)
Ph
ycod
rys
rube
ns
(Lin
nae
us)
B
atte
rsW
est A
ngl
e B
ay, P
emb
roke
shir
e,
UK
; Ju
l 22,
199
7-
-A
F25
7429
Lin
et a
l. (2
001a
)
Pol
yneu
ra b
onn
emai
son
ii (
C. A
gard
h)
Mag
gs &
Ho
mm
ersa
nd
Ile
Vert
e, R
osc
off
, Bri
ttan
y,
Fran
ce; J
un
22,
199
3-
-A
F25
7437
Lin
et a
l. (2
001a
)
Pol
yneu
ra la
tiss
ima
(Har
vey)
K
ylin
Seal
Ro
ck, O
rego
n,
USA
; May
16,
199
9-
-A
F25
7438
Lin
et a
l. (2
001a
)
Pol
yneu
ra ja
pon
ica
(Yam
ada)
M
ikam
iO
har
a, C
hib
a, Ja
pan
; Mar
26,
200
8-
-C
Q47
9943
Lin
et a
l. (2
001a
)
Wom
ersl
eya
mon
anth
os (
J. A
gard
h)
Pa
pen
fuss
Pt.
Lan
sdal
e, V
icto
ria,
A
ust
ralia
; Ju
l 30,
199
5-
-A
F25
7457
Lin
et a
l. (2
001a
)
Kang & Kim Erythroglossum hyacinthinum sp. nov.
11 http://e-algae.kr
Tabl
e 2.
Com
paris
on o
f dis
tingu
ishi
ng m
orph
olog
ical
cha
ract
ers
amon
g Er
ythr
oglo
ssum
hya
cint
hinu
m s
p. n
ov. a
nd p
utat
ive
rela
tives
Ch
arac
teri
stic
sE
. hya
cin
thin
um
sp
. nov
.
E. l
atu
m
E. m
inim
um
E. p
inn
atu
m
Pol
yneu
ra ja
pon
ica
Hei
ght
3-6
cmU
p to
10
cmU
p to
2.5
cm
5-10
cm
Up
to 1
2 cm
Wid
th o
f bla
de
1.5-
2.5
cm1.
2 cm
1-2
mm
5-10
mm
1.0-
1.5
cm
Bla
de
shap
e E
llip
tica
l to
ob
ovat
eLa
nce
ola
te to
lin
ear
la
nce
ola
teLi
nea
r o
r le
nea
r-la
nce
ola
teB
road
ly li
nea
rB
road
ly li
nea
r
Shap
e o
f lat
eral
bra
nch
Elli
pti
cal t
o o
bov
ate
Bro
ad la
nce
ola
te to
ovo
idO
bov
ate
or
linea
r-o
blo
ng
Bro
adly
lin
ear
Bro
adly
lin
ear
Bla
de
mar
gin
Nu
mer
ou
s m
icro
sco
pic
d
enta
teW
eakl
y u
nd
ula
teA
few
mic
rosc
op
ic te
eth
Min
ute
teet
hIr
regu
larl
y fi
ne
teet
h
Bas
e o
f lat
eral
bra
nch
No
t tap
erin
gSl
igh
tly
tap
erin
gTa
per
ing
Tap
erin
g to
cu
nea
te b
ase
No
t tap
erin
g
Bra
nch
ing
pat
tern
Sim
ple
or
di-
tric
ho
to-
mo
usl
yA
lter
nat
ely
pin
nat
eP
inn
ate
wit
h 1
-2 ti
mes
Pin
nat
e w
ith
2-3
tim
esR
epea
ted
ly d
i-tr
i-p
in-
nat
ely
Ho
ldfa
sts
Dis
coid
, cyl
ind
rica
l cr
eep
ing
bra
nch
-R
oo
t pro
cess
es fr
om
bas
al
ud
er-s
urf
ace
and
mar
gin
Scu
tate
dis
cSc
uta
te d
isc,
sle
nd
er
sto
lon
-lik
e ro
ots
Stip
eC
ylin
dri
cal
--
-C
ylin
dri
cal
Vein
sM
idri
b, a
lter
nat
e ve
ins
Mid
rib
Ab
sen
tM
idri
bM
idri
b, a
lter
nat
e ve
ins
Bla
de
stru
ctu
reM
on
ost
rom
atic
exc
ept
vein
sP
oly
stro
mat
ic e
xcep
t bla
de
mar
gin
Mo
no
stro
mat
ic e
xcep
t me-
dia
n p
ort
ion
Poly
stro
mat
ic e
xcep
t b
lad
e m
argi
nPo
lyst
rom
atic
exc
ept
bla
de
mar
gin
Irid
esce
nce
B
uli
sh-v
iole
t-
--
-
Typ
e lo
cali
ty
Ch
uja
do,
Jeju
, Ko
rea
Shio
yaza
ki, F
uku
shim
a,
Jap
anC
hib
a, Ja
pan
Ch
iba,
Jap
anO
har
a, C
hib
a, Ja
pan
Ref
eren
ces
Th
is s
tud
y Yo
shid
a an
d M
ikam
i (19
97)
Oka
mu
ra (
1932
b),
Mik
ami (
1976
),
Yosh
ida
(199
8)
Oka
mu
ra (
1932
b),
Mik
ami (
1977
),
Yosh
ida
(199
8)
Yam
ada
(193
0),
Oka
mu
ra (
1932
a),
Mik
ami (
1973
)
Algae 2014, 29(1): 1-13
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eronema japonica in 1930, and was then transferred to
the genus Nienburgia in 1935 by Kylin (Yamada 1930, Mi-
kami 1973). Later, Mikami (1973) confirmed that this spe-
cies had Phycodrys-type apical organization and Polyneu-
ra-type procarp. At that time, Mikami was aware that the
specimens had both midrib and lateral veins; however, he
transferred the species to the genus Polyneura citing the
presence of a midrib in Polyneura gmelinii (J. V. Lamour-
oux) Kylin. In Korea, Kim and Nam (1994) mentioned the
possibility of establishing a new taxonomic group for P.
japonica and P. gmelinii because these species have a
midrib, lateral veins, and distinctive apical organization
differently other species of Polyneura. However, P. gmeli-
nii is currently treated as a taxonomic synonym of Eryth-
roglossum laciniatum (Lightfoot) C. A. Maggs & Hommer-
sand (Maggs and Hommersand 1993). The results of our
molecular analyses demonstrate the possibility that P. ja-
ponica is more closely related with the genus Erythroglos-
sum than Polyneura (Fig. 3). To fully resolve this issue, we
need more morphological and molecular evidence from
other taxonomic groups, including the type species of
each genus, Erythroglossum, Polyneura, and Sorella.In conclusion, we collected a new species that has
similar morphological features with P. japonica. To con-
firm the taxonomic position of that species, we collected
other species of the tribe Phycodryeae and performed
morphological and molecular analyses. As the result, we
identified a new species, E. hyacinthinum sp. nov. In ad-
dition, our molecular phylogenetic results highlight the
problems of taxonomic position in the genera of the tribe
Phycodryeae, namely showing the polyphyletic taxon
among Polyneura-type procarp group.
ACKNOWLEDGEMENTS
We thank the staff of Dadohaehaesang National Park
for assisting with transportation and underwater guid-
ance, and we thank Miss H. S. Choi for her transla-
tion of Japanese articles. This work was supported by a
grant from the National Institute of Biological Resources
(NIBR), as funded by the Ministry of Environment (MOE)
of the Republic of Korea (NIBR No. 2013-02-001 for the
collecting samples, and 1834-302 for molecular analyses).
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