New red algal species, Erythroglossum hyacinthinum ... · PDF fileThis alga resembles...

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


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


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


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.


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


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


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


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

)


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


quibus algarum regnum constituitur. Vol. 3, part 3. De

dispositione Delesseriearum curae posteriores. CWK Gle-

erup, Lund, 239 pp.

Díaz-Tapia, P., Berecibar, E., Bárbara, I., Cremades, J. & San-

tos, R. 2009. Biology and taxonomic identity of Eryth-

roglossum lusitanicum (Delesseriaceae, Rhodophyta)

from the Iberian Peninsula. Bot. Mar. 52:207-216.

Guiry, M. D. & Guiry, G. M. 2013. AlgaeBase. World-wide

electronic publication, National University of Ireland,

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anatomy of Polyneura japonica (Yamada) Mikami (De-

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trib. nov. (Delesseriaceae, Rhodophyta) from southern

Australia. Phycologia 40:135-146.

Lin, S. -M., Hommersand, M. H. & Nelson, W. A. 2007. An

assessment of Haraldiophyllum (Delesseriaceae, Rho-

dophyta), inclucing H. crispatum (J.D. Hooker et Har-

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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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