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Molecular evidence for verifying the distribution of Chondracanthus chamissoi and C. teedei (Gigartinaceae, Rhodophyta)


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Chondracanthus teedei and C. chamissoi are regarded as cosmopolitan and endemic species, respectively. To verify the geographic distribution of these two species, we analyzed specimens of C. teedei from the Western Pacific and the Atlantic coast of Spain, as well as of C. chamissoi from Chile using plastid rbcL and mitochondrial COI genes. The phylogenetic tree of rbcL revealed that “C. teedei” from Asia and Chondracanthus sp. from France are conspecific with C. chamissoi from Chile, but distinct from the clade of C. teedei from the Atlantic. These results indicate that C. chamissoi is not exclusively distributed in the southeastern Pacific, but is also found in Korea/Japan and France, whereas C. teedei is found in the Atlantic and not in the western Pacific region. This study demonstrated that the range of C. chamissoi is wider than previously thought, raising interesting questions regarding the transportation vector and the absence of C. teedei from Korea and Japan. In addition, we confirmed that molecular analyses can be used to examine the geographic distribution of marine macroalgae.
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Botanica Marina 2015; 58(2): 103–113
*Corresponding author: Myung Sook Kim, Department of Biology,
Jeju National University, 102 Jejudaehakno, Korea,
Mi Yeon Yang: Department of Biology, Jeju National University, 102
Jejudaehakno, Korea
Erasmo C. Macaya: Laboratorio de Estudios Algales (ALGALAB),
Departmento de Oceanografía, Casilla 160-C, Universidad de
Concepción, Concepción, Chile
Mi Yeon Yang, Erasmo C. Macaya and Myung Sook Kim*
Molecular evidence for verifying the distribution
of Chondracanthus chamissoi and C. teedei
(Gigartinaceae, Rhodophyta)
Abstract: Chondracanthus teedei and C. chamissoi are
regarded as cosmopolitan and endemic species, respec-
tively. To verify the geographic distribution of these two
species, we analyzed specimens of C. teedei from the West-
ern Pacific and the Atlantic coast of Spain, as well as of
C. chamissoi from Chile using plastid rbcL and mitochon-
drial COI genes. The phylogenetic tree of rbcL revealed that
C. teedei” from Asia and Chondracanthus sp. from France
are conspecific with C. chamissoi from Chile, but distinct
from the clade of C. teedei from the Atlantic. These results
indicate that C. chamissoi is not exclusively distributed in
the southeastern Pacific, but is also found in Korea/Japan
and France, whereas C. teedei is found in the Atlantic and
not in the western Pacific region. This study demonstrated
that the range of C. chamissoi is wider than previously
thought, raising interesting questions regarding the trans-
portation vector and the absence of C. teedei from Korea
and Japan. In addition, we confirmed that molecular anal-
yses can be used to examine the geographic distribution
of marine macroalgae.
Keywords: Chondracanthus; C. chamissoi; C. teedei; COI;
distribution; rbcL; Rhodophyta.
DOI 10.1515/bot-2015-0011
Received 26 January, 2015; accepted 5 March, 2015; online first 21
March, 2015
Molecular studies have provided evidence for a change
in the known distribution of macroalgae; thus, several
studies have been performed to identify cryptic species
composed of regional groups among widespread species
(Zuccarello et al. 2002, Saunders and Lehmkuhl 2005,
Le Gall and Saunders 2010b). For example, Zuccarello
et al. (2002) reported two distinct lineages in the red
cosmopolitan alga, Caulacanthus ustulatus (Mertens ex
Turner) Kützing, in the Pacific and Atlantic on the basis
of the sequences from cox2–3 spacer and rubisco spacer.
Another cosmopolitan species, Plocamium cartilagineum
(Linnaeus) Dixon, has been found in at least eight diver-
gent cryptic species from different geographic loca-
tions using large subunit ribosomal RNA gene analysis
(Saunders and Lehmkuhl 2005).
Alternatively, an endemic species may actually be
widespread. For example, the genus Capreolia Guiry and
Womersley, considered endemic to Australasia, has been
found on the coast of central Chile, based on rbcL and cox1
sequences and morphological analysis (Boo et al. 2014).
Pyropia koreana (M.S. Hwang et I.K. Lee) M.S. Hwang, H.G.
Choi, Y.S. Oh et I.K. Lee, described previously from Korea,
has been reported in the Mediterranean Sea (Vergés etal.
2013) and New Zealand (Nelson et al. 2014) using rbcL
analyses. Molecular studies have also been conducted to
examine those species that have been introduced in many
regions (Verlaque etal. 2005, Nelson etal. 2013, Raffo etal.
2014). These results indicate that molecular studies could
aid in uncovering the geographic distribution of macroalgal
species, focusing on cosmopolitan and endemic species.
The red algal genus Chondracanthus Kützing (Gigar-
tinaceae) contains approximately 20 species, with the
majority distributed in the Pacific Ocean (Hommersand
etal. 1994). The genus has a soft to firmly cartilaginous
thallus, which is bladelike to variously branched, and
bears reproductive structures on the ordinary branches,
pinnules, and spines (Hommersand et al. 1994). The
species within the genus are mainly distinguished on the
basis of morphological features, such as the thallus shape,
size, colour, texture, and branching patterns (Hommer-
sand etal. 1994). However, the similarity and high degree
of plasticity in the morphology of Chondracanthus have
led to controversy over the taxonomy of the genus and
related genera (Hommersand etal. 1994).
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104 M.Y. Yang etal.: Distribution of two Chondracanthus species
Chondracanthus teedei is a widespread species as
it has been reported in Asia, Europe, North and South
America, and Africa (Guiry and Guiry 2014). Recently,
the specimens recognized as C. teedei from Bermuda has
been assigned to a new species, Chondracantus saunder-
sii, using the rbcL marker (Schneider and Lane 2005), but
the taxonomic status and geographic distribution of C.
teedei from other regions have not been explored using
molecular analyses. In comparison, the endemic species,
Chondracanthus chamissoi (C. Agardh) Kützing, is found
along the Pacific coast of South America (from 5° to 42°S;
Ramírez and Santelices 1991). This species has been har-
vested commercially in Chile as a raw material for car-
rageenans and exported for human consumption to the
Asian market (Bulboa etal. 2013).
Hughey and Hommersand (2008) employed rbcL
and ITS sequences to investigate the taxonomic status of
Chondracanthus species from the Pacific coast of North
America. Although Atlantic C. teedei was included in the
molecular analysis (Hommersand etal. 1994), the taxon-
omy of species from the Northwest Pacific has relied on
morphological features (Mikami 1965, Kang 1968). Accord-
ing to Hommersand etal. (1994), the centre of speciation
of the genus was East Asia and Pacific North America, but
it later spread into the Atlantic Ocean.
The main objective of the present study was to clarify
the geographic distribution of two Chondracanthus species
using plastid rbcL and mitochondrial COI (5 end of cyto-
chrome c oxidase I gene). More specifically, our goal was
1) to compare the taxonomic status of the widespread
species, C. teedei, by analyzing samples from Spain and
the Northwest Pacific; 2) to confirm the distribution of C.
chamissoi, which is known to be endemic in the South-
eastern Pacific; and 3) to describe in detail the morphol-
ogy of specimens from the Northwest Pacific region. In
addition, we explored the phylogenetic relationships of
related species in the genus Chondracanthus.
Materials and methods
We obtained samples of Chondracanthus across its dis-
tributional range of the Northwest Pacific and Atlantic
(Figures 1–6). We collected “C. teedei” from Korea and
Japan (Figures 3–5) and C. teedei from Spain (Figure 6).
Chondracanthus chamissoi was collected from the type
locality, Chile (Figures 1 and 2). Information on the col-
lection sites for the samples used in this study is pre-
sented in Table 1. Field-collected samples were put into
an icebox with seawater and transported to the laboratory.
Specimens were pressed onto herbarium paper, and por-
tions of dried specimens were deposited in silica gel for
molecular analyses. Voucher specimens were deposited
in the herbarium of Jeju National University, Jeju, Korea.
Samples for anatomical studies were fixed in 5% forma-
lin/seawater and were sectioned by hand or using a freez-
ing microtome (NK-101-II; Nippon Optical Works Co., Ltd.,
Tokyo, Japan). Sections were stained with 1% aqueous
aniline blue acidified with a drop of 1% HCl and mounted
in 40% Karo corn syrup. Photomicrographs were taken
using a QImaging 1394 camera (QImaging, Surrey, BC,
Canada) attached to a BX50 microscope (Olympus, Tokyo,
Japan). All images were imported into the Adobe Photo-
Shop 5.5 software (Adobe Systems Inc., San Jose, CA, USA)
for plate assembly.
DNA extraction, PCR amplification, and sequenc-
ing were performed as described by Yang et al. (2013).
Specific primer pairs for the rbcL gene were rbcLF145-
rbcLR898 and rbcLF762- rbcLR1442 (Kim etal. 2010). For
the COI gene, we used GazF2 (Lane etal. 2007) and GazR1
(Saunders 2005) or GWSFn (Le Gall and Saunders 2010a)
and GWSRx (Clarkston and Saunders 2012). Sequences
obtained from rbcL and COI genes were edited using
Chromas version 1.45 software (Technelysium Pty Ltd,
South Brisbane, QLD, Australia). Multiple sequence align-
ments were made in BioEdit (Hall 1999) and aligned man-
ually. We assessed the level of variation in the sequences
of two genes; uncorrected (p) pair-wise genetic distances
were estimated using the MEGA 4.0 software (Tamura
etal. 2007). To construct the tree of rbcL and COI dataset,
we performed maximum likelihood (ML) analysis based
on the GTR + Γ evolutionary model using RAxML version
7.2.6 software (Stamatakis 2006). To identify the best tree,
we constructed 200 independent tree inferences using the
-# option with default -I (automatically optimized Subtree
Pruning-Regrafting rearrangement) and -c (25 distinct rate
categories) software options. We performed 1000 replica-
tions using the same software and settings to generate
bootstrap values for each gene.
Molecular analyses
RbcL sequence data of Chondracanthus were determined
for 41 specimens from this study and from GenBank,
including two outgroups: Gigartina grandifida and Rho-
doglossum gigartinoides (Table 1). A total of 21 sequences
were generated in this study. The length of rbcL sequences
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M.Y. Yang etal.: Distribution of two Chondracanthus species105
was 1395 bp, the number of variable sites was 282, and
182 (64%) sites were phylogenetically informative. In the
phylogenetic tree of rbcL (Figure 7), five specimens of “C.
teedei” from Korea and Japan, 12 specimens of C. chamis-
soi from Chile, and JQ405738 from France were clus-
tered as conspecifics with 100% bootstrap supporting
value. The highest intraspecific sequence divergence was
0.7% between the specimens from Chile (AF146193) and
Korea (KP059087). No sequence divergence was detected
between “C. teedei” from Japan and Chondracanthus sp.
from France (JQ405738). Chondracanthus teedei from
Spain was grouped with previously published sequences
from France (U03024) and Brazil (U02945). The clade of C.
teedei from three regions, including the Atlantic coasts of
Spain, showed intraspecific variation of 0.9% with 100%
bootstrap supporting value. “Chondracanthus teedei” from
the Northwest Pacific (Korea and Japan) differed from
the Spanish C. teedei with 2.8% sequence divergence and
from the French Chondracanthus sp. (JQ405738) with 4.0%
divergence. “Chondracanthus teedei” from Korea and Japan
was also separated from C. intermedius and C. tenellus from
Japan with 5.2–5.8% sequence divergences. Chondracan-
thus chamissoi was separated from Atlantic C. teedei with
2.7–3.8% sequence divergence. The clade of C. teedei was
sister to the Pacific species, namely, C. exasperatus, C. spi-
nosus and C. squarrulosus, with 2.0–2.9% divergences. The
low interspecific divergence among closely related species
was 0.1% between C. kjeldsenii and C. canaliculatus, 0.3%
between C. exasperatus and C. spinosus, 0.4% between
C. canaliculatus and C. bajacalifornicus, and 0.7% between
C. serratus and C. bajaclifornicus. The average value of
interspecific divergence was 4.7% in the rbcL gene.
COI sequence data were aligned for 20 specimens of
Chondracanthus from Korea, Japan, Spain, Chile, Canada
and USA. A total of 17 sequences were newly generated
in this study (Table 1). The length of COI sequences was
664 bp and the number of variable sites was 85, where
23 (27%) were phylogenetically informative. In the ML
tree of COI (Figure 8), five specimens of “C. teedei” from
Korea and Japan and eight of C. chamissoi from Chile were
Figures 1–6:External morphology of two Chondracanthus species, C. chamissoi and C. teedei, used in this study. (1) C. chamissoi, habit of
thalli growing in the intertidal zone from Chile. (2) C. chamissoi, specimen from Chile. (3) C. chamissoi, specimen from Busan, Korea, veg-
etative plant. (4) C. chamissoi, specimen from Misaki, Japan, female plant. (5) C. chamissoi, specimen from Enoshima, Japan, female plant.
(6) C. teedei, specimen from Spain, vegetative plant. Scale bars: 1–4=5 cm, 5=2 cm, 6=3 cm.
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106 M.Y. Yang etal.: Distribution of two Chondracanthus species
Table 1:List of taxa used in the rbcL and COI analyses, their sources and GenBank accession numbers.
Collection information
GenBank accession number
Chondracanthus acicularis (Roth) Fredericq Roscoff, Brittany, France, ( March ) U Hommersand etal. 
Roche, Cádiz, Andalucía, Spain ( April ) KP This study
Masonboro Inlet, New Hanover County, North California,
USA ( March )
KJ Unpublished
C. bajacalifornicus Hughey et Hommersand Baja California, Mexico ( July ) DQ Hughey and Hommersand 
C. canaliculatus(Harvey) Guiry Monterey Co., California ( July ) DQ Hughey and Hommersand 
C. chamissoi (C. Agardh) Kützing Lechagua, Chiloé, Chile ( February ) AF Hughey and Hommersand 
Cocholgüe, Biobío Region, Chile ( February ) KP This study
Cocholgüe, Biobío Region, Chile ( February ) KP KP This study
Cocholgüe, Biobío Region, Chile ( February ) KP This study
Cocholgüe, Biobío Region, Chile ( February ) KP KP This study
Iquique, Tarapacá Region, Chile( November ) KP This study
Iquique, Tarapacá Region, Chile ( November ) KP This study
Chonchi, Los Lagos Region, Chile ( February ) KP This study
Chonchi, Los Lagos Region, Chile ( February ) KP KP This study
Chonchi, Los Lagos Region, Chile ( February ) KP This study
Puerto Aldea, Coquimbo Region, Chile ( January ) KP This study
Puerto Aldea, Coquimbo Region, Chile ( January ) KP KP This study
Lebu, Biobío Region, Chile ( February ) KP KP This study
Huayquique, Tarapacá Region, Chile ( November ) KP KP This study
Gwangan-ri, Busan, Korea ( December ) KP KP This study
Gwangan-ri, Busan, Korea ( December ) KP KP This study
Misaki, Chiba Pref. Japan ( April ) KP KP This study
Enoshima, Kanagawa Pref. Japan ( March ) KP KP This study
Enoshima, Kanagawa Pref. Japan ( March ) KP KP This study
Toulindac, Gulf of Morbihan, France ( May ) JQ Mineur etal. 
C. chapmanii (J.D. Hooker et Harvey) Fredericq Wellington, New Zealand ( May ) U Hommersand etal. 
C. corymbiferus (Kützing) Guiry Kitsap Co., Washington ( June ) DQ Hughey and Hommersand 
Canada GQ Le Gall and Saunders a
C. exasperatus (Harvey et Baily) Hughey Baja California, Mexico ( July ) DQ Hughey and Hommersand 
Canada GQ Le Gall and Saunders a
C. harveyanus (Kützing) Guiry Sonoma Co., California ( July ) DQ Hughey and Hommersand 
C. intermedius (Suringar) Hommersand Chiba Pref., Japan ( May ) U Hommersand etal. 
C. kjeldsenii Hughey et Hommersand Los Angeles Co., California ( February ) DQ Hughey and Hommersand 
C. saundersii C.W. Schneider et C.E. Lane Hamilton Parish, Bermuda AY Schneider and Lane 
C. serratus (Gardner) Hughey et Hommersand San Diego Co., California ( March ) DQ Hughey and Hommersand 
C. spinosus (Kützing) Guiry U Hommersand etal. 
C. squarrulosus (Setchell et Gardner) Hughey, Silva et
Gulf of California, Mexico () DQ Hughey and Hommersand 
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M.Y. Yang etal.: Distribution of two Chondracanthus species107
Collection information
GenBank accession number
C. teedei (Martens et Roth) Kützing Roscoff, Brittany, France ( March ) U Hommersand etal. 
Itanhaém, Estado de Sào Paulo, Brazil (April ) U Hommersand etal. 
San Fernando, Cádiz, Spain ( April ) KP KP This study
San Fernando, Cádiz, Spain ( April ) KP KP This study
San Fernando, Cádiz, Spain ( April ) KP KP This study
San Fernando, Cádiz, Spain ( April ) KP KP This study
C. tenellus (Harvey) Hommersand Tateyama Bay, Chiba Pref. Japan ( June ) AF Hughey and Hommersand 
Gigartina grandifera J. Agardh Waitai West, Chatham Is., New Zealand DQ Unpublished
Rhodoglossum gigartinoides (Sonder) Edyvane et
Flinders Jetty, Victoria, Australia JN Schneider etal. 
Table 1 (Continued)
included in one clade with 90% bootstrap value, and
they showed 0–0.9% intraspecific divergence. Within this
clade, three intra-clades were found: one with samples
from Korea and Japan, and two others from Chile. The
intra-clade from Asia displayed 0.3% sequence variation,
whereas there was 0.8–0.9% variation between the two
intra-clades from Chile. This clade, including samples
from Korea, Japan and Chile, was quite distinct from
Spanish C. teedei, showing no intraspecific sequence vari-
ation, with 4.1–4.7% sequence divergence. The average
value of interspecific divergence was 7.0% in the COI gene.
Morphological analysis
The thalli of Chondracanthus chamissoi from Korea and
Japan were solitary, erect or occasionally prostrate,
9–25 cm in height, flaccid cartilaginous in texture, and
reddish purple or sometimes yellowish, arising from small
discoid holdfasts (Figures 3–5). Main axes are slightly or
broadly flattened, 2–6mm in width, and alternately or
oppositely branched at the margin. Branches are slightly
flexuose, acuminate, with shorter and longer inter-
mixed, and covered in short spine-like ramuli. The tips
of spine-like ramuli sometimes have a swelling (Figure
9). Anatomical structures are multiaxial and composed
of cortical, subcortical, and medullary layers. The cortex
is composed of densely arranged pigment cells consist-
ing of 5–6 rows of globular to ellipsoidal cells (Figure 10).
The surface cells are elongated in transverse sections and
are rounded in surface view (Figures 10, 11). The subcorti-
cal layer consists of 2–3 series of rather loosely arranged
irregular cells (Figure 12). The medullary layer consists of
loosely packed filamentous cells (Figure 12). Cystocarps
protrude externally, 950–1700 μm in diameter, subspheri-
cal or hemispherical, and are borne on the side of second-
ary branches or the short ramuli. Mature cystocarps are
composed of a large central carposporangial mass, 600–
1000 μm in diameter (Figure 13). Special medullary fila-
ments are developed through secondary cell division of
the medullary cell and are arranged in a circle around the
gonimoblast masses (Figure 14). Carposporangia are sub-
globuse, 32–45 μm in diameter and are aggregated with
sterile gonimoblasts, which are irregular and polygonal
(Figure14). Spermatangia were not found in the analyzed
material. Tetrasporangial sori appear as small deeply col-
oured swellings on the branch margins and are embedded
on the secondary or tertiary axes (Figure 15). Tetraspo-
rangia are formed in intercalary chains in the subcortex
(Figure15). Mature tetrasporangia were cruciately divided,
and 15–20μm × 28–40 μm in size (Figure 16).
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108 M.Y. Yang etal.: Distribution of two Chondracanthus species
This study redefined the geographic distribution of two
Chondracanthus species: C. teedei has been known as a
widespread species (Hommersand et al. 1994), whereas
C. chamissoi is endemic in Chile and Peru (Ramírez and
Santelices 1991). Our analyses of the plastid rbcL and
mitochondrial COI genes clearly demonstrated that the
specimens considered as “C. teedei” from Korea and
Japan belonged to C. chamissoi, including the specimen
JQ405738 (Minuer etal. 2012) from France. Therefore, we
confirmed that the incorrect record of “C. teedei” in the
Figure 7:Maximum likelihood phylogenetic tree of Chondracanthus inferred from rbcL sequences. Bootstrap values for ML are shown for
each clade. Bold type indicates sequences generated in the present study. Scale indicates substitutions per site.
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M.Y. Yang etal.: Distribution of two Chondracanthus species109
western Pacific was due to misidentification arising from
very similar morphology (Mikami 1965, Dixon and Irvine
1977). At the same time, the occurrence of C. chamissoi in
this region has been verified as a new distribution record.
Our collections of C. teedei from Spain were clustered as
a clade with the samples from France and Brazil (U03024
and U02945 in Hommersand etal. 1994) in our rbcL tree.
Chondracanthus chamissoi was originally described
as Sphaerococcus chamissoi C. Agardh (type material from
Chile), and then once assigned to Gigartina chamissoi
(C. Agardh) J. Agardh (1842). This species is known as an
endemic species in the Southeastern Pacific, having been
described from Peru (5°S) to Chile (40°S) and has a highly
variable morphology (Acleto 1986, Ramírez and Santelices
1991). In the present study, the specimens assigned to
C. chamissoi from the Northwest Pacific, France and Chile
also showed morphological variations (Figures 1–5),
similar to those reported by Mineur et al. (2012). The
specimens from the Northwest Pacific (Korea and Japan)
have been confused with C. teedei (Mikami 1965, Kang
1968, Yoshida 1998, Lee and Kang 2001) because they
share some morphological features, including plant size,
flattened fronds with similar width of main axes, cruci-
ately divided tetrasporangia, and protruding cystocarps
on the branches (Mikami 1965, Dixon and Irvine 1977).
However, the cystocarps of C. chamissoi from Korea/Japan
and Chile are borne on the side of branches, whereas C.
teedei from Europe have cystocarps in the apex of lateral
branches (Table 2).
The occurrence of Chondracanthus chamissoi on
the Atlantic coasts was confirmed in the present study.
Mineur et al. (2012) reported four new exotic red algae
on European shores, including one Chondracanthus
species (as Chondracanthus sp., JQ405738). According to
Mineur et al. (2012), Chondracanthus sp. resembled the
native C. teedei, but the rbcL sequences were related to C.
chamissoi from southern Chile. The rbcL sequence from
Chondracanthus sp. (JQ405738) was conspecific with C.
chamissoi from Korea, Japan, and Chile (Figure 7). In this
study, the sequences from France (JQ405738) were iden-
tical to those from Japan. Therefore, C. chamissoi is not
endemic to South America, and is distributed globally:
South America (Chile, Peru), East Asia (Korea, Japan), and
Europe (France).
Figure 8:Maximum likelihood tree of Chondracanthus inferred from COI sequences. Bold type indicates sequences generated in the
present study. Scale indicates substitutions per site.
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110 M.Y. Yang etal.: Distribution of two Chondracanthus species
The rbcL data supported the monophyly of Chond-
racanthus (Hommersand et al. 1994). Chondracanthus
chamissoi is closely related to the clade including C.
teedei, but is not as closely related to the other two western
Pacific species, C. tenellus and C. intermedius (Figure 7).
Although C. chamissoi shares a distribution range with
C. tenellus and C. intermedius in the western Pacific, it is
distinguished from the latter two species by having often
broadly flattened main axes and spine-like branchlets on
the margin of the frond (Mikami 1965). The interspecific
sequence divergences among the eastern Pacific species
were conspicuously low, ranging from 0.1% (C. kjeldsenii
and C. canaliculatus) to 0.7% (C. serratus and C. bajacali-
fornicus). These values were lower than the intraspecific
divergence of several species, such as of C. chamissoi
(0.7%), C. teedei (0.9%) and C. acicularis (0.6%). Low
interspecific divergence was also apparent in the genus
Mazzaella (0.9%) of the family Gigartinaceae (Hughey and
Hommersand 2010).
To support species boundaries using molecular data,
it is necessary to include other genes that exhibit more
variation, such as COI (Yang etal. 2013). Our COI data set
including five species of Chondracanthus revealed higher
interspecific divergences than the rbcL sequences. By
Figures 9–16:Chondracanthus chamissoi from Korea and Japan. (9) Spine-like ramuli with swelling at the tips. (10) Transverse section of
thallus showing cortical and subcortical layers. (11) Surface view showing rounded cortical cells. (12) Transverse section of thallus showing
filamentous medullary cells. (13) Transverse section of cystocarp showing large central carposporangial mass surrounded by special medul-
lary filaments. (14) Close-up of carposporangial mass. (15) Transverse section of tetrasporophyte. (16) Cruciately divided mature tetraspo-
rangia. Scale bars: 9, 12 and 14=500 μm, 10=50 μm, 11=30 μm, 13 and 15=300 μm, 16=100 μm.
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M.Y. Yang etal.: Distribution of two Chondracanthus species111
Table 2:Comparison of morphological characteristics of C. chamissoi from Korea/Japan and Chile and C. teedei from Europe.
C. chamissoi from Korea/Japan C. chamissoi from Chile C. teedei from Europe
Habitat Protected areas, tide pools in
intertidal zone and to m depth in
subtidal zone
Intertidal to m depth in
subtidal zone, especially in
protected habitats
Upper sublittoral to at least m
depth in subtidal zone
Height – cm Up to  cm Up to  cm
Main axes Flattened, –mm width Flattened, –mm width Flattened, –mm width
Tetrasporangia Cruciately divided, –×– μmCruciately divided, ×– μmCruciately divided, –×
– μm
Cystocarps Subspherical or hemispherical,
– μm in diameter, developed
on the side of secondary branches or
the short ramulus
Spherical,  μm in diameter,
developed along the margins of
primary and secondary branches
– μm in diameter,
developed in apices of lateral
Carpospornagial mass– μm in diameter – μm in diameter
Carposporangia Subglobuse, (.) – μmRounded-hexagonal or elongated
hexagonal, – ()×–
() μm
References This study, Mikami  (as C. teedei)Hoffmann and Santelices  Dixon and Irvine , Guiry 
comparing the same taxa using both types of molecular
data, we found that interspecific divergences ranged from
4.5–9.0% in COI and 2.5–7.0% in rbcL. These findings are in
agreement with the results of DNA barcoding in the Gelidi-
ales (Freshwater etal. 2010). Their results indicate that the
COI marker is useful for molecular-assisted identification,
especially in the case of closely related species when the
more conserved rbcL may be uninformative (Freshwater
etal. 2010).
Similar problems of identification are also apparent
in several macroalgal species. Nemalion helminthoides,
which was thought to be a cosmopolitan species, was
actually divided into five distinct genetic lineages by bio-
geographic distribution patterns (Le Gall and Saunders
2010b). Zuccarello etal. (2002) demonstrated that Spyridia
filamentosa from different regions worldwide existed as
several cryptic species. The combined data from biogeo-
graphic patterns and phylogenetic relationships can be
used to increase our understanding of long-distance dis-
persal or red algal evolution (Zuccarello etal. 2002). Our
results indicate that careful observations with molecular
analyses are required to identify species correctly under
the name of C. teedei from other regions.
In conclusion, the present study increases our under-
standing of the geographic distribution of two Chondra-
canthus species. Chondracanthus chamissoi, previously
known as an endemic species in Chile, is now found glob-
ally. Long-distance dispersal of seaweeds may be rein-
forced by inter-oceanic shipping, which may transport
species beyond their historical range (Carlton and Hodder
1995). Although the population analyses with enough
sampling from many different sites were not conducted
in this study, it is possible that C. chamissoi has spread
from Chile to Asian countries because dried and salted
infertile or tetrasporic fronds have been exported during
recent years (Avila et al. 2011). Other dispersal vectors
(e.g., boat traffic) cannot be ruled out because Japan and
Korea represent one of the most important destinations for
maritime traffic from Chile (Boletín Estadístico Marítimo
2014). In addition, given that C. chamissoi can grow under
a wide range of temperatures ranging from 10°C–2 5°C
(Bulboa and Macchiavello 2001), it can easily become
established under different conditions. Secondary attach-
ment discs can help the formation of new individuals and
play an important role as a vegetative propagation strat-
egy (Sáez et al. 2008). Consequently, the cosmopolitan
species C. teedei is not distributed in the western Pacific
region. We discovered the cryptic introduction because
of the morphological similarity among Chondracanthus
species, which can lead to misidentification of the species
in another region. Therefore, it is important to integrate
molecular and morphological analyses to verify macroal-
gal distribution worldwide.
Acknowledgments: We thank Dr. Kikuchi, Dr. Kondo and
Dr. Lacida for helping in the collection of samples in Japan
and Spain, and members of the Molecular Phylogeny of
Marine Algae Laboratory at Jeju National University. This
study was supported by a grant from the National Insti-
tute of Biological Resources (NIBR), funded by the Minis-
try of Environment (MOE) of the Republic of Korea (NIBR
No. 2013-02-001 for collecting samples and NIBR No. 2013-
02-013 for molecular phylogeny of Korean major taxa) and
FONDECYT/CONICYT project N° 11110437 to ECM.
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Mi Yeon Yang
Department of Biology, Jeju National
University, 102 Jejudaehakno, Korea
Mi Yeon Yang is a PhD student at Jeju National University, Jeju,
Korea. She was awarded a MSc in Molecular Phylogeny for her work
on the molecular phylogeny and DNA barcoding of the Gracilari-
aceae (Marine Algal Laboratory, Jeju National University). For her
PhD research, Ms. Yang is working on the red algal order Gigartina-
les from Korea.
Erasmo C. Macaya
Laboratorio de Estudios Algales (ALGALAB),
Departmento de Oceanografía, Casilla 160-C,
Universidad de Concepción, Concepción,
Erasmo C. Macaya has been an Assistant Professor at Concep-
ción University, Chile since 2010. He received a primary degree in
Marine Biology and a Master’s in Marine Sciences from Universidad
Catolica del Norte, Coquimbo, Chile. He obtained a PhD in Marine
Biology from Victoria University, Wellington, New Zealand studying
the dispersal patterns, connectivity, taxonomy and genetic diversity
of the giant kelp, Macrocystis pyrifera. His research focuses on the
different aspects of macroalgae, such as ecology, taxonomy and
phylogeography, among others.
Myung Sook Kim
Department of Biology, Jeju National
University, 102 Jejudaehakno, Korea
Myung Sook Kim is a Professor of Biology at Jeju National Univer-
sity, Jeju, Korea. She was awarded a PhD in Algal Systematics by
the Seoul National University, Korea for her work on the taxonomic
revision of Polysiphonia. She has studied systematics in Rhodo-
phyta for over 15 years, especially in the family Rhodomelaceae.
Her more recent research has concentrated on establishing a DNA
barcode database for Korean seaweeds to identify species and
genus correctly.
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... The taxonomic identification of red algae using only morphological characters is insufficient because of the algae's high plasticity (Saunders, 2005 South Korea and Japan, and Chondracanthus sp. from France have proven to be C. chamissoi, when they were phylogenetically evaluated with mitochondrial and plastidial DNA sequences (Yang, Macaya, & Kim, 2015). Therefore, the known distribution of C. chamissoi has undergone changes and is no longer considered endemic to Peru and Chile. ...
... Le Roux, Stegenga, Verlaque, and Maggs (2012) for their discussion regarding possible origin and vectors of introduction in the Gulf of Morbihan, France, and the study of Yang et al. (2015). Using the same DNA markers, a recent study also revealed no genetic differentiation among specimens from f. lessonii and f. chauvinii sampled in southern Chile (Rodríguez, Tellier, Pérez-Araneda, & Otaíza, 2021). ...
... Our results also confirm that specimens identified morphologically as C. chamissoi from Chile share haplotypes with specimens from Peru, and are closely related (ML = 96, intraspecific distance = 0.005) to C. chamissoi from South Korea and Japan in the COI analysis (Figure 1), and to C. chamissoi from South Korea, Japan, and France in the rbcL analysis ( Figure 2). Therefore, the present study completes the mitochondrial and plastidial phylogenetic analyses by Yang et al. (2015) where material from Chile, Japan, and France were proven to be C. chamissoi. Additionally, three (H1, H4, and H7) of the seven COI haplotypes found on the Peruvian coast and two (R1 and R2) of the three rbcL haplotypes were shared among specimens identified as C. chamissoi and as C. glomeratus. ...
Full-text available
Chondracanthus chamissoi is part of the diet of coastal people from Peru and is exported dehydrated to Asian countries for direct consumption. Although it is considered endemic to Peru and Chile, its range has extended to distant regions, such as Korea, Japan, and France. Using morphological and molecular approaches, we examined specimens from Peru assigned to C. chamissoi (including the taxon of uncertain status Chondracanthus glomeratus) to improve phylogenetic and geographical information and characterize its morphological variability. Twenty‐one localities on the Peruvian coast were sampled, obtaining 102 COI and 27 rbcL sequences. To differentiate both entities, morphological characters such as thallus size, consistency, arrangement of main and secondary axes, branching patterns and location of reproductive structures, were analyzed on 46 specimens. While morphological characteristics are clearly contrasting among the two groups, both COI and rbcL phylogenies revealed a well‐supported clade with no genetic differentiation between the two morphologies. Therefore, the phylogenies indicate that C. chamissoi and C. glomeratus form a single taxonomic entity with high morphological variability, large geographic distribution and at least two morphological forms. The smaller form of C. chamissoi can be identified as C. chamissoi f. glomeratus. Such morphological variability can be of interest for future aquaculture development.
... distribución conocida de C. chamissoi ha sufrido cambios recientes, dejando de ser considerada como endémica de Perú y Chile. Los individuos identificados morfológicamente como C. chamissoi de Chile, C. teedei de Corea del Sur y Japón y Chondracanthus sp. de Francia, han demostrado ser C. chamissoi al ser evaluados filogenéticamente con secuencias de ADN mitocondrial y cloroplástico (Yang et al. 2015). ...
... Esta diversidad debe ser considerada al momento de establecer cultivos, así como ante traslado de material genético entre praderas para repoblamiento, por ejemplo. Futuros estudios también deberían incluir el resto de la distribución geográfica de la especie, tanto la costa chilena como las costas de Japón y Corea del Sur donde se ha confirmado recientemente la presencia de la especie (Yang et al., 2015). ...
... Chondracanthus chamissoi (Rhodophyta, Gigartinaceae) is a common red seaweed along the coasts of the southeastern Pacific Ocean, extending from Per u (5°S) to Ancud, Chile (42°S; Ram ırez and Santelices 1991, Hoffmann andSantelices 1997), although recent molecular evidence indicates that this species is also present in Korea, Japan, and France (Yang et al. 2015). It has been used as a source for the extraction of the commercially important phycocolloid carrageenan (Hoffmann and Santelices 1997), but more recently, it has been exported as an edible seaweed to Asia (Bulboa and Macchiavello 2006). ...
... Nevertheless, molecular analyses done on female gametophytes of both forms collected at the same three localities studied here revealed a single haplotype for the rbcL marker and only two haplotypes for the COI marker. They were identical to sequences in GenBank found by Yang et al. (2015) from C. chamissoi collected in Chile. Moreover, samples of both forms can share the same haplotype and be found in the same locality (Rodr ıguez 2018), supporting the interpretation that the two forms correspond to the same species. ...
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The red seaweed Chondracanthus chamissoi shows high morphological variability. Initially, three species were identified based on the width of the main axis of their blades. Later, all of them were included in a single species with two morphological groups. Recently, quantitative studies demonstrated the existence of two forms in C. chamissoi: f. lessonii and f. chauvinii. It was also shown that these two forms occur in sympatry, growing side by side. These forms were not associated with either a life cycle phase or the sex of the blades. This study aimed to determine whether the two forms could represent different species. We evaluated the forms' taxonomic position using COI and rbcL markers, including samples from three localities in southern Chile. All specimens shared a single rbcL haplotype, whereas the two COI haplotypes differed by four base pairs and were present in blades of both forms and life cycle phases. The two morphological types correspond to intraspecific forms. This species is of commercial importance, and its main market is aimed at human consumption with a marked preference for f. lessonii.
... Chondracanthus chamissoi (C.Agardh) Kützing, 1843, conocida como "yuyo" o "mococho" es la macroalga roja comercial más abundante en el Perú y ha sido registrada desde el intermareal mixto (piedras y arena) hasta profundidades cercanas a los 15 m (Hoffmann y Santelices, 1997) sobre fondo duro con presencia de moluscos bivalvos, galerías de poliquetos y otras macroalgas (observación personal). Esta especie se encuentra en el Océano Pacífico (Corea, Japón, Perú y Chile) y el Mediterráneo (Francia) (Yang et al., 2017). En el Pacífico Suroriental su distribución es continua desde Paita, Perú (5°S) hasta Ancud, Chile (42°S) (Acleto, 1971;Ramírez y Santelices, 1991;Arbaiza, 2016). ...
... Chondracanthus chamissoi (C.Agardh) Kützing, 1843, known locally as yuyo or mococho is the most abundant commercial red macroalgae in Peru and has been recorded from the mixed intertidal (stones and sand) to depths close to 15 m (Hoffmann & Santelices, 1997) on a hard bottom with the presence of bivalve mollusks, polychaetes, and other macroalgae (personal observation). This species is found in the Pacific Ocean (Korea, Japan, Peru, and Chile) and the Mediterranean (France) (Yang et al., 2017). In the Southeast Pacific, its distribution is continuous from Paita, Peru (5°S) to Ancud, Chile (42°S) (Acleto, 1971;Ramírez & Santelices, 1991;Arbaiza, 2016). ...
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Características bioecológicas de la macroalga roja Chondracanthus chamissoi (C. Agardh) Kützing (Rhodophyta, Gigartinaceae) en la zona intermareal del norte del Perú. Bol Inst Mar Perú. 35(2): 271-293.-En La Libertad en invierno y primavera 2018 se estudió a la macroalga roja Chondracanthus chamissoi "yuyo" o "mococho". En Trujillo, Paiján y Pacasmayo se determinaron 21 praderas; se realizaron transectos perpendiculares a la línea de costa y tres cuadratas (0,25 m 2 c/u) en cada transecto; de cada cuadrata se extrajeron los organismos. Las muestras de macroalgas fueron separadas para su análisis, y las de flora y fauna acompañante fueron registradas in situ como presente/ ausente. Se describió tipo de sustrato y características físico-químicas del agua (temperatura, oxígeno y pH). En laboratorio las muestras de C. chamissoi fueron lavadas con agua corriente, se determinó tallas, densidad (ind./m 2), biomasa y se diferenciaron ejemplares con estructuras reproductivas (cistocarpos). Los resultados muestran que la mayoría de praderas se encuentran en orilla de arena y fondo de piedras. Los estadísticos de tendencia central y de dispersión calculados fueron similares. Las densidades en promedio fueron de 322 plantas/m 2 en invierno y 292 plantas/m 2 en primavera. La biomasa en invierno se estimó en 277,77 t y en primavera 650,69 t. El grupo taxonómico con mayor frecuencia fue Rhodophyta (algas rojas) y las especies más representativas C. chamissoi, Asterfilopsis furcellata (=Gymnogongrus furcellata) y Grateloupia doryphora. La biomasa de otras macroalgas se estimó en 11,17 t para Chondracanthus glomeratus "clavo" o "yuyo clavo" en invierno y 3,53 t en primavera, para Gracilariopsis lemaneiformis "pelillo" se estimó en 1,77 t. ABSTRACT Uribe R, Atoche-Suclupe D, Paredes-Paredes J, Seclén J. 2020. Bioecological features of the red macroalgae Chondracanthus chamissoi (C. Agardh) Kützing (Rhodophyta, Gigartinaceae) in the intertidal zone of northern Peru. Bol Inst Mar Peru. 35(2): 271-293.-In winter and spring 2018, we studied the red macroalgae Chondracanthus chamissoi known as yuyo or mococho in La Libertad Region. In Trujillo, Paiján, and Pacasmayo, we determined 21 meadows by performing transects perpendicular to the coastline and three quadrats (0.25 m 2 each) in every transect. We extracted the organisms from each quadrat, then we separated the samples of macroalgae for analysis from those of accompanying flora and fauna which were recorded in situ as present/absent. We described the type of substrate and the physical-chemical characteristics of the water (temperature, oxygen, and pH). In the laboratory, the samples of C. chamissoi were washed with running water. We determined sizes, density (ind./m 2), biomass, and differentiated specimens with reproductive structures (cystocarps). The results show that the majority of the meadows are found on sandbanks and stone bottoms. The central tendency and dispersion statistics calculated were similar. The mean densities were 322 plants/m 2 in winter and 292 plants/m 2 in spring. In winter, the biomass was estimated at 277.77 t and in spring at 650.69 t. The most frequent taxonomic group was Rhodophyta (red algae) and the most representative species were C. chamissoi, Asterfilopsis furcellata (=Gymnogongrus furcellata), and Grateloupia doryphora. The biomass of other macroalgae was estimated at 11.17 t for Chondracanthus glomeratus clavo or yuyo clavo in winter and 3.53 t in spring, for Gracilariopsis lemaneiformis pelillo it was estimated at 1.77 t.
... Recently, the specimens recognised as C. teedei from B e r m u d a h a s b e e n a s s i g n e d t o a n e w s p e c i e s , Chondracanthus saundersii, using the ribulose-1,5bisphosphate carboxylase/oxygenase (RuBisCo) gene (RbcL) marker (Schneider and Lane 2005 Yang et al. (2015), using molecular data (cytochrome c oxidase I gene-COI and RbcL), reported that many samples classified in North Pacific as C. teedei are, in fact, specimens of Chondracanthus chamissoi, previously known as an endemic species in Chile. In their study, specimens from Spain and Brazil were grouped as C . ...
... It has also been reported from Japan (Mikami 1965), the Indian Ocean (Silva et al. 1996), Venezuela (Taylor 1942(Taylor , 1960 and Brazil (Joly 1957, Ugadim 1975). According to Yang et al. (2015), C. teedei is found in the Atlantic and not in the western Pacific region, being absent in Korea and Japan (Fig. 4). ...
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Chondracanthus teedei is a small red alga, whose fronds are cartilaginous membranous, dark crimson to red black, turning yellow-green due to decay. The main frond axes, due to their broad branches, reach 1 cm in the oldest portions in the case of the lusitanicus variety. Originally described in Portugal, C. teedei is found widespread in the Atlantic, Mediterranean, Black Sea and Indian Ocean. This cosmopolitan species is typical of lower intertidal and shallow subtidal habitats, of sciophilous habitats in semi-exposed or protected areas, and tolerates the presence of mud and sand. Its composition may vary according to the geographical location of origin, and the time of year when it is collected. It has relatively high levels of protein (14.66%), ash (28.68%), fibres (2.21%), lipids (1.82%) and moisture (86.73%), making this alga able to be considered for its implementation as food. In addition, C. teedei var. lusitanicus produces hybrid carrageenans belonging to the lambda family (xi-theta hybrid carrageenans), in the case of tetrasporophytes, and kappa-iota-mu-nu hybrid carrageenans, in the case of female gametophytes and non-fruited thalli. The carrageenans extracted from this species have antifungal and antiviral activities, and the dry ground biomass of C. teedei has potential to be used in cosmetic formulations.
... Morphological plasticity, intraspecific variation in form due to environmental differences, makes identifying specimens difficult [33][34][35][36][37]. Similarly, different species with convergent morphologies are difficult to distinguish [38][39][40]. ...
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Cosmopolitan Acrosorium species with hook-forming thalli have been merged under the name of Acrosorium ciliolatum (Harvey) Kylin through a long and complicated nomenclatural history. We examined the specimens of ‘A. ciliolatum’ and related taxa from the northwestern (NW) Pacific, the UK, southern Spain, Australia, New Zealand, and Chile, using morphological and molecular analyses. We confirmed that these specimens are separated into four clades based on rbcL phylogeny, and the absence or presence of terminal hook-like structures represent intraspecific variation. Our results indicated that Acrosorium flabellatum Yamada, Cryptopleura hayamensis Yamada, Cryptopleura membranacea Yamada and the entities known as ‘A. ciliolatum’ in the NW Pacific are conspecific; the name A. flabellatum is the oldest and has priority. This taxon exhibits extreme variations in external blade morphology. We also confirmed that the position of the tetrasporangial sori is a valuable diagnostic characteristic for distinguishing A. flabellatum in the NW Pacific. We also discussed the need for further study of European and southern hemisphere specimens from type localities, as well as the ambiguous position of California specimens.
... Its distribution ranges from Paita, Perú (5°S) to Ancud, Chile (42°S) (Ramírez & Santelices, 1991). However, the last phylogenetic study over different population evidences that this species is not endemic to the South Pacific region, sharing mitochondrial COI gene and plastid rbcL with populations located in the coasts of Korea, Japan and France (Yeon et al., 2015). ...
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This work aims at the development of a bottom culture technology for the red algae Chondracanthus chamissoi, its main results are summarized in terms of production of biomass, length and critical variables for culture technologies development in this specie, such as the formation of secondary attachments discs and biofouling amount. Additionally, emphasis is placed on the main improvements that must be incorporated and the future challenges that the development of the aquaculture of C. chamissoi must face for the mass production of this commercial interest red seaweed.
... Because it is not possible to determine the exact origin of these species in the study area, they should be considered cryptogenic species (Carlton 1996). Determining the origin of the introduction as well as the colonization route for small species such as the ones found in this study is a challenging task, which in general requires an approach from molecular genetics (e.g., Holland 2000;Saltonstall 2002;Estoup and Guillemaud 2010;Rius et al. 2015;Yang et al. 2015;Geoffroy et al. 2016). ...
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We report new records of four macroalgae species in the Eastern Tropical Pacific, specifically from the rocky reefs of northern Chocó, Colombian Pacific. Among them, three species of Rhodophyta are included-Crouania mageshi-mensis Itono, 1977; Monosporus indicus Børgesen, 1931; Jania articulata N'Yeurt & Payri, 2009-and one species of Chlorophyta-Ulothrix subflaccida Wille, 1901. The new records increase the knowledge of tropical marine algae in the Pacific, open the discussion about possible dispersal mechanisms, and recall the importance of conducting molecular studies to define phylogenetic and biogeographic aspects of macroalgae. Citation: Rincón-Díaz N, Gavio B, Sánchez Muñoz JV, Chasqui L (2020) Crouania mageshimensis Itono, 1977 (Ceramiales, Rhodophyta) and new records for three other species of macroalgae from the Eastern Tropical Pacific. Check List 16 (5):
... Chondracanthus chamissoi (C. Agardh) Kützing is an alga in the order Gigartinales which is distributed on the coasts of Perú and Chile from 5 to 42°S, as well as in France, Korea, and Japan (Yang et al. 2015;Ramírez et al. 2018). It is exploited for extraction and commercialization of carrageenans and its biomass has also been intended for human consumption during the last 10 years (Bulboa et al. 2013), with prices that fluctuate between 23 and 32 USD$ per kg (dry weight) of raw material (Aduana 2018). ...
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In this study, we report for the first time the appearance of an endophytic filamentous red alga, living associated with the edible red seaweed Chondracanthus chamissoi. Molecular identification, growth, and reproduction of this endophyte are described under different conditions: photoperiod (8:16, 12:12, and 16:08 L:D), temperature (10 and 15 °C), and photon flux density (40 and 20 μmol photons m−2 s−1). Filaments of this endophyte were isolated and cultured, and the growth was recorded according to the number of ramifications and its reproduction, by quantifying the emergence of monosporangia. By sequencing the COI gene and through phylogeny reconstruction using maximum likelihood, we determined that this endophyte corresponded to Colaconema daviesii. The highest growth was recorded under the treatment 16:08 (L:D), 10 °C, and 20 μmol photons m−2 s−1, reaching up to 100 branches after 18 days. On the other hand, over 80% of branches with monosporangia were observed at 12:12 (L:D), 10 °C, and 20 μmol photons m−2 s-1. This is the first record of C. daviesii on the South-eastern Pacific as an endophyte on thalli of C. chamissoi. The results of the in vitro cultures showed that once C. daviesii is isolated, the filaments are able to grow and reproduce independently of C. chamissoi. This suggests that there may not be a strict relationship between both algae and reveals the possibility of finding C. daviesii living without a host or associated with other species
... Several phylogeographic studies on seaweed species have suggested their low capacity for dispersal (Leliaert et al 2018;Yang and Kim 2018); however, some seaweeds species seem to have undergone long-distance dispersal, such as Agarum clathratum Dumortier (Boo et al. 2011), Mastocarpus latissimus (Harvey) S.C. Lindstrom, Hughey & Martone , and Chondracanthus chamissoi (C. Agardh) Kützing (Yang et al. 2015). In addition, our results also showed genetically differentiated subgroups in the NW Pacific (A1-A2-A3), with levels of variation on par with that between the NW and NE Pacific subgroups (A3-A4) (Fig. 3). ...
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Genetic diversity patterns around the North Pacific received attention for marine organisms and have been used to infer biodiversity “hotspots” in the region. We conducted a phylogeographic study of the red alga Gloiopeltis furcata, investigating cryptic species diversity and comparing population genetic structure in the north Pacific. A phylogenetic tree and haplotype networks were constructed on the basis of 201 mitochondrial COI-5P sequences and 149 plastid rbcL sequences from G. furcata specimens. Eight distinct cryptic lineages (A–H) were identified within G. furcata. These lineages showed high genetic diversity and complex geographic distributions. All eight lineages of G. furcatasensulato were present in the NW Pacific; however, only a single lineage (A) was present in the NE Pacific, suggesting that the NW Pacific is a center of genetic diversity for G. furcatasensulato. Habitat discontinuities of G. furcatasensulato in the high rocky intertidal zone may have been responsible for the high level of genetic differentiation of G. furcatasensulato in the NW Pacific by impeding genetic exchange between adjacent populations. Our phylogenetic diversity suggests that the NW Pacific, especially Jeju Island, was a genetic diversity hotspot involving species diversity of Gloiopeltis.
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An epiphytic bladed member of the Bangiales was found growing in Christchurch (South Island, New Zealand) Molecular sequence data and morphological comparisons revealed that the New Zealand specimens belong to the species Pyropia koreana (M. S. Hwang & I. K. Lee) M. S. Hwang, H. G. Choi, Y. S. Oh & I. K. Lee. This is the first record of Py. koreana in the southern hemisphere and a new record of an introduced species in New Zealand.
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Chondracanthus chamissoi is a carragenophyte and edible red seaweed which is harvested from its natural habitat, and which, in recent years, has been diminishing along the Chilean coast. It is in this location that a new technique for the vegetative propagation of C. chamissoi by secondary attachment disc (SAD) was evaluated for the first time. In order to achieve this, SAD development was seasonally analyzed on natural and artificial substrata submitted to different water exchange and nutrient enrichment conditions. Cystocarpic and vegetative thalli were able to reattach and develop SADs during winter and summer under all tested conditions. However, higher values were observed in vegetative thalli cultivated in the winter. The size of shoots formed from SADs was higher in summer than in winter, varying in length between 0.2 and 5.4 mm after 40 days of cultivation. A continuous seawater exchange was the appropriate condition for SAD development whereas nutrient enrichment was not necessary. High adaptability of these structures to outdoor conditions was observed, as expressed in high survival rates of SADs. This study shows the technical feasibility of culturing C. chamissoi by means of SADs, which could be an alternative to spore and macrofragmentation strategies.
The red algal family Kallymeniaceae was surveyed along the west coast of Canada using the DNA barcode (COI-5P - 59 region of the mitochondrial cytochrome c oxidase I gene) as a species identification tool. A total of 253 specimens field identified as Pugetia spp. subsequently resolved as five genetic species groups, although only two are reported in the flora. Additionally, COI-5P data were available for the Chilean P. chilensis, which resolved as a distinct species. Subsequent analysis of the internal transcribed spacer of the ribosomal cistron and the universal plastid amplicon (domain V of the 23S rRNA gene) resolved the same groups as COI-5P. Phylogenetic relationships of the Canadian groups were investigated using large-subunit ribosomal DNA (LSU) and a combined analysis of LSU and COI-5P data. One species was divergent from the Pugetia spp. in all analyses and grouped closely with the kallymeniacean genera Erythrophyllum (Kallymeniaceae, Gigartinales) and Kallymeniopsis (Kallymeniaceae, Gigartinales) 2 it is here described as Beringia wynnei sp. nov. The other 'Pugetia' species fell into two divergent clusters in phylogenetic analyses, differing also in blade thickness, carpogonial branch morphology and the association of the auxiliary cell relative to the carpogonium (procarpic vs nonprocarpic). We retain the genus Pugetia for the type species P. fragilissima and P. cryptica sp. nov. and describe the new genus Salishia for Salishia firma (Kylin) comb. nov., S. sanguinea (Montagne) comb. nov. and S. chilensis (J. Agardh) comb. nov. Finally, we completed morphological and anatomical examinations of other Pugetia species to provide a comprehensive review of the genus.