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ISSN : 1226-9999(Print)
ISSN : 2287-7851(Online)
Korean J. Environ. Biol. Vol.44 No.1 pp.14-25
DOI : https://doi.org/10.11626/KJEB.2026.44.1.014

Boreolithothamnion astragaloi (Hapalidiales, Florideophyceae): A new record of coralline algae from Korea

Gwan Woung Kim, So Young Jeong1, Tae Oh Cho, Boo Yeon Won*
Department of Biological Science, Chosun University, Gwangju 61452, Republic of Korea
1Australian Rivers Institute-Coast & Estuaries and Coastal and Marine Research Centre, School of Environment and Science, Nathan Campus, Griffith University, Nathan, Queensland, Australia
*Corresponding author Boo Yeon Won Tel. 062-230-7983 giving_won@hanmail.net

Contribution to Environmental Biology


▪ We report Boreolithothamnion astragaloi as a new addition to the catalog of macroalgal flora in Korea.


▪ The known distribution of Boreolithothamnion astragaloi, formerly restricted to the northeastern Pacific (British Columbia, Canada), is herein extended to the northwestern Pacific.


13/06/2025 27/02/2026 10/03/2026

Abstract


Boreolithothamnion astragaloi, a coralline algal species belonging to the family Hapalidiaceae and order Hapalidiales, has been reported from British Columbia, Canada in northeastern Pacific. In this study, we report B. astragaloi for the first time from Korea, representing its initial record in the northwestern Pacific. Morpho-anatomical and molecular analyses of Lithothamnion-like specimens collected in Korea revealed that some individuals correspond to B. astragaloi. This species is characterized by flattened, flared epithallial cells; subepithallial initials as long as or longer than the cells immediately subtending them; cell fusions; multiporate tetrasporangial conceptacles; and tetra/bisporangial conceptacles developed from small groups of subepithallial initials. Phylogenetic analyses based on the COI-5P, psbA, and rbcL datasets place B. astragaloi firmly within the Boreolithothamnion clade, supporting its identification and extending the known distribution of the species to the northwestern Pacific.


Boreolithothamnion astragaloi , Hapalidiaceae , Hapalidiales

초록

    1. INTRODUCTION

    Coralline algae are some of the most distinctive and recognizable benthic marine plants, frequently dominating both intertidal and subtidal ecosystems (Woelkerling 1988). Despite their abundance, they are not easily identified and have been relatively understudied (Maneveldt et al. 2008). In Korea, these algae are widespread in intertidal and subtidal zones, with their documentation in the region’s marine flora mainly relying on morphological studies (Lee and Kang 2002; Lee 2008;Jeong et al. 2018). Recent advancements in molecular research have uncovered numerous new insights, deepening our understanding of coralline algae (Coutinho et al. 2021;Gabrielson et al. 2023;Kim et al. 2024;Jeong et al. 2025;Romero-Orozco et al. 2025;Taylor and Saunders 2025). This trend toward molecular methodologies underscores the need to combine traditional morphological techniques with modern approaches to fully grasp the diversity and ecological significance of coralline algae.

    The coralline alga Lithothamnion Philippi had been identified as a polyphyletic group through molecular studies (Peña et al. 2014;Richards et al. 2016). Gabriel- son et al. (2023) proposed a new genus, Boreolithothamnion P.W.Gabrielson, Maneveldt, Hughey & V.Peña, based on molecular and morphological data after Jeong et al. (2021) sequenced the type species of Lithothamnion, L. muelleri Lenormand ex Rosanoff. Molecular analyses have shown that Boreolithothamnion forms a well-supported monophyletic clade of cold-water, northern hemisphere species that are only distantly related to Lithothamnion. Although the two genera share several fundamental morphological features, such as a monomerous, non-coaxial thallus construction, flared epithallial cells, elongate intercalary meristematic cells, cell fusions, and multiporate tetrasporangial conceptacles, Gabrielson et al. (2023) showed that Lithothamnion species often possess squarish, flattened, and flared epithallial cells, with all three cell shapes sometimes present within a single specimen, whereas Boreolithothamnion has only flared epithallial cells. In addition, trichocytes are commonly to occasionally found in Lithothamnion but have not been reported in Boreolithothamnion. Boreolithothamnion now includes fourteen species: B. acuminatum S.Y.Jeong, B.Y.Won, Fredericq & T.O.Cho, B. astragaloi K.E.Taylor & G.W.Saunders, B. coralloides (P.Crouan & H.Crouan) V.Peña & P.W.Gabrielson, B. giganteum (L.R.Mason) P.W.Gabrielson, Maneveldt, Hughey & V.Peña, B. glaciale (Kjellman) P.W.Gabrielson, Maneveldt, Hughey & V.Peña, B. homoglobosum S.Y.Jeong, B.Y.Won, Fredericq & T.O.Cho, B. japonicum (Foslie) S.Y.Jeong, B.Y.Won, Fredericq & T.O.Cho, B. jejuensis S.Y.Jeong, B.Y.Won, Fredericq & T.O.Cho, B. lemoineae P.W.Gabrielson, Maneveldt, Hughey & V.Peña, B. phymatodeum (Foslie) P.W.Gabrielson, Maneveldt, Hughey & V.Peña, B. sonderi (Hauck) P.W. Gabrielson, Maneveldt, Hughey & V.Peña, B. soriferum (Kjellman) P.W.Gabrielson, Maneveldt, Hughey & V. Peña, B. tanuense K.E.Taylor & G.W.Saunders and B. tophiforme (Esper) P.W.Gabrielson, Maneveldt, Hughey & V.Peña (Guiry and Guiry 2026). Among these, B. acuminatum, B. coralloides (initially described as Lithothamnion coralloides), B. giganteum (initially described as Lithothamniongiganteum), B. homoglobosum, B. japonicum (initially described as Lithothamnion japonicum), B. jejuensis, B. lemoineae (initially described as Lithothamnion lemoineae), and B. phymatodeum (initially described as Lithothamnionpacificum) are known to occur in Korea (Lee and Kang 2002;Lee 2008;Jeong et al. 2018, 2025).

    Boreolithothamnionastragaloi was originally described from British Columbia, Canada (Taylor and Saunders 2025). Its type locality is the channel between Murchison Island and Faraday Island, Haida Gwaii, Canada, and the species has been regarded as endemic to the northeastern Pacific. However, there have been no reports of this species in the northwest Pacific.

    Three Lithothamnion-like specimens were collected from subtidal areas at depths 20-30 m along the eastern coast of Korea, including Ulleungdo. In this study, we examined their morpho-anatomical characteristics and generated DNA sequences of the mitochondrial COI-5P and chloroplast psbA and rbcL genes. Based on an integrative analysis of morphological and molecular data, all specimens were identified as Boreolithothamnionastragaloi, a species previously known only from British Columbia and newly recorded here for Korea.

    2. MATERIALS AND METHODS

    2.1. Morphological analyses

    Samples were collected from subtidal zones in Korea at depths of approximately 20 m to 30 m. The specimens were photographed to capture their external morphological features using a waterproof digital camera (Nikon COOLPIX AW100; Nikon Corp., Tokyo, Japan). The airdried samples were broken into pieces using a chisel and a small hammer. These fragments were then affixed to aluminum stubs with double-sided adhesive carbon tape (Nisshin EM Co., Ltd., Japan), and coated with gold or platinum for 10 min using digital ion coater (SPT-20; COXEM Co., Ltd., Korea). The samples were analyzed using a Hitachi S-4800 scanning electron microscope (FE-SEM; Hitachi, Ltd., Tokyo, Japan) or a COXEM EM-30 PLUS+ scanning electron microscope (Mini SEM; COXEM Co., Ltd., Korea) at an accelerating voltage of 15 or 20 kV. The resulting digital images were edited for clarity using Adobe Photoshop software v.6.1 (Adobe Systems Inc., San Jose, CA, USA). Representative voucher specimens from this study were deposited in the herbarium of Chosun University (CUK) and the Marine Biodiversity Institute of Korea (MABIK).

    2.2. Molecular analyses

    DNA was extracted from 3 silica gel-dried specimens using the NucleoSpin Plant II Kit (Macherey-Nagel, Düren, Germany). The COI-5P, rbcL and psbA genes were amplified via polymerase chain reaction (PCR) using HelixAmp Ready-2x-Go premix from NanoHelix Co., Ltd. (Daejeon, Korea). GazF1/GazR1 for COI-5P (Saunders 2005), two primer sets for psbA (psbAF1/ psbAR2 or psbAF1/psbA600R; Yoon et al. 2002) and four primer sets for rbcL (F57/RrbcStart, F57/R753, F577/ R1150, and F993/RrbcStart; Freshwater and Rueness 1994) were used for amplification. PCR was carried out on a Veriti 96-well Thermal Cycler (Applied Biosystems, Waltham, MA, USA). Two COI-5P, three psbA, and two rbcL sequences from the coralline algae obtained in this study have been submitted to GenBank. The newly generated COI-5P, rbcL, and psbA sequences, together with those retrieved from GenBank, were used for phylogenetic analysis. All sequences were aligned using Geneious Prime® (v.2023.0.1; Biomatters Ltd., Auckland, New Zealand). Genetic distances were calculated using the p-distance method in Mega X (Kumar et al. 2018). Phylogenetic trees were generated for individual dataset. The optimal partitioning scheme and substitution models were selected using PartitionFinder 2 (Lanfear et al. 2017) under the Akaike Information Criterion (AIC). Maximum-likelihood (ML) analysis were conducted under the GTR+I+G model with 1,000 bootstrap replicates using RAxMLGUI v1.5 (Silvestro and Michalak 2012) for all datasets. Bayesian inference was carried out with MrBayes 3.2.6 (Ronquist and Huelsenbeck 2003) involving Markov chain Monte Carlo runs performed for 2,000,000 generations with one cold and three heated chains, utilizing the GTR+G+I evolutionary model. Trees were sampled every 1,000 generations, and summary trees were created with a burn-in value of 25%.

    3. RESULTS AND DISCUSSION

    Division Rhodophyta Wettstein, 1901 홍조식물문

    Class Florideophyceae Cronquist, 1960 진정홍조강

    Order Hapalidiales W.A. Nelson, J.E. Sutherland, T.J. Farr & H.S. Yoon, 2015 방석산호말목 Family Hapalidiaceae J.E. Gray, 1864 방석산호말과 Genus Boreolithothamnion P.W.Gabrielson, Maneveldt, Hughey & V.Peña 2023 북방쩍속 (국명신칭)

    Boreolithothamnionastragaloi K.E.Taylor & G.W.Saunders 주사위북방쩍 (신칭) (Fig. 1A-K)

    Holotype. UNB GWS048600, coll. G.W.Saunders & C. Brooks, 2 August 2022, subtidal (5 m).

    Type locality. Channel between Murchison Island and Faraday Island, Haida Gwaii, BC, Canada, 52.59687N, 131.47513W.

    Specimens examined. CUK14763 (MABIK AL00103 529) (Yeolgi-ri, Dongsan-ri, Hyeonnam-myeon, Yangyang- gun, Gangwon-do, Korea, subtidal at 20 m, collected by S.Y.Jeong & T.O.Cho, 31 July 2014), CUK19076, CUK19088 (Taeha 1-gil, Seo-myeon, Ulleung-gun, Gyeongsangbuk-do, Korea, subtidal 30 m, collected by T.O.Cho & S.Y.Jeong & Jose & Antony & G.C.Choi, 29 August 2018).

    Geographic distribution. Skidegate Inlet and northeast Gwaii Hanaas, Haida Gwaii, BC, Canada (Taylor and Saunders 2025) and the east coast of Korea including Ulleungdo (present study).

    Vegetative structure. Plants are non-geniculate encrusting to warty, with short and terete individual protuberances. Protuberances are less than 0.3 cm high, unbranched, acuminate or cylindrical in shape, up to 2.0- 2.3 mm long and 1.1-1.3 mm wide, with rounded apex (Fig. 1A, B). Thallus margins are composed of plumose medullary filaments. Thalli are monomerous, dorsiventrally organized in the crustose portion (Fig. 1C), and radially organized in protuberances. Haustoria are absent. Thalli are 280-432 μm thick and composed of hypothallium and perithallium (Fig. 1C). Epithallial cells arranged in a single layer flattened and flared, 0.7- 2.7 μm long, and 3.3-7.1 μm wide (Fig. 1D). Subepithallial initials are rectangular with rounded corners, as long as or longer than the cells immediately subtending them, 3.1-7.7 μm long, and 4.3-8.7 μm wide (Fig. 1D). Perithallium comprises the bulk of thallus thickness, 121-665 μm, and usually constitutes over 47-65% of thallus thickness, except near the margins. Perithallial cells are ovoid to rectangular with rounded corners, 4.4-9.4 μm long and 3.8-9.0 μm wide (Fig. 1E). Hypothallium is plumose (non-coaxial), comprising about 11-20 cell layers, 101-168 μm thick, and usually less than 34-52% of the mature thallus thickness. Hypothallial cells are rectangular to elongated with rounded corners, 8.6-23.1 μm long, and 3.0-8.3 μm wide (Fig. 1F, G). Downward curving medullary filaments terminate in more-or less triangular-shaped cells (Fig. 1C). Cell fusions are abundant among cortical and medullary cells, frequently occupying most of the walls of adjoining cells (Fig. 1D, E, G). Secondary pit connections and trichocytes were not observed.

    Reproductive structure. Tetrasporangial conceptacles are multiporate, slightly raised to low-domed, 47-74 μm, lacking rims, not fused with adjacent conceptacles, and occur above the surrounding thallus surface (Fig. 1H), 350 μm in external diameter. Pore plates are rounded and approximately 145 μm in diameter, with up to 8 pores (Fig. 1H). Pores are irregularly distributed on the pore plate, surrounded by a rosette of 6-7 cells, and neither protruded nor sunken in relation to the cells of the surrounding roof surface (Fig. 1I). Tetrasporangial conceptacle chambers are rounded to elliptical, 246 μm in diameter, and 126 μm in height (Fig. 1J). Roofs of tetrasporangial conceptacles are 25 μm in thickness and composed of pore canals, pore canal lining filaments, and roof filaments (Fig. 1K). Gametangial thalli were not observed in this study.

    Habitat and phenology. The plants collected in Korea attached to shell (epizoic) and rock (epilitic) from subtidal at 20-30 m depth. Tetrasporangial plants collected in July.

    Phylogenetic analyses. A total of 663 base pair (bp) sequences of COI-5P, 1,351 bp of rbcL, and 845 bp of the psbA sequences were obtained from coralline algal specimens collected in Korea. The newly generated COI-5P, rbcL, and psbA sequences, together with sequences downloaded from GenBank, were aligned. Phylogenetic trees were inferred separately using COI- 5P, rbcL, and psbA datasets (Figs. 2-4). Sporolithon spp. was included as outgroup taxa. Phylogenetic analyses based on each dataset consistently revealed that the specimens from Korea are nested within the Boreolithothamnion clade (Figs. 2-4). Additionally, the newly generated rbcL sequences clustered with the holotype sequence of Boreolithothamnionastragaloi (GenBank accession no. PQ882800) with high support (bootstrap value=97; Bayesian posterior probability=1.0), as reported by Taylor and Saunders (2025). This clustering pattern, recovered in COI-5P and psbA datasets, further confirms the identity of the studied specimens as B. astragaloi.

    Remarks. The genus Boreolithothamnion was established by Gabrielson et al. (2023), and within this genus, Boreolithothamnionastragaloi was recently described by Taylor and Saunders (2025) from British Columbia, Canada. The Korean specimens examined in the present study conform well to the diagnostic characters of B. astragaloi, including: 1) flattened, flared epithallial cells (Fig. 1D); 2) subepithallial initials that are as long as or longer than the cells immediately subtending them (Fig. 1D); 3) the presence of cell fusions (Fig. 1E, G); the absence of trichocytes (Fig. 1D, E); and 4) multiporate tetrasporangial conceptacles (Fig. 1H). At present, no stable interspecific morphological characters can be identified among Boreolithothamnion species, likely due to high morphological plasticity and the absence of female and male reproductive structures (Taylor and Saunders 2025). A comparison of morphoanatomical characters among Boreolithothamnion species reported from Korea is summarized in Table 1. Consequently, molecular phylogenetic data provide a more robust framework for species delimitation within the genus.

    Phylogenetic analyses based on the COI-5P, rbcL, and psbA gene datasets further support the identification of the Korean specimens as Boreolithothamnionastragaloi (Figs. 2-4). In the phylogenetic tree, B. astragaloi was consistently resolved as the sister taxon to B. soriferum across all gene datasets. The genetic divergence between the B. astragaloi and its sister taxon, B. soriferum was 4.5-4.8% for COI-5P, 0.8-2.2% for rbcL, and 0.5-0.7% for psbA respectively and these values fall within the range of genetic distances typically observed among species of coralline algae (Taylor and Saunders 2025).

    Boreolithothamnionastragaloi has previously been reported only from North America, specifically British Columbia (Taylor and Saunders 2025). According to Taylor and Saunders (2025), this species occurs in subtidal zones at depths of 5-22 m, where it grows either as epizoic crusts on shells or as free-living rhodoliths. In the present study, B. astragaloi was collected from deeper subtidal habitats (20-30 m depth) off the coast of Yangyang and Ulleungdo, representing the first confirmed record of this species from Korea. This new record from the East Sea of Korea substantially extends the known geographic distribution of B. astragaloi into the northwestern Pacific region (Fig. 5).

    ACKNOWLEDGEMENTS

    This study was supported by the Ministry of Ocean and Fisheries (Marine Biotics Project, 20210469), and by the management of Marine and Fishery Bio-resources Center (2026) funded by the National Marine Biodiversity Institute of Korea (MABIK).

    CRediT authorship contribution statement

    GW Kim: Formal analysis, Writing-Original draft. SY Jeong: Formal analysis, Writing-Original draft. TO Cho: Funding acquisition, Supervision. BY Won: Writing-Original draft, Visualization.

    Declaration of Competing Interest

    The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

    Figure

    KJEB-44-1-14_F1.jpg

    Vegetative and reproductive structures of Boreolithothamnionastragaloi. (A) Specimen forming protuberances growing on a shell. (B) Specimen growing directly on rock without protuberances. (C) Vertical section of thallus showing hypothallial filaments (white square brackets) and peripheral cells (black square brackets) with monomerous construction. (D) Vertical section of thallus surface showing flared epithallial cells (arrowhead), subepithallial initials (asterisk), and cell fusions (f). (E) Vertical section of peripheral cell layer with cell fusions (f). (F) Hypothallial layer curved outward. (G) Hypothallial cells with cell fusions (f). (H) Surface view of multiporate tetrasporangial conceptacles. ( I ) Pore of tetrasporangial conceptacle surrounded by rosette cells. (J) Vertical section of tetrasporangial conceptacle. (K) Vertical section of pore canal showing rosette cells aligned with the thallus surface.

    KJEB-44-1-14_F2.jpg

    Phylogenetic tree based on Maximum likelihood (ML) and Bayesian analysis with COI-5P dataset. Value above branches=Maximum likelihood bootstrap values (BS) in % ≥50/Bayesian posterior probabilities (BPP) ≥0.75. Values lower than BS 50 or BPP 0.75 are indicated by hyphens (-). Values of BS 100 or BPP 1.00 are indicated by asterisks (*). Sequences generated in this study are highlighted in bold.

    KJEB-44-1-14_F3.jpg

    Phylogenetic tree based on Maximum likelihood (ML) and Bayesian analysis with psbA dataset. Value above branches=Maximum likelihood bootstrap values (BS) in % ≥50/Bayesian posterior probabilities (BPP) ≥0.75. Values lower than BS 50 or BPP 0.75 are indicated by hyphens (-). Values of BS 100 or BPP 1.00 are indicated by asterisks (*). Sequences generated in this study are highlighted in bold.

    KJEB-44-1-14_F4.jpg

    Phylogenetic tree based on Maximum likelihood (ML) and Bayesian analysis with rbcL dataset. Value above branches=Maximum likelihood bootstrap values (BS) in % ≥50/Bayesian posterior probabilities (BPP) ≥0.75. Values lower than BS 50 or BPP 0.75 are indicated by hyphens (-). Values of BS 100 or BPP 1.00 are indicated by asterisks (*). Sequences generated in this study are highlighted in bold.

    KJEB-44-1-14_F5.jpg

    Distributions of Boreolithothamnionastragaloi based on newly generated sequences and sequences retrieved from GenBank.

    Table

    Comparison of morpho-anatomical features among Boreolithothamnion species in Korea

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    Vol. 40 No. 4 (2022.12)

    Journal Abbreviation 'Korean J. Environ. Biol.'
    Frequency quarterly
    Doi Prefix 10.11626/KJEB.
    Year of Launching 1983
    Publisher Korean Society of Environmental Biology
    Indexed/Tracked/Covered By

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