Algal Flora of Korea

Volume 4, Number 2 Rhodophyta: Florideophyceae: Nemaliophycidae: Acrochaetiales, Colaconematales, Palmariales, Nemaliales Nemalian Red Algae

Flora and Fauna of Korea

National Institute of Biological Resources Ministry of Environment National Institute of Biological Resources Ministry of Environment Russia

CB Chungcheongbuk-do CN Chungcheongnam-do HB GB Gyeongsangbuk-do China GG Gyeonggi-do YG GN Gyeongsangnam-do GW Gangwon-do HB Hamgyeongbuk-do JG HN Hamgyeongnam-do HWB Hwanghaebuk-do HN HWN Hwanghaenam-do PB JB Jeollabuk-do JG Jagang-do JJ Jeju-do JN Jeollanam-do PN PB Pyeonganbuk-do PN Pyeongannam-do YG Yanggang-do HWB HWN GW East Sea GG GB (Ulleung-do) Yellow Sea CB CN GB

JB GN JN

JJ South Sea Algal Flora of Korea

Volume 4, Number 2 Rhodophyta: Florideophyceae: Nemaliophycidae: Acrochaetiales, Colaconematales, Palmariales, Nemaliales Nemalian Red Algae

2011

National Institute of Biological Resources Ministry of Environment

Algal Flora of Korea

Volume 4, Number 2 Rhodophyta: Florideophyceae: Nemaliophycidae: Acrochaetiales, Colaconematales, Palmariales, Nemaliales Nemalian Red Algae

Published by the National Institute of Biological Resources Environmental Research Complex, Gyeongseo-dong, Seo-gu Incheon 404-708, Republic of Korea www.nibr.go.kr

All rights reserved. No part of this book may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise, without the prior permission of the National Institute of Biological Resources.

ISBN : 9788994555621-96470 Government Publications Registration Number 11-1480592-000163-01

Printed by Junghaengsa, Inc. in Korea on acid-free paper

Publisher : Chong-chun Kim Project Staff : Hong-Yul Seo, Ye Eun, Joo-Lae Cho

Published on February 28, 2011

The Flora and Fauna of Korea logo was designed to represent six major target groups of the project including vertebrates, invertebrates, insects, algae, fungi, and bacteria. The book cover and the logo were designed by Jee-Yeon Koo. Preface

Biological resources are important elements encompassing organisms, genetic resources, and parts of organisms which provide potential values essential for human lives. The creation of high-valued products such as new varieties of organisms, new substances, and the development of new drugs by harnessing biological resources is now widely perceived to be one of the major indices of national competitiveness. In the wake of the “Convention of Biological Diversity”, which was adopted in 1992 in recognition of national sovereignty over indigenous biological and genetic resources, all the countries of the world are now concerting their efforts on the discovery of original materials for the bio-industry, initiating international competition in the 21st century. Competition among countries for biological resources is now entering an intense phase following the adoption of the ABS (Access to genetic resources and Benefit-Sharing) international regime in Nagoya in 2010. For this reason, the National Institute of Biological Resources of the Korean Ministry of Environment recognizes the preservation and management of the biological resources in Korea for the bio-industry as a first priority project for the future, and has begun publication of Flora and Fauna of Korea for the systematic preservation and efficient management of our biological resources. Korea has been acclaimed as a country with a high level of biological diversity, the total number of described species in Korea to date being about 37,000. Beginning in 2006, the National Institute of Biological Resources embarked on the publication of Flora and Fauna of Korea which, containing comprehensive and diverse information on our invaluable native species, has become the standard textbook of native species. The systematic survey of diverse taxa in all parts of Korea led by a group of professionals in the field of taxonomy over the past four years has finally come to fruition and culminated in the appearance of 16 monographs in the 2010 volumes of Flora and Fauna of Korea encompassing 1,037 species in 158 families belonging to 9 phyla, along with further volumes of Flora and Fauna of Korea encompassing 1,163 species in 112 families belonging to 7 phyla due to appear this year. This is the first volume of Flora and Fauna of Korea in which a taxon of organisms of the Korean Peninsula is extensively treated at the level of species. Flora and Fauna of Korea will contribute to raising the standard of Korean taxonomy and improve pride in the management of our biological resources through enhanced understanding of the true nature of our native species. In addition, I am confident that the ongoing publication of Flora and Fauna of Korea will significantly contribute to paving the way for sustainable, wise use of biological resources. I would like to express my sincerest gratitude to Dr. Il-Ki Hwang, National Fisheries Research and Development Institute and Professor Hyung-Seop Kim, Gangneung-Wonju National University, who are responsible for writing this publication of Flora and Fauna of Korea. This series will play a pivotal role in the census of native Korean species, which are estimated to number 100,000. By promoting innovative and taxonomic research for the identification of the totality of native Korean species and by continuously publishing such results in Flora and Fauna of Korea, I sincerely hope that a valuable foundation will be laid for the sustainable use of our national biological resources through the exten- sive research, development and for their profitable use by a prosperous bio-industry in the creation of high-valued products such as natural products, medicines, cosmetics and essential supplements in our country.

Chong-chun Kim, Ph. D. President NIBR 1

Contents

List of Taxa 2 Introduction 4 Materials and Methods 7 Taxonomic Notes 9 1. Acrochaetium canariense Børgesen 12 2. Acrochaetium catenulatum Howe 13 3. Acrochaetium densum (Drew) Papenfuss 15 4. Acrochaetium inkyui Lee 17 5. Acrochaetium microscopicum (Nägeli ex Kützing) Nägeli 19 6. Acrochaetium sancti-thomae Børgesen 22 7. Acrochaetium scapae (Lyle) Papenfuss 24 8. Acrochaetium secundatum (Lyngbye) Nägeli in Nägeli et Cramer 25 9. Acrochaetium terminale (Nakamura) Lee 28 10. Acrochaetium virgatulum (Harvey) Batters 30 11. Rhodochorton purpureum (Lightfoot) Rosenvinge 33 12. Colaconema bispora (Børgesen) Hwang et Kim, comb. nov. 37 13. Colaconema codicola (Børgesen) Stegenga, Bolton et Anderson 39 14. Colaconema codii (Crouan et Crouan) Hwang et Kim, comb. nov. 42 15. Colaconema compta (Børgesen) Hwang et Kim, comb. nov. 44 16. Colaconema daviesii (Dillwyn) Stegenga 47 17. Colaconema dictyotae (Collins) Hwang et Kim, comb. nov. 51 18. Colaconema elegans (Drew) Hwang et Kim, comb. nov. 54 19. Colaconema hyalosiphoniae (Nakamura) Hwang et Kim, comb. nov. 56 20. Colaconema infestans (Howe et Hoyt) Woelkerling 58 21. Colaconema pacificum (Kylin) Woelkerling 61 22. Colaconema thuretii (Bornet) Gabrielson in Gabrielson, Widdowson, Lindstrom, Hawkes et Scagel 64 23. Rhodonematella subimmersa (Setchell et Gardner) Clayden et Saunders 69 24. Actinotrichia fragilis (Forsskål) Børgesen 74 25. Dichotomaria apiculata (Kjellman) Kurihara et Masuda 76 26. Dichotomaria falcata (Kjellman) Kurihara et Masuda 79 27. Galaxaura rugosa (J. Ellis et Solander) J.V. Lamouroux 83 28. Tricleocarpa cylindrica (J. Ellis et Solander) Husiman et Borowitzka 84 29. Helminthocladia australis Harvey 88 30. Nemalion vermiculare Suringar 91 Literature Cited 94 Index to Korean Names 105 Index to Korean Names as Pronounced 106 Index to Scientific Names 108 2

List of Taxa

Subclass Nemaliophycidae Christensen 1978 Order Acrochaetiales Feldmann emend. Harper et Saunders 2002 Family Acrochaetiaceae Fritsch emend. Harper et Saunders 2002 Genus Acrochaetium Nägeli enmed. Harper et Saunders 2002 Acrochaetium canariense Børgesen 1927 Acrochaetium catenulatum Howe 1914 Acrochaetium densum (Drew) Papenfuss 1945 Acrochaetium inkyui Lee 1987 Acrochaetium microscopicum (Nägeli ex Kützing) Nägeli 1861 Acrochaetium sancti-thomae Børgesen 1915 Acrochaetium scapae (Lyle) Papenfuss 1945 Acrochaetium secundatum (Lyngbye) Nägeli in Nägeli et Cramer 1858 Acrochaetium terminale (Nakamura) Lee 1987 Acrochaetium virgatulum (Harvey) Batters 1902 Genus Rhodochorton Nägeli emend. Harper et Saunders 2002 Rhodochorton purpureum (Lightfoot) Rosenvinge 1900 Order Colaconematales Harper et Saunders 2002 Family Colaconemataceae Harper et Saunders 2002 Genus Colaconema Batters enmend. Harper et Saunders 2002 Colaconema bispora (Børgesen) Hwang et Kim, comb. nov. Colaconema codicola (Børgesen) Stegenga, Bolton et Anderson 1997 Colaconema codii (Crouan et Crouan) Hwang et Kim, comb. nov. Colaconema compta (Børgesen) Hwang et Kim, comb. nov. Colaconema daviesii (Dillwyn) Stegenga 1985 Colaconema dictyotae (Collins) Hwang et Kim, comb. nov. Colaconema elegans (Drew) Hwang et Kim, comb. nov. Colaconema hyalosiphoniae (Nakamura) Hwang et Kim, comb. nov. Colaconema infestans (Howe et Hoyt) Woelkerling 1973 Colaconema pacificum (Kylin) Woelkerling 1971 Colaconema thuretii (Bornet) Gabrielson in Gabrielson, Widdowson, Lindstrom, Hawkes et Scagel 2000 Order Palmariales Guiry et Irvine in Guiry 1978 Famliy Rhodophysemataceae Saunders et McLachlan 1989 Genus Rhodonematella Clayden et Saunders 2010 Rhodonematella subimmersa (Setchell et Gardner) Clayden et Saunders 2010 Order Nemaliales Schmitz in Engler 1892 Family Galaxauraceae Parkinson 1983 Genus Actinotrichia Decaisne 1842 Actinotrichia fragilis (Forsskål) Børgesen 1932 Genus Dichotomaria Lamarck 1816 gen. emend. Wang et al. 2005 Dichotomaria apiculata (Kjellman) Kurihara et Masuda 2005 List of Taxa 3

Dichotomaria falcata (Kjellman) Kurihara et Masuda 2005 Genus Galaxaura Lamouroux 1812 Galaxaura rugosa (J. Ellis et Solander) J.V. Lamouroux 1816 Genus Tricleocarpa Huisman et Browitzka 1990 Tricleocarpa cylindrica (J. Ellis et Solander) Husiman et Borowitzka 1990 Family Liagoraceae Kützing 1843 Genus Helminthocladia J. Agardh 1852 Helminthocladia australis Harvey 1863 Genus Nemalion Duby 1830 Nemalion vermiculare Suringar 1874 4

Introduction

Recent molecular systematic works have resolved five higher-level lineages within the red algal class Florideophyceae. The five lineages are ranked as sub-classes, Hindenbrandiophycidae, Nemaliophycidae, Corallinophycidae, Ahnfeltiophycidae, and Rhodymeniophycidae (Le Gall and Saunders 2007; Verbruggen et al. 2010). The subclass Nemaliophycidae Christensen (1978: 66) comprises 9 orders (Acrochaetiales, Balbianiales, Balliales, Batrachospermales, Colaconematales, Nemaliales, Palmariales, Rhodachlyales, Thoreales) that includes those with cells possessing two cap layers on the cytoplasmic faces of the pit plugs (Saunders and Hommersand 2004; Le Gall and Saunders 2007). Systematics within the subclass Nemaliophycidae has improved profoundly with the aid of molecular tools. Ragan et al. (1994) provided the first molecular support for a controversial alliance between the Acrochaetiales, Nemaliales and Palmariales base on SSU-rDNA sequence data. Saun- ders et al. (1995) added the Batrachospermales to this lineage and transferred an acrochaetioid genus Rhodothamniella to the Palmariales. Subsequently, Saunders and Bailey (1997) provided supporting evidence for an alliance of this lineage to the Corallinales and Rhodogorgonales. The acrochaetioid genera Balbiania and Rhododraparnaldia were assigned to a new order Balbianiales (Sheath and Müller 1999) and added to this lineage. Furthermore, the marine filamentous Ballia

Bangiophyceae Hildenbandiophycidae Palmariales Acrochaetiales Colaconematales Nemaliales Balliales Balbianiales Rhodachlyales Nemaliophycidae Thoreales

Batrachospermales Florideophyceae

Corallinophycidae Ahnfeltiophycidae

: domed outer cap Rhodymeniophycidae : thin outer cap with cap membrane : inner cap lost

Fig. 1. Summarized phylogenetic tree of Rhodophyta base on multi-gene DNA sequence data of subclass Nemaliophycidae (references papers: Le Gall and Saunders 2007; Verbruggen et al. 2010). Explain the dotted line enclosing the Nemaliophycidae and bold type order names represent marine members. Introduction 5

Table 1. Updated classification systems of Nemaliophycidae.

Acrochaetiales Feldman 1953 Fam. 1. Acrochaetiaceae Fritsch ex W.R. Taylor 1957 [Acrochaetium Nägeli in Nägeli et Cramer 1858, Audouinella Bory de Saint-Vincent 1823, Grania (Rosenvinge) Kylin 1944, Rhodochorton Nägeli 1862]

Balbianiales Sheath and Müller 1999 Fam. 2. Balbianiaceae Sheath et Müller 1999 [Balbiania Sirdot 1876, Rhododraparnaldia Sheath, Whittick et Cole 1994]

Balliales Choi, Kraft et Saunders 2000 Fam. 3. Balliaceae Choi, Kraft and Saunders 2000 [Ballia Harvey 1840]

Batrachospermales Pueschel et Cole 1982 Fam. 4. Batrachospermaceae Fries 1825 [Balliopsis Saunders et Necchi 2002, Batrachospermum Roth 1797, Kumanoa Entwisle, Vis, Chiasson, Necchi et Sherwood 2009, Nothocladus Skuja 1934, Petrohua Saunders in Vis, Harper and Saunders 2007, Sirodotia Kylin 1912, Tuomeya Harvey 1858] Fam. 5. Lemaneaceae Roemer 1845 [Lemanea Bory de Saint-Vincent 1808, Paralemanea (Silva) Vis and Sheath 1992, Sacheria Sirodot 1872] Fam. 6. Psilosiphonaceae Entwisle, Sheath, Roemer, Müller et Vis 1996 [Psilosiphon Entwisle 1989]

Colaconematales Harper et Saunders 2002 Fam. 7. Colaconemataceae Harper et Saunders 2002 [Colaconema Batters 1896]

Nemaliales Schmitz 1892 Fam. 8. Galaxauraceae Parkinson 1983 [Actinotrichia Decaisne 1842, Dichotomaria Lamarck 1816, Galaxaura Lamou- roux 1812, Tricleocarpa Huisman et Borowitzka 1990] Fam. 9. Liagoraceae Kützing 1843 [Akalaphycus Huisman, Abbott et Sherwood 2004, Cumagloia Setchell et Gardner 1917, Cylindraxis Kraft et Huisman 1989, Dermonema Harvey ex Heydrich 1894, Dotyophycus Abbott 1976, Ganonema Fan et Wang 1974, Gloiotrichus Huisman et Kraft 1994, Helminthocladia J. Agardh 1852, Helminthora J. Agardh 1852, Izziella Doty 1978, Liagora Lamouroux 1812, Liagoropsis Yamada 1944, Nemalion Duby 1830, Patenocarpus Yoshizaki 1987, Sinocladia Tseng et Li in Zeng et al. 2000, Stenopeltis Itono et Yoshizaki 1992, Trichogloea Kützing 1847, Trichogloeopsis Abbott et Doty 1960, Yamadaella Abbott 1970] Fam. 10. Scinaiaceae Huisman, Harper et Saunders 2004 [Gloiophloea J. Agardh 1872, Nothogenia Montagne 1843, Scinaia Bivona-Bernardi 1822, Whidbeyella Setchell et Gardner 1903]

Palmariales Guiry and Irvine 1978 Fam. 11. Meiodiscaceae Clayden et Saunders 2010 [Meiodiscus Saunders et McLachlan 1991, Rubrointusa Clayden et Saunders 2010] Fam. 12. Palmariaceae Guiry 1974 [Devaleraea Guiry 1982, Halosaccion Kützing 1843, Neohalosacciocolax Lee et Kurogi 1978, Palmaria Stackhouse 1801] Fam. 13. Rhodophysemataceae Saunders et McLachlan 1990 [Coriophyllum Setchell et Gardner 1917, Pseudorhodo- discus Masuda 1976, Rhodonematella Clayden et Saunders 2010, Rhodophysema Batters 1900, Rhodophysemopsis Masuda 1976] Fam. 14. Rhodothamniellaceae Saunders 1995 [Camontagnea Pujals 1981, Rhodothamniella Feldmann in Christensen 1978]

Rhodachlyales Saunders, Clayden, Scott, West, Karsten et West in West et al. 2008 Fam. 15. Rhodachlyaceae Saunders, Clayden, Scott, West, Karsten et West in West et al. 2008 [Rhodachlya West, Scott, West, Karsten, Clayden et Saunders in West et al. 2008] 6 Algal Flora of Korea·Nemalian Red Algae

(Ceramiaceae), which was separated to the order Balliales by Choi et al. (2000) and added a member of this lineage. The distinctness of genus Thorea from Batrachosperales was resolved (Vis et al. 1998), and later was established Thoreales (Müller et al. 2002). A detail molecular survey of the Acrochaetiales resolved two divergent lineages, resulting in recognition of an additional order Colaconematales (Harper and Saunders 2002). Recently, West et al. (2008) established a new order Rhodachlyales base on a new marine acrochaetioid genus Rhodachlya. On the other hand, Le Gall and Saunders (2007) removed the Corallinales and Rhodogorgonales from the Nemaliophycidae and placed them in a segregate subclass, Corallinophycidae. Furthermore, two unusual Colaconema species, C. membranaceum and C. subimmersum, were reinterpretated base on life history and molecular phylogeny and removed to Palmariales by establishing two new genera Rubrointrusa and Rho- donematella for the two species (Clayden and Saunders 2010). The members of orders Balbianiales, Batrachospermales and Thoreales occur exclusively in freshwater. The marine components of this lineage comprise various morphological types, from simple or branched monosiphonous (Acrochaetiales, Colaconematales, Rhodachlyales), multiaxial pseudoparenchymatous (Nemaliales) and true parenchymatous (Palmariaceae, Palmariales), were distinguished by the synapomorphic character state of “thin outer cap with cap membranes” pit plugs (Pueschel 1987; Le Gall and Saunders 2007). The phylogenetic tree of Nemaliophycidae based on recent molecular works (Le Gall and Saunders 2007; Verbruggen et al. 2010) was summerarized in Fig. 1, and is usued in the updated classification system in Table 1. 7

Materials and Methods

Korean members of Namaliophycidae have been collected mainly by the authors during the past 5 years. Specimens investigated were preserved in 10% formaldehyde-sea water, with voucher specimens deposited in the Herbarium of the National Institute of Biological Resources, Korea. Some micro-filamentous groups were cultured at temperature 20°C, using fluorescent lighting of 10-20 μmol photons m-2 s-1 with 14:10 LD photoperiod. For microspopic observations, the thalli were stained with 1% aniline blue solution, and mounted in 15-30% Karo syrup mixed with phenol.

Taxonomic Notes

Subclass Nemaliophycidae Christensen 1978. Guk-su-na-mul-a-gang (국수나물아강)

Thallus organization variable, from filamentous, multiaxial pseudoparanchymtous to parenchymatous. Plastids various in shape, with or without pyrenoid. Golgi association with endoplasmic reticulum and mitochondria. Pit plug with two caps on the cytoplasmic faces of the pit plugs, with or without membrane. Reproduction asexual by monosporangia or sexual by simple type carpogonium without a carpogonial branch or axiliary cell. Life cycles biphasic or triphasic.

REMARKS: See introduction part of this text.

Order Acrochaetiales Feldmann emend. Harper et Saunders 2002: 470. Bul-geun-som-mok (붉은솜목)

Thalli filamentous, simple or branched. Plastid variable from single axial stellate with a pyrenoid (Acrochaetium type) or multiple parietal discoid to band-shaped without pyrenoid (Rhodochorton type), or to multiple parietal lobed to discoid without pyrenoid (Audouinella type), or spiral (Grania type). Asexual reproduction primarily by monosporangia (Acrochaetium, Audouinella), or asexual tetrasporangia (Rhodochorton). Sexual life histories biphasic when known, heteromorphic alternation of generations with reduced gametophyte (Acrochaetium, Rhodochorton), or triphasic isomorphic alternation of generations.

REMARK: A contemporary classification scheme is proposed base on recent DNA sequences phylogenic researches (Harper and Saunders 1998, 2002; Le Gall and Saunders 2007; Clayden and Saunders 2008). DNA sequencing phylogenic results led to the traditional Acrochaetiales being divided into two orders, Acrochaetiales and Colaconematales (Harper and Saunders 2002), and also provided new generic delineations of some acrochaetioid genera. Furthermore, some traditional marine acrochaetioid represents were separated into new order Rhodachlyales (West et al. 2008) or new families Rhodothamniellaceae (Saunders et al. 1995) and Meiodiscaceae (Clayden and Saunders 2010) into Palmariales. In this work, the new concepts of acrochaetioid classification scheme are accepted for Korean representatives. 10 Algal Flora of Korea·Nemalian Red Algae

Family Acrochaetiaceae Fritsch emend. Harper et Saunders 2002: 470. Bul-geun-som-gwa (붉은솜과)

Characters as for the order. Type genus: Audouinella Bory de Saint-Vincent 1823: 340.

GENERA AND SPECIES: 4 genera (ca. 150, but uncountable at present). DISTRIBUTION: Worldwide in marine to freshwater. KEY REFERENCE: Harper and Saunders (2002).

Key to the genera of family Acrochaetiales

1. Monosporangia unknown, reproducing primarily by asexual tetrasporangia; where known, sexual life history biphasic and heteromorphic with reduced gametophytes··········Rhodochorton - Monosporangia common, primarily reproduction sources; where known, sexual life history triphassic and isomorphic with similar morphologically gametophytes···································2 2. Living in freshwater, plastid multiple parietal lobed to discoid without pyrenoid ······················ ·······································································································································Audouinella - Living in marine, plastid single axial stellate with a conspicuous central pyrenoid ····················· ·······························································································································Acrochaetium

REMARKS: The acrochaetioid red algae possess simple vegetative and reproductive structures. The vegetative thalli are usually heterotrichous, simple or branched monosiphous filaments composed of relatively smaller sized cells than other Florideoidean algae. Most members of this group repro- duce asexually by production of monospores. Where sexual reproduction is known, a carpogonial branch is absent and the fertilized carpogonium develops directly into a carposporophyte without the formation of an auxiliary cell. It is this apparent simplicity that arguably has led to one of the most chaotic taxonomic histories in red algal systematics. Nevertheless, numerous species (ca. 400) were described in this simple morphological group “Acrochaetioid”. Thus, an attempt to identify or describe new species leads one into a maze of literature and specific epithets. At least 17 generic names have been used in association with acrochaetioid algae: Acrochaetium, Audouinella, Balbiania, Byssus, Chantransia, Chantransiella, Chromastrum, Colaconema, Grania, Kylinia, Liagorophila, Pseudoacro- chaetium, Pseudochantransia, Rhodochorton, Rhodothamniella, Thamnidium and Trentepohlia. Of these, only five are currently available for use with species of Acrochaetiaceae: Acrochaetium, Audouinella, Grania, Kylinia, Rhodochorton. At least ca. 20 different classification schemes involving taxa of generic and higher rank have been proposed and these concepts have ranged from recognizing a single genus (Rosenvinge 1909; Drew 1928; Garbary 1978) to many as eight genera (Feldmann 1962). These discrepancies in the literature have led to multiple concepts associated with each genus. For example, Drew (1928) advocated a monogeneric Acrochaetiaceae and placed all marine and freshwater acrochaetioid taxa within Rhodochorton. Thus, this genus comprised species in which the plastids varied from one to many per cell, and were parietal, stellate, ribbon-shaped, discoid or radiating lobes. On the other hand, Papenfuss (1945) and Feldmann (1962) recognized multiple genera and restricted Rhodochorton to asexual marine species possessing few to many discoid Nemaliophycidae: Acrochaetiales: Acrochaetiaceae 11 plastids, and others arranged into Acrochaetium and Audouinella. Y.P. Lee (1987) distinguished three generic schemes based on reproductive mode and plastid morphology; Acrochaetium (monosporangia/triphasic sexual life history/stellate plastid), Audouinella (monosporangia/triphasic life history/ parietal laminate or ribbon shaped), and Rhodochorton (non-monosporangia/diphasic life history) base on Korean materials.

Genus Acrochaetium Nägeli enmed. Harper et Saunders 2002: 470. Bul-geun-som-sok (붉은솜속)

Thalli monosiphonous, simple or branched, epiphytic on various marine algae, heterotrichous with prostrate and erect filamnets, or only erect filaments with a single celled mucilagious attaching basal cell. Growth apical. Branches subdichotomous when present. Plastid single in each cell, stellate with single conspicuous central pyrenoid. Unicelluar hairs occurring frequently. Asexual reproduction predominantly by monospores. Sexual reproduction, if present, with a triphasic life history. Type species: Acrochaetium secundatum (Lyngbye) Nägeli in Nägeli et Cramer 1858: 532.

SPECIES ca. 100 spp. (10 spp. in Korea; Y. Lee 1987). DISTRIBUTION: Cosmopolitan, widely distributed in temperate and tropical waters. KEY REFERENCE: Harper and Saunders (2002), Y. Lee (1987).

Key to the species of genus Acrochaetium

1. Basal system composed of a single cell·····················································································2 - Basal system composed of multicellular branched filaments ····················································4 2. Erect filaments decumbent, simple usually with terminal hair······································A. scapae - Erect filaments upright and arcuate by secundly arranged branches ········································3 3. Branches terminate with one-celled hairs, monosporangia solitary ··················A. microscopicum - One-celled hair absent, monosporangia concatenate············································A. catenulatum 4. Thallus minute, simple or sparsely branched, epiphytic on uticles of Codium···························5 - Thallus much branched············································································································6 5. Erect filaments simple with terminal monosporangia ··············································A. terminale - Erect filaments branched in low portion with lateral monosporangia···························A. inkyui 6. Monopsorangial short laterals arranged oppositely ·································································7 - Laterals arranged alternately to secundly·················································································8 7. Erect filaments usually terminate with elongate one-celled hairs ·······················A. sancti-thomae - Erect filaments tapered without one-celled hairs ···················································A. virgatulum 8. Erect filaments branched secundly without formation of terminal hairs···································9 - Erect filaments branched alternately or irregularly with formation terminal hairs······················· ································································································································A. canariense 9. Cells of erect filaments 7-9 μm broad and basal system loosely coherent ···················A. densum - Cells of erect filaments 10-12 μm and basal system closely coherent to form disc ······················· ·····························································································································A. secundatum 12 Algal Flora of Korea·Nemalian Red Algae

REMARKS: The generic circumscription of Acrochaetium has been continuously changed as in case of other genus of acrochaetioid. According to the protologue of Nägeli (1861: 402), Acrochaetium was characterized by the branched filaments which usually arose from monostromatic crustose basal disc and reproducing by means of zoospores (=monospores). Rhodochorton Nägeli (1861: 355), which established in the same paper, was distinguished by the branched prostrate basal filaments and reproducing by means of tetraspores from Acrochaetium. After that, Acrochaetium included all asxual acrochaetioid algae (Bornet 1904; Collins 1906) or all marine acrochaetioid algae (Børgesen 1915). Later, Papenfuss (1945) reviewed Acrochaetium and related gerera on the focus of plastidal feature as another criterion and that he restricted the genus Acrochaetium whose cells contain one parietal or laminate plastid. Originally, the generally cited lectotype species of Acrochaetium was A. daviesii (Papenfuss 1945). However, Woelkerling (1983) designated A. secundatum as the lectotype. Harper and Saunders (2002) confirmed that the generic type species is clustered with Audouinella hermannii and Rhodochorton purpureum as a monophyletic clade (=Acrochaetiaceae) separted from Colaconema clade (=Colaconemataceae) on the basis of SSU-LSU rDNA sequence phylogeny. Now, Acrochaetium is defined restrictly by the presence of cells with a single stellate plastid with a central pyrenoid as observable in A. secundatum.

1. Acrochaetium canariense Børgesen 1927: 17 (Figs. 2, 3). Da-bal-bul-geun-som (다발붉은솜: 개칭)

Børgesen 1927: 17. f. 9-11. Lee 1987: 24. f. 10A-F.

SYNONYM: Audouinella canariensis (Børgesen) Garbary 1979: 490. Chromastrum canariense (Børgesen) Stegenga et Mulder 1979: 305.

Thalli epiphytic, caespitose, composed of basal and erect filaments, 500-850 μm high. Basal filaments creeping, irregularly and more or less coherently branched, forming pseudoparenchymatous discs. Erect filaments branched irregualarily to subdichomo- tously in middle to upper portion, oftenly terminating with one-celled hairs. Cells of erect filaments cylindrical, not constricted at jungtion, 7-9 μm broad, and 9-13 μm long. Plastids stellate with a central pyrenoid. Monosporangia lateral or terminal, sessile or on one to two- celled stalks, born in secund arrangement, solitary on a stalk cell, ovoid, ca. 9 μm in diameter.

TYPE: Playa de Santa Catalina, Canary Islands (?). SEASONALITY: Winter to Summer (see Lee 1987). DISTRIBUTION: Worldwide (?). Fig. 2. Distribution of Acrochaetium cana- KOREA: East coast. riense. Nemaliophycidae: Acrochaetiales: Acrochaetiaceae 13

ABC

Fig. 3. Acrochaetium canariense. A. Habit of detached thalli from Zostera sp.; B, C. Detail of branching mode and monospoangia and one-celled hairs.

SPECIMEN EXAMINED: KN1047 (Deoksan harbor, Samcheok: 16.vii.2009). REMARKS: Acrochaetium canariense was reported in Korea by Lee (1987) from Youngil Bay in Korea which grown on Codium fragile. However, the Lee’s specimens had not terminal hairs on erect filaments and different in branching mode compared to protologue of this species. According to Børgesen (1927), the original specimens are small about 350 μm high, epiphytic on Gelidium sp., and the erect filaments subdichotomously branched with one-celled terminal hair. Our specimens are almost similar with the descriptions and illustrations given by Børgesen (1927, f. 10). However, this species is very similar with Acrochaetium densum except the presence of terminal hairs, thus further studies are needed to certify the independence of both species.

2. Acrochaetium catenulatum Howe 1914: 84 (Figs. 4, 5). Ae-gi-son-bul-geun-som (애기손붉은솜: 개칭)

Howe 1914: 84-86. pl. 31. f. 12-18. Stegenga et al. 1997: 236. pl. 72. f. 3, 4.

SYNONYM: Acrochaetium microscopicum sensu Lee 1987: 30. f. 14. Acrochaetium moniliforme sensu Lee 1987: 32. f. 15A-C (exclude f. 15D-G.). Audouinella catenulata (Howe) Garbary 1979: 490. Chantransia catenulata (Howe) De Toni 1924: 44. Chromastrum catenulatum (Howe) Stegenga and Mulder 1979: 299. Kylinia catenulata (Howe) Kylin 1944: 13. Rhodochorton catenulatum (Howe) Nakamura 1941: 273. 14 Algal Flora of Korea·Nemalian Red Algae

ABDE

FG H

Fig. 4. Acrochaetium catenulatum. A-E. Field plants on Sphacelaria sp.; F-H. Cultured plants in laboratory, attached by mucilaginous basal cell.

Thalli epiphytic usually on Sphacelaria sp., with single-celled base and usually a single erect filament, 50-100 μm high in field. Basal attachting cell single, globose and slightly appressed, having thick mucilaginous refractive wall in attaching portion. Erect filaments much branched, heavily arcurated by first and second branchlets which arranged secundly, terminating with monosporan- Nemaliophycidae: Acrochaetiales: Acrochaetiaceae 15 gia without hairs. Cells of erect filaments barrel shaped to moniliform, 5-10 μm wide, 3-7 μm long. Plastid stellate with a central pyrenoid. Hairs absent in wild and field materials. Monosporangia terminal or lateral on erect filaments and branches, solitary or concatenate with 2-3 sporangia, ovoid, 5-7 μm wide, 6-9 μm long. Sexual reproduction not observed.

TYPE: La Punta, Peru (Holotype: New Botanical Gardner ?). SEASONALITY: Rarely observed in summer (June to September). DISTRIBUTION: Worldwide cosmopolitan. KOREA: South and east coasts. SPECIMEN EXAMINED: KN1041 (Sanggye islet, Yeosu: 24.vi.2009), KN1042 (Dongsan harbor, Yangyang: 18. viii.2009). REMARKS: The type materials of Acrochaetium catenulatum were epiphytes on Chaetomorpha sp. (Howe 1914), but Korean materials are usually occur on Sphacelaria sp. growing on Sargassum sp. in this studies. Our specimens in wild and room cultures are almost identical with the original descriptions of Howe (1914), Fig. 5. Distribution of Acrochaetium cate- who characterized his new species as having one erect nulatum. filament and concatenate monosporangia without terminal hairs. Woelkerling (1971; 1972) synonymized this species with A. microscopicum. However, A. catenulatum is distinguished from A. microscopicum by producing concatenate monosporangia without terminal hairs and arcuated axis by first and second ordered secundly arranged branches. Lee (1987) reported A. moniliforme which attached on Chaetomorpha sp., however, some of his specimens (Lee 1987, f. 15A-D) to be considered a misidenti- fication of this species because they had not terminal hairs. Stegenga and Vroman (1976) regarded A. catenulatum to be the gametangial plant of A. densum. Thus, the taxonomic entity of this species is obscure at present and needed to be certified base on molecular studies.

3. Acrochaetium densum (Drew) Papenfuss 1945: 308 (Figs. 6, 7). Ae-gi-bul-geun-som (애기붉은솜: 개칭)

Lee 1987: 27. f. 12. Lee and Yoshida 1997: 181. f. 8.

BASIONYM: Rhodochorton densum Drew 1928: 168. pl. 38. f. 17-24. Kang 1966: 59. Lee and Lee 1974: 38. f. 2. SYNONYM: Acrochaetium canariense sensu Lee 1987: 24. f. 10. Audouinella densa (Drew) Garbary 1979: 490. Chromastrum densum (Drew) Stegenga and Mulder 1979: 299. Rhodochorton densum Drew 1928: 168. 16 Algal Flora of Korea·Nemalian Red Algae

ABC

DEF

Fig. 6. Acrochaetium densum. A-D. Habit of detached thalli from Sargassum fusiforme (Gangneung); E, F. Cultured plants isolated from fishing wire from Incheon harbor.

Thalli epiphytic, caespitose, composed of basal and erect filaments, 400-700 μm high. Basal filaments creeping, coherently branching, forming pseudoparenchymatous and monostromatic basal discs. Erect filaments scarcely tapering towards apices, issuing branches secundly on upper portion, more or less feeble. Cells of erect filaments cylindrical, slightly constricted at junction, 7- Nemaliophycidae: Acrochaetiales: Acrochaetiaceae 17

9 μm wide, 9-13 μm long. Plastids stellate with a central pyrenoid. Monosporangia terminal or lateral on branches and erect filaments, sessile or rarely on one-celled stalks, ovoid to ellipsoid, 8-10 μm wide, 10-12 μm long.

TYPE: Fort Point, San Francisco, California, U.S.A (UC no. 294560; Gardner no. 4607; Herb. Univ. of Cali- fornia, Berkely, California). SEASONALITY: Summer (?, Lee 1987). DISTRIBUTION: Worldwide. KOREA: Around all coasts of Korea. SPECIMEN EXAMINED: KN1043 (Incheon harbor: 9.i. 2009), KN1045 (Sacheon, Gangneung: vi.2009). REMARKS: Acrochaetium densum was reported since Kang (1966) and has been listed up from several local- ities in Korea (see Lee and Kang 2001). The morphological characteristics of Korean materials were illustrated by Lee and Lee (1974, f. 2; as named Rhodochor- ton densum) and taken another close look by Lee (1987, f. 12), which agree quite well with the description and illustrations given by Drew (1928). It is considered Fig. 7. Distribution of Acrochaetium den- that the described specimens of Acrochaetium canari- sum. ense from Korea (Lee 1987: 24. f. 10) to be a tetasporo- phytic stage of this species. The plastid morphology of this species, which has been ambiguous since Drew, was discussed in detail by Lee (1987). And also, Lee and Yoshida(1997) reported the phenological observations of this species from Hokkaido.

4. Acrochaetium inkyui Lee 1987: 27 (Figs. 8, 9). Je-ju-bul-geun-som (제주붉은솜: 개칭)

Lee 1987: 27. f. 13A-F.

Thalli epiphytic on Codium fragile, caespitose, composed of basal prostrate and erect filaments, 40-120 μm high. Basal prostrate filaments creeping, branching subdichotomously, more or less confluent together, forming a pseudoparenchymatous monostromatic basal disc. Erect filaments composed of 7-12 cells, simple or subdichotomously branched in lower portion. scarcely tapering towards apex without terminal hairs. Cells of erect filaments cylindrical to barrel-shaped, 6-8 μm wide, 6-13 μm long. Plastid stellate with a central pyrenoid. Monosporangia terminal or lateral on erect filaments, solitary, sessile or pedicellated, globose, 10-11 μm in diameter. Sexual reproduction not observed.

TYPE: Seongsan, Jeju, Korea (slide LYP 185-5, Herbarium of the CNU, see Lee 1987). SEASONALITY: Autumn to early winter. 18 Algal Flora of Korea·Nemalian Red Algae

ABC

DE FG

Fig. 8. Acrochaetium inkyui. A. Epiphytic on the tip of utricles of Codium fragile show prostrate basal system (arrow) and simple erect filaments (arrow head); B. Some erect filaments branched subdichotomously in low portion (arrow); C. Monosporangia produced laterally on erect filaments (arrow); D-G. Developmental series from earlier (D) to mature stage (F) and liberation of monospore (G; arrow). Nemaliophycidae: Acrochaetiales: Acrochaetiaceae 19

DISTRIBUTION: Korea. KOREA: South and east coasts. SPECIMEN EXAMINED: KN1044 (Jangilri, Pohang: 22. xii.2009). REMARKS: Acrochaetium inkyui was described as a new species from Jeju Isalnd (Lee 1987). This reporting is the second time of this species from Korea. This species are growing restrictly on the tip of the utricles of Codium fragile usually with together Colaconema codicola. According to the description and figures given by Lee (1987), the monosporangia of this species are sessile on simple erect filaments. However, some specimens show subdichotomous branching in lower portion and pedicellate lateral monosporagia on erect filaments in our observation. In spite of this discre- pancy, our specimens are treated to this species based on the overall similarity between them. This species is very similar with A. terminale (Nakamura) Lee (1987), which is clearly distinguished by the terminal monosporangia contrast with the lateral monosporangia of this species (see Lee 1987).

Fig. 9. Distribution of Acrochaetium inkyui.

5. Acrochaetium microscopicum (Nägeli ex Kützing) Nägeli 1861: 407 (Figs. 10, 11). Gul-geun-ja-ru-bul-geun-som (굵은자루붉은솜: 개칭)

Woelkerling 1972: 85. f. 1-14.

BASIONYM: Callithamnion microscopicum Nägeli ex Kützing 1849: 640. SYNONYM: Acrochaetium catenulatum sensu Lee 1987: 24. f. 11A-G. Acrochaetium crassipes (Børgesen) Børgesen 1915: 20. Audouinella crassipes (Børgesen) Garbary 1979: 490. Audouinella microscopica (Nägeli ex Kützing) Woelkerling 1971: 9. Audouinella moniliformis (Rosenvinge) Garbary 1979: 490. Chantransia crassipes Børgesen 1909: 1. Chantransia microscopica (Nägeli ex Kützing) Batters in Schiffner 1916: 136. Chantransia moniliformis Rosenvinge 1909: 99. f. 28-29. Chromastrum crassipes (Børgesen) Papenfuss 1945: 321. Chromastrum microscopicum (Nägeli ex Kützing) Papenfuss 1945: 322. Chromastrum moniliforme (Rosenvinge) Papenfuss 1945: 322. Kylinia crassipes (Børgesen) Kylin 1944: 13. 20 Algal Flora of Korea·Nemalian Red Algae

A B C

DE F

Fig. 10. Acrochaetium microscopicum. A-C. Epiphytic thalli habits on Sphacelaria sp.; D-F. Cultured plants in laboratory (Sanggye islet, Yeosu).

Kylinia microscopica (Nägeli ex Kützing) Kylin 1944: 13. Kylinia moniliformis (Rosenvinge) Kylin 1944: 13, 16. Rhodochorton microscopicum (Nägeli ex Kützing) Drew 1928: 163. Rhodochorton moniliforme (Rosenvinge) Drew 1928: 164. Nemaliophycidae: Acrochaetiales: Acrochaetiaceae 21

Thalli epiphytic usually on Sphacelaria sp., composed of a single-celled base and an erect filament, 50 μm high. Basal attacting cell single, globose and slightly appressed, having thicker mucilaginous refractive wall toward base, 5-6 μm wide, 4-5 μm long. Erect filaments usually born singly on a basal cell, 5-7 celled long, simple or with secund branches, terminating with monosporangia or one-celled hairs. Branches rare in field, usually arose secundly from every cells in culture. Cells of erect filaments oblong to barrel-shaped, slightly longer towards apex, 5-6 μm wide, 4-5 μm long. Plastid stellate with a central pyrenoid. Monosporangia sessile or pedicellate on an one-celled stalk, lateral or terminal on erect filaments or branches, globose to ovoid, 5-7 μm wide, 6-7 μm long.

TYPE: Torquey, Devon England(Holotype: L. no. 940, 285, 306; Rijksherbarium, Leiden, Netherlands; cf. Woelkerling 1972, 1973b; Dixon and Irvine 1977). SEASONALITY: Rarely observed in summer (June to September). Fig. 11. Distribution of Acrochaetium mic- DISTRIBUTION: Worldwide cosmopolitan. roscopicum. KOREA: South and west coasts. SPECIMEN EXAMINED: KN1046 (Udo, Jeju: 4.vi.2009), KN1103 (Sanggye islet, Yeosu: 22.vi.2009). REMARKS: Korean plants of Acrochaetium microscopicum were found epiphytic on Sphacelaria species growing on Sargassum thunbergii or epizoic on hydroids in western and southern coasts and Jeju Islands (Lee 1987). Woelkerling (1972) has presented a detailed account of this species including description sexual stages and he synonymized six species related to A. microscopicum after examina- tions of their type collections; A. crassipes Børgesen, A. catenulatum Howe, A. collopodum (Rosenvinge) Kylin, A. compactum Jao, and A. moniliforme (Rosenvinge) Børgesen. Stegenga and Mulder (1979) and Lee (1987), however, did not accept the Woelkerling’s suggestion of conspecificity. Our specimens of A. microscopicum are quitely identical with the type materials which were reexamined by Woelkerling(1972, f. 1-6). In this study, we confirmed the morphological variations of A. microscopicum base on field and room cultured materials. A. microscopicum is characterized by a single basal cell showing almost same morphology as that of vegetative cells, 5-10 celled short erect filaments, secundly branching and oftenly teminating with one-celled hair. These charcters are considered to be stable which are expressed in both conditions in wild and room cultures. And also, these features are almost identical with the diagnoses of Chantransia moniliformis (Rosenvinge 1909, f. 28) and A. crassipes (Børgesen 1915, f. 11-13). However, the Korean samples of A. catenulatum, which is almost identical with the original description (Howe 1914, f. 12-18), is clearly distinguished by the hairless filaments and concatenate monosporangia (see Fig. 4). Thus, we considered that C. moniliformis Rosenvinge and A. crassipes Børgesen should be treated as synonyms of this species. 22 Algal Flora of Korea·Nemalian Red Algae

Lee (1987) made a mistake in identifications of A. catenulatum, A. microscopicum and A. moniliforme in Korean materials. His figures of A. microscopicum (Lee 1987, f. 14A-F) are very similar with A. catenulatum which has no terminal hairs. Furthermore, his A. moniliforme (Lee 1987, f. 15) is also very different from the original protologue (Rosenvinge 1909, f. 28) by having opposite branches without terminal hairs (Lee 1987, f. 15E-G).

6. Acrochaetium sancti-thomae Børgesen 1915: 30 (Figs. 12, 13). Cheon-sa-bul-geun-som (천사붉은솜: 개칭)

Børgesen 1915: 30. f. 23, 24. Nakamura 1941: 280. f. 5-7. Lee and Lee 1974: 39. f. 3a-e. Lee 1987: 34. f. 16.

SYNONYM: Audouinella sancti-thomae (Børgesen) Garbary 1979: 490. Chantransia sancti-thomae (Børgesen) De Toni 1924: 53. Rhodochorton sancti-thomae (Børgesen) Nakamura 1941: 280.

Thalli epiphytic, caespitose, composed of basal and erect filaments, 350-500 μm high. Basal filaments creeping, irregularly and more or less coherently branched, forming pseudoparenchymatous discs. Cells of basal filaments oblong to fusi- form in side view, 4-7 μm broad, 5-7 μm long. Erect filaments simple or seldomly producing laterals from lower portion, producing frequently short determi- nate branches secundly or oppositely which producing monosporangia, usually terminating one-celled hairs. Cells of erect filaments cylindrical to club- shaped, constricted at jungtion, gradually shortened toward apices, 8-13 μm broad, and 13-15 μm long. Plastids stellate with a central pyrenoid. Monospo- rangia lateral or terminal on short branches, sessile or on one-celled stalks, born in secund arrangement, solitary on a stalk cell, ovoid, ca. 8 μm wide, 11-13 μm long.

TYPE: St. Thomas, Virgin Islands (?). SEASONALITY: Summer (?, see Lee 1987). DISTRIBUTION: Worldwide (?). KOREA: Around all coasts of Korea. SPECIMEN EXAMINED: KN1048 (Sacheon, Gangneung: 18.vi.2009). REMARKS: Acrochaetium sancti-thomae was reported in Korea by Lee and Lee (1974) and Lee (1987) from Jeju Island which grown on Sargassum fusiforme or Fig. 12. Distribution of Acrochaetium Myelophycus simplex. However, our specimens are sancti-thomae. Nemaliophycidae: Acrochaetiales: Acrochaetiaceae 23

ABC

DEFG

Fig. 13. Acrochaetium sancti-thomae. A. Habit of detached thalli from Phyllospadix sp.; B, C. Detail of branching mode and axial cell shape; D. Detail of base; E, F. Detail of terminal hairs and mature monosporangium (arrow); G. Liberated monospore (arrow).

different from those of Jeju Island. Furthermore, our specimens are also very similar with the description and illustration of Acrochaetium densum provided by Garbary et al. (1982, f. 7). Accord- ing to Børgesen (1915), the original specimens are small about 200-300 μm high, the erect filaments usually terminating with long hairs and slightly branched or often not at all, but the short branch- 24 Algal Flora of Korea·Nemalian Red Algae lets on the other hand are often present. Moreover, the short branchlets of our specimens are usually arranged oppositely as like in original specimens. Thus, the species boundary for A. sancti- thomae used here is strictly restricted to the description of Børgesen (1915) excluding some confused descriptions.

7. Acrochaetium scapae (Lyle) Papenfuss 1945: 307 (Figs. 14, 15). Bi-nyeo-bul-geun-som (비녀붉은솜: 개칭)

Papenfuss 1945: 307. Lee 1987: 36. f. 17A-H.

BASIONYM: Kylinia scapae Lyle 1929: 245. SYNONYM: Audouinella scapae (Lyle) Dixion in Parke and Dixon 1976: 590. Dixion and Irvine 1977: 113. f. 38.

Thalli epiphytic, composed of a single-celled base and two decumbent opposite erect filaments, to 20 μm high. Basal cell globose to subglobose, 7-8 μm in diameter, usually producing two decumbent filaments oppositely. Erect filaments simple, decumbent, 2-8 celled long, terminating with one-celled hairs. Cells of erect filaments barrel-shaped to moniliform, 4-6 μm wide, 5-8 μm long. Plastids stellate with a central pyrenoid. Hairs short, terminal on erect filaments, ca. 1 μm

AB C D

Fig. 14. Acrochaetium scapae. A. Epiphytic decumbent habit on Sphacelaria sp (arrow); B. Single- celled basal cell (arrow); C. Two decumbent filaments from single basal cell (arrow); D. Terminal hair (arrow). Nemaliophycidae: Acrochaetiales: Acrochaetiaceae 25 thick, to 60 μm long. Monosporangia sessile, terminal or lateral, ovoid to ellipsoid, 4-5 μm wide, 6-7 μm long.

TYPE: Orkney, Scapa Flow, England(BM, slide 3713). SEASONALITY: Summer (see Lee 1987). DISTRIBUTION: Worldwide cosmopolitan. KOREA: South coast. SPECIMEN EXAMINED: KN1049 (Dwiyeo islet, Yeosu, 22.vi.2009). REMARKS: Acrochaetium scapae was reported one time by Lee (1987), and this reporting is the second time in Korea. The morphological features of our specimens are almost similar with the description and photographes of Lee (1987, f. 17), but there are some differences in the cell dimensions and presence of terminal hairs. Furthermore, Korean specimens are different from the type materials which reexamined by Dixon and Irvine (1977, f. 38). Thus, further studies need to determine the status of this species including A. microscopicum more precisely although their con- tinued independence is highly unlikely. Fig. 15. Distribution of Acrochaetium scapae.

8. Acrochaetium secundatum (Lyngbye) Nägeli in Nägeli et Cramer 1858: 532 (Figs. 16-18). Eol-rae-gi-bul-geun-som (얼레기붉은솜: 개칭)

Coppejans and De Vos 1979: 64. pl. 1. f. 1-18. pl. 2. f. 1-6. Kornmann and Sahling 1977: 176. f. 96. Stegenga et al. 1997: 240. pl. 73. f. 4, 5. Lee 1987: 35. f. 18.

BASIONYM: Callithamnion daviesii var. secundatum Lyngbye 1819: 129. SYNONYM: Audouinella secundata (Lyngbye) Dixon in Parke and Dixon 1976: 590. Ceramium secundatum (Lyngbye) C. Agardh 1824: 132. Chantransia secundata (Lyngbye) Thuret in Le Jolis 1863: 106. Chromastrum secundatum (Lyngbye) Papenfuss 1945: 323. Colaconema secundatum (Lyngbye) Woelkerling 1973a: 94. Kylinia secundata (Lyngbye) Papenfuss 1947: 437.

Thalli epiphytic, caespitose, with basal creeping and erect filaments, to 1 mm height. Basal creeping filaments coherently branching, forming a pseudoparenchymatous basal disc. Cells of creeping filaments rather smaller than the cells of erect filaments, oblong to globose in shape seen from below, 6-7 μm wide, 6-8 μm long. Erect filaments much branched, arcuated by secundly arranged branches, sometimes grown sympodially by terminal monosporangium. Cells of erect filaments 26 Algal Flora of Korea·Nemalian Red Algae

ABC

DE F

Fig. 16. Acrochaetium secundatum. A. Habits of detached thalli from Phyllospadix sp.; B. Detail of secundly arranged branches; C, D. Sympodial growing axis by the formation of terminal monosporangium; E. Basal system and erect axis; F. Detail of stellate plastid with a central pyrenoid and monosporangium. Nemaliophycidae: Acrochaetiales: Acrochaetiaceae 27

ABCD

EFG

Fig. 17. Acrochaetium secundatum. A-G. Room culturing plants. A. Habits of crustose basal and erect filaments; B-D. Developmental series from monospores; E, F. Secundly arranged branches; G. Detail of stellate plastid with a central pyrenoid. 28 Algal Flora of Korea·Nemalian Red Algae cylindrical to rectangular, slightly constricted at junc- tions, 10-12 μm wide, 15-20 μm long. Plastids stellate with a central pyrenoid, occupying the distal part of cells. Hairs absent. Monosporangia sessile or on one- or two-celled stalks, usually sollitary, born in secund arrangment, ellipsoid to ovoid, 8-9 μm wide, 12-16 μm long.

TYPE: Quiving, Faeroes Isl. (cf. Dixon and Irvine 1977). SEASONALITY: Observed June to October. DISTRIBUTION: Worldwide. KOREA: East coast. SPECIMEN EXAMINED: KN1051 (Ingu, Namae, Gang- neung: 26.viii.2009). Epiphytic on Sargassum fusiforme or other algae. REMARKS: Acrochaetium secundatum is easily distinguished from other Korean acrochaetioid algae by secund branching, typical stellate plastid and pyrenoid, and bearing monosporangia in a secund or pin- nate arrangement. A multilayered basal disc of this species was reported by several authors and a mono- Fig. 18. Distribution of Acrochaetium sec- layered one by others (Børgesen 1902; Kylin 1907; undatum. Rosenvinge 1909; Hamel 1927; Taylor 1957; Woelkerling 1973a, 1973b; Pedersen 1976; Dixon and Irvine 1977). Børgesen (1902) and Woelkerling (1973a) regarded the multilayered basal system as distinctive character of the species. However, Woelkerling(1973b: 580) recorded the basal system of the species as “... but the majority are single layered.” Such a multilayered basal system was not observed in the plants from Korea. The developmental series from monospores of this species was illustrated in detail by Coppejans and De Vos (1979) which are almost identical with our observations (Fig. 17). Woelkerling (1973b) synonymized A. virgatulum with A. secundatum after examination of the type collections of the two taxa. Dixon and Irvine (1977) thought the two taxa to be related each other, but treated them independently pending further investigations. The plants of the two taxa examined here can be easily distinguished by the branching pattern. Woelkerling (1983) designated A. secundatum as the lectotype species of the genus Acrochaetium.

9. Acrochaetium terminale (Nakamura) Lee 1987: 39 (Figs. 19, 20). Sang-tu-bul-geun-som (상투붉은솜: 개칭)

Lee 1987: 39. f. 19.

BASIONYM: Rhodochorton terminale Nakamura 1944: 99. f. 1. Lee and Lee 1974: 37. f. 1. SYNONYM: Audouinella terminalis (Nakamura) Garbary 1979: 490. Yoshida 1998: 469. Kylinia terminalis (Nakamura) Noda and Yokayama 1971: 17. Nemaliophycidae: Acrochaetiales: Acrochaetiaceae 29

AB C

DE F

Fig. 19. Acrochaetium terminale. A, B. Habits of crustose basal cells and upright erect filaments; C. Basal creeping cells forming crustose patches on terminal portion of utricles; D. Upright erect filaments from crustose basal cells; E. Producing monosporangia from terminal cell of erect filaments; F. Simple erect filament without branchs. 30 Algal Flora of Korea·Nemalian Red Algae

Thalli epiphytic usually on utricles of Codium contractum, caespitose, forming patch-like specklings, composed of creeping basal crustose cells and upright erect filaments, upto 200 μm hight. Creeping basal filaments developing radially from the daughter cells of germinating spores, coherently branching, forming a monostromatic and crustose basal discs. Cells of creeping filaments various in shape, 2-4 μm wide, 4- 6 μm long. Erect filaments arising from the basal cell from the middle region of the basal disc, usually simple or rarely branched on upper part. Cells of erect filaments cylindrical to barrel-shaped, 5-7 μm wide, 5-10 μm long. Plastids stellate with a central pyrenoid. Monosporangia terminal or subterminal, or rarely lateral near the apices of erect filaments, ellipsoid, 7-8 μm wide, 10-12 μm long. Tetrasporangia or sexual reproductive structures not observed.

TYPE: Hayama, Japan (SAP: Herbarium, Department of Botany, Hokkaido University, Sapporo, Japan). SEASONALITY: Winter to spring. DISTRIBUTION: Korea, Japan. Fig. 20. Distribution of Acrochaetium ter- KOREA: Jeju Island. minale. SPECIMEN EXAMINED: KN5237 (Seobsum islet, Jeju: 7.i.2009, Epiphytic on the tips of the utricles of Codium contractum). REMARKS: Acrochaetium terminale is growing on tips of the utricles of Codium contractum in Korea rather than on C. subtubulosum in Japan (Nakamura 1944), but Korean specimens collected at Jeju Island agree well with the lectotype collections (Lee 1987; this work), except the branching pattern. This species shares some morphological characteristics with A. inkyui and A. humile (Rosenvinge) Børgesen, especially in sharing the growth on tips of uticles of Codium. However, this species is clearly distinguished by the simple erect filaments and the terminal monosporangia from A. inkyui and also differed from A. humile by the absent of terminal uni-cellular hair (see Lee and Yoshida 1997: 184). Lee and Lee (1974) described the plastid morphology of A. terminale as parietal laminate, but it was revealed that the misreading of the stellate morphology (Lee 1987). Lee and Lee (1974) shown some branched thalli on upper portion of erect filament of this species, however, we did not observed the branched thalli.

10. Acrochaetium virgatulum (Harvey) Batters 1902: 58 (Figs. 21, 22). Mae-deub-bul-geun-som (매듭붉은솜: 개칭)

Kornmann and Sahling 1977: 174. f. 95A-F. Lee 1987: 39. f. 20.

BASIONYM: Callithamnion virgatulum Harvey 1833: 349. Nemaliophycidae: Acrochaetiales: Acrochaetiaceae 31

A BC

DE F

Fig. 21. Acrochaetium virgatulum. A, B. Habits of epiphytic thalli on Chaetomorpha moniligera; C. Thalli composed of prostrate (arrow) and erect (arrow head) filaments; D. Detail of prostrate basal system shown stellate plastid with a central pyrenoid (arrow); E. Erect filaments occasionally branched subdichotomously in lower portion (arrow); F. Detail of stellate plastid and oppositely arranged monosporangial stalk cells. 32 Algal Flora of Korea·Nemalian Red Algae

SYNONYM: Audouinella virgatula (Harvey) Dixon in Parke and Dixon 1976: 590. Chantransia virgatula (Harvey) Thuret in Le Jolis 1863: 106. Chromastrum virgatulum (Harvey) Papenfuss 1945: 323. Kylinia virgatula (Harvey) Papenfuss 1947: 437. Rhodochorton virgatulum (Harvey) Drew 1928: 152. Trentepohlia virgatula (Harvey) Farlow 1881: 109.

Thalli epiphytic, caespitose, composed of basal and erect filaments, to 600 μm high. Basal filaments creeping, coherently and alternately branched, forming monostromatic and pseudoparenchymatous basal discs. Erect filaments simple or subdichotomously branched, feeble, tapered to the apex without terminal hairs. Cells of erect filaments cylindrical to club- shaped, constricted at junction. 8-11 μm wide, 18-23 μm long. Plastids stellate with a central pyrenoid, occupying the distal part of cells. Monosporangia born secundly or oppositely, or occasionally helical in arrangement, solitary or in pairs on one-celled stalk or sessile, ellipsoid to ovoid, 6-8 μm wide, 9-10 μm Fig. 22. Distribution of Acrochaetium vir- long. gatulum.

TYPE: Devon shore, England (TCD; School of Botany, Trinity College, Dublin, Ireland). SEASONALITY: Autumn to winter. DISTRIBUTION: Worldwide. KOREA: South and East coasts (see Lee 1987). SPECIMEN EXAMINED: KN1053 (Sacheon, Gangneung: 22.xii.2009, epiphytic on Chaetomorpha moniligera). REMARKS: Acrochaetium virgatulum was recorded by Lee (1987) and this reporting is the second time in Korea. Korean plants of A. virgatulum agree quite well with the original description and figures (Harvey 1833, 1846: 314). Dixon (in Parke and Dixon 1976) and Dixon and Irvine (1977) regarded A. virgatulum to be conspecific with Acrochaetium secundatum (see also Woelkerling, 1973b). However, the two taxa are easily distinguished each other by the branching pattern (see Kornmann and Sahling 1977; this study). Borsje (1973) reported that the gametangial plants of A. virgatulum showed a morphology similar to Acrochaetium parvulum (Kylin) Hoyt or Acrochaetium rhipidandrum (Rosenvinge) Hamel under culture conditions. According to Lee (1987), the tetrasporangial plants of this species have short fructiferous laterals along the erect filaments and branches which are closely related to those of Acrochaetium plumosum (K.M. Drew) G.M. Smith. Nemaliophycidae: Acrochaetiales: Acrochaetiaceae 33

Genus Rhodochorton Nägeli emend. Harper et Saunders 2002: 471 Ja-ri-pul-sok (자리풀속)

Marine algal member not occuring freshwater. Thalli composed of monosiphonous simple or branched filaments. Plastid multiful parietal discoid to band-shaped per cell, lacking pyrenoids. Reproduction usually asexual, primarily by apparently asexual tetrasporangia, monosporangia unknown. Sexual reproduction, if present, with a biphasic life history and heteromorphic with reduced gametophyte. Type species: Rhodochorton purpureum (Lightfoot) Rosenvinge 1900: 75.

SPECIES uncountable at present (1 spp. in Korea). DISTRIBUTION: Cosmopolitan, widely distributed in temperate and tropical waters. KEY REFERENCE: Harper and Saunders (2002), Y. Lee (1987). REMARKS: The generic concept of Rhodochorton has varied greatly since Nägeli (1861: 355) described the genus for acrochaetioid algae reproducing tetraspores. Drew (1928) treated all described acrochaetioid algae as members of this genus. Drew’s single generic concept was connected to Dixon and Irvine (1977) and Garbary et al. (1982) with accepting Audouinella Bory (1823: 340) having priority over Rhodochorton Nägeli. Meanwhile, Papenfuss (1945), Kylin (1956), and Feldmann (1962) recognized 4 or 8 separated genera for acrochaetioid. Lee and Lee (1988) suggested three generic concepts as Acrochaetium, Audouinella and Rhodochorton based on the characteristics of life cycles and plastid morphology. However, the classical classification system of the acrochaetioid has been changed profoundly with evidences of molecular phylogenetic works (Harper and Saunders 2002; Clayden and Saunders 2008, 2010). Now, the genus Rhodochorton was certified by only one type species R. purpureum (Lightfoot) Rosenvinge (1900: 75). Within the Acrochaetiaceae, this genus is easily distinguished from Acrochaetium in lacking monosporangia and stellate plastids, from Grania by spiral or ribbon-shaped plastid, and from Audouinella in being marine and lacking monosporangia. In Korea, two Rhodochorton has been known after Lee (1987), R. purpureum and R. subimmersum Setchell et Gardner, however, the later was transferred to a new genus Rhodonematella under family Rhodophysemataceae of Palmariales based on molecular phylogenetic study (Clayden and Saun- ders 2010).

11. Rhodochorton purpureum (Lightfoot) Rosenvinge 1900: 75 (Figs. 23, 24). Yung-dan-ja-ri-pul (융단자리풀)

Lee 1987: 39. f. 19. Lee and Yoshida 1997: 199. f. 15. West et al. 2006: 103. f. 1-10.

BASIONYM: Byssus purpurea Lightfoot 1777: 1000. SYNONYM: Audouinella purpurea (Lightfoot) Woelkerling 1973: 536. f. 52-55. Callithamnion purpureum (Lightfoot) Harvey 1841: 116. Callithamnion rothii (Turton) Lyngbye 1819: 129. Ceramium rothii (Turton) Agardh 1824: 133. Conferva purpurea (Lightfoot) Dillwyn 1802-1809: 56. pl. 40. Conferva violacea Roth 1797: 190. pl. 4. f. 1. 34 Algal Flora of Korea·Nemalian Red Algae

ABCD

EF GH

Fig. 23. Rhodochorton purpureum. A-H. Room cultured plants (culture strain of J.A. West, No 4187; isolated from Jeju Island by Y.P. Lee); A. Habits of tetrasopophyte at 14:10 LD, 20°C with rhizoidal portion (arrow); B. Thallus consisted of long decumbent and short erect filaments; C. Datail of rhizoidal portion; D. Irregular branches (arrow); E. Erect filament arose perpendicularly from middle portion of decumbent cell; F, G. Tetrasporangia on terminal of short branches; H. Detail of discoidal plastid without pyrenoid. Nemaliophycidae: Acrochaetiales: Acrochaetiaceae 35

Rhodochorton intermedium (Kjellman) Kjellman 1883: 231. pl. 15. f. 8. Rhodochorton islandicum Rosenvinge 1900: 75. f. 1-4. Rhodochorton parasiticum Batters 1896: 389. Rhodochorton rothii (Turton) Nägeli 1861: 356. Rhodochorton tenue Kylin 1925: 44. Thamnidium intermedium Kjellman 1875: 28. f. 10. Thamnidium rothii (Turton) Thuret in Le Jolis 1863: 111. Trentepohlia purpurea (Lightfoot) C. Agardh 1824: 36. Trentepohlia rothii (Turton) Harvey 1836: 218.

Thalli epilithic, caespitose, composed of decumbent and erect filaments, 700-900 μm high. Decumbent filaments having obtuse apices and rhizoidal portion, growing long simple filament at first, branched sparsely and issusing short erect filaments almost right angle from middle portion of decumbent cell, coherently entangled at later, forming massive basal plates. Rhizoidal cells irregular in shape, elongated and contorted, contained few plastids, rarely developed in culture condition. Cells of erect filaments cylindrical, 9-11 μm wide, 30-55 μm long. Erect filaments branched subdichotomously to alternately, usually producing tetrasporangia on terminal cells of branches. Cells of erect filament cylindrical, 10-12 μm wide, 15-25 μm long, not constricted at junction. Hairs absent. Plastids several, discoid without pyrenoids. Tetrasporangia terminal on branchlet or erect filaments, obovoid, cruciately divided, 17-18 μm wide, 28-30 μm long in mature.

TYPE: Ruined Abbey, Island of Iona, Scotland(?). SEASONALITY: Winter to spring. DISTRIBUTION: Cosmoploitan. KOREA: Jeju Island. SPECIMEN EXAMINED: West-4187 (culture strains: isolated from supralittoral zone at Seongsanpo, Jeju: 28. ix.2001 by Y.P. Lee, see West et al. 2006). REMARKS: Rhodochorton purpureum has been studied intensively with culture by Knaggs (1965, 1966, 1967, 1968), by West (1969, 1970, 1972), and by Ohta and Kurogi (1979). Althought, Lee (1985) reported the sexual and asexual reproduction of the plants in Hok- kaido population, but Korean materials in wild have been reported without reproductive structures from only Jeju Island (Lee 1987). Accoding to Lee, the Kor- ean materials grow on rocks facing north and shelter- ed of crevice with dim light in the supralittoral zone. We can observe this species based on the culture strain (J.A. West; no 4187), which isolated from Jeju Island by Lee (West et al. 2006). The cultured plants are somewhat different in branching mode, but are almost identical with descriptions of Lee (1987) in plastid morphology and cell dimension except the reproduction. The Jeju strain also produced tetrasporangia in our Fig. 24. Distribution of Rhodochorton pur- culture conditions as like the result of West et al.(2006). pureum. 36 Algal Flora of Korea·Nemalian Red Algae

Order Colaconematales Harper et Saunders 2002: 471. Hong-da-bal-sol-mok (홍다발솔목: 신칭)

Thalli filamentous, monosiphonous, simple or branched. Plastid variable, one to several per cells, parietal lobed to irregular but never stellate with or without pyrenoid. Asexual reproduction primarily by monosporangia or not observed. Sexual life histories occasionally observed, when known its alternation of triphasic isomorphic generations.

REMARKS: Harper and Saunders (2002) established this order mainly by the DNA sequencing phylogeny. In phylogenetic tree based on a SSU-LSU r-DNA combined data set, this order is clearly separated as a sister of Acrochaetiales-Palmariales complex (Fig. 1). However, it is very difficult to distinguish this order from Acrochaetiales based on morphological characters.

Family Colaconemataceae Harper et Saunders 2002: 471. Hong-da-bal-sol-gwa (홍다발솔과: 신칭)

Diagnosis same as the order. Type genus: Colaconema Batters 1896: 8.

GENERA AND SPECIES: 1 genus (ca. 100 species). DISTRIBUTION: Worldwide in marine to freshwater. KEY REFERENCE: Harper and Saunders (2002). REMARKS: Usually the members of this family have a larger and elongate cell type than those of Acrochaetiales. At present, only one genus is known in this family. Harper and Saunders (2002) pointed out that the current Colaconema generic circumscription is not a final solution to the pro- blems of generic delineation within the Colaconematales, but it is a foundation for further research to build a revised and comprehensive classification for these organisms. The newly emended genus Audouinella (Acrochaetiaceae) was restricted to the members of occurring in freshwater (Harper and Saunders 2002). Thus, the reported species of the marine Audouinella in Korea (Y. Lee 1987) agreed well with the characteristics of this family.

Genus Colaconema Batters enmend. Harper et Saunders 2002: 473. Hong-da-bal-sol-sok (홍다발솔속: 신칭)

Marine acrochaetioid algae. Cells with one to several parietal plastids of varying shaped (lobed to spiral to irregular of laminate) with or without pyrenoids. Asexual reproduction primarily by monosporangia. Sexual reproduction with triphasic life history occasionally observed. Type species: Colaconema bonnemaisoniae Batters 1896: 8. Nemaliophycidae: Colaconematales: Colaconemataceae 37

SPECIES uncountable at present (ca. 12 species in Korea). DISTRIBUTION: Cosmopolitan, widely distributed in temperate and tropical waters. KEY REFERENCE: Harper and Saunders (2002), Lee (1987). REMARKS: The genus Colaconema was established by Batters (1896a) for acrochaetioid with prostrate, irregular filaments, and monosporangia formed on terminal or on intercalary cup-like cells. Accord- ing to the original description, the cup-like monosporangial-bearing cells is regarded a diagnostic character, however, several species of Colaconema do not have it (see Harper and Saunders 2002).

Key to the species of genus Colaconema

1. Thallus epiphytic to epi-endophytic or epizoic·······································································2 - Thallus endozoic in fibers of sponge········································································C. infestans 2. Two to more pyrenoids per cell of erect filaments, growing exclusively on Codium ················3 - Mostly single pyrenoid per cell of erect filaments···································································4 3. Usually one or two pyrenoids per cell of erect filaments················································C. codii - Usually 4-6 pyrenoids per cell of erect filaments ·······················································C. codicola 4. Lateral branches arranged distichously on one side ···················································C. compta - Lateral branches arranged alternately or irregularly·······························································5 5. Monosporangia seem to be bisporangia by swelling of the stack cell ·························C. bispora - Monosporangia sessile or pedicellate without swelling of the stack cell ·································6 6. Lateral branches terminate with long hyaline hairs ·········································C. hyalosiphoniae - Hyaline hairs absent···············································································································7 7. Basal cells rounded, single-celled at least in early stage···········································C. dictyotae - Basal system filamentous consisting of contorted cylindrical cells··········································8 8. Monosporangia densely clustered on short lateral branches······································C. daviesii - Monosporangia seriated on pedicels of erect axis or branches ················································9 9. Thallus much branched and the branches reach almost same height ·························C. thuretii - Thallus less branched and produced serial short laterals of monosporangia ·························10 10. Monosporangia arranged exclusively on one side ··················································C. pacificum - Monosporangia arranged alternately to secundly ······················································C. elegans

12. Colaconema bispora (Børgesen) Hwang et Kim, comb. nov.(Figs. 25, 26) Du-po-ja-hong-da-bal-sol (두포자홍다발솔: 신칭)

BASIONYM: Chantransia bispora Børgesen 1910: 178. f. 1. SYNONYM: Acrochaetium bisporum (Børgesen) Børgesen 1915: 43. f. 41. Audouinella bispora (Børgesen) Garbary 1979: 490. Schneider 1983: 5. f. 1c-f.

Thalli epiphytic, caespitose, composed of basal prostrate and erect filaments, up to 500-700 μm high. Basal filaments creeping horizontally, branched irregular interval, forming loosely entangled mass, bright red colour. Cells of endophytic filaments cylindrical, slightly contorted, including laminate plastid with a cental pyrenoid, 6-8 μm wide, 15-25 μm long. Erect filaments arising at right angles from basal prostrate filaments, issuing alternate branches in lower to middle portion. Branches usually simple, tapering toward apices and usually terminating with terminal monosporangia without hairs. Cells of erect filaments cylindrical, slightly constricted at cross wall, 6-8 μm 38 Algal Flora of Korea·Nemalian Red Algae

ABC

DE F

Fig. 25. Colaconema bispora. A-C. Cultured plants isolated from epiphytes of Eisenia bicyclis (Ulleung Island); D-F. Cultured plants isolated from Phyllospadix sp. (Daeksan harbor, Samcheok).

wide and 15-20 μm long. Plastids of erect filaments parietal laminate with one pyrenoid. Monospo- rangia born secundly on branches or on terminal portion of branches, usually solitary on one-celled swollen stalks which seem to be bisporangia, ellipsoid, 8-10 μm wide, 13-18 μm long. Regeneration occurring in empty monosporangia. Tetrasporangia or sexual reproductive structures unknown. Nemaliophycidae: Colaconematales: Colaconemataceae 39

TYPE: St. Thomas, Virgin Islands (C. see Schneider 1983). SEASONALITY: Summer. DISTRIBUTION: West Indian, North Carolina, Korea. KOREA: East coast. SPECIMEN EXAMINED: KN1102 (Jukdo, Ulleung Island: 29.vii.2009, epiphytic on blade of Eisenia bicyclis), KN1103 (Deoksan harbor, Samcheok: 16.vii.2009, epiphytic on Phyllospadix sp.). REMARKS: This is the first report of this taxon from outside of the North Atlantic Ocean (West Indies and North Carolina). This species is clearly distinguished by the peculiar monosporangia which look like bisporangia by the swollen stalk cell. The monosporangial feature of Korean specimens is almost same with the descriptions and illustrations provided by Børgesen (1910, f. 1) and Schneider (1983, f. 1c-f). Although, Korean specimens are not congruent with the earlier descriptions in branching mode and some other charc- teristics, but we considered the biosporangious monosporangial features of Korean plants is a sufficient evidence for identifying this species. Fig. 26. Distribution of Colaconema bispora.

13. Colaconema codicola (Børgesen) Stegenga, Bolton et Anderson 1997: 248 (Figs. 27-29). Cheong-gak-hong-da-bal-sol (청각홍다발솔: 개칭)

Stegenga, Bolton and Anderson 1997: 248. pl. 76. f. 1, 2.

BASIONYM: Acrochaetium codicola Børgesen 1927: 33. f. 18-20. SYNONYM: Audouinella codicola (Børgesen) Garbary 1979: 490. Y. Lee 1987: 7. f. 1. Lee and Kang 2001: 249. Rhodochorton codicola (Børgesen) Nakamura 1944: 113. Kang 1962: 75. Lee and Lee 1974: 49. f. 8. Rhodochorton rhizoideum sensu Lee and Lee 1974: 49. f. 9.

Thallus epi-endophytic on Codium sp., caespitose, composed of basal and erect filaments, up to 3 mm high, purple red colour. Basal filaments endophytic, developing downwards as well as horizontally between utricles of host, loosely entangled. Cells of endophytic filaments cylindrical, slightly contorted, including 2-3 irregularly robed or patch-like plastids with pyrenoids, 15-22 μm wide, 40-80 μm long. Erect filaments arising at right angles from horizontally developed endophytic basal filaments, issuing secund or alternate branches at various intervals. Branches tapering toward apices and sometimes terminating with thin, feeble, hair-like prolongations. Cells of erect filaments cylindrical, not constricted at cross wall, 10-15 μm wide and 50-100 μm long. Plastids in 40 Algal Flora of Korea·Nemalian Red Algae

ABC D

EFG H

Fig. 27. Colaconema codicola. A. Detached thallus from Codium fragile; B. Endophytic basal filament and lower portion of erect filaments; C. Irregular shaped plastids in cells of basal filament; D. Branches and monosporangia on erect filaments; E, F. Detail of pyrenoids in cells of erect filaments; G, H. Detail of monosporangia in room cultures.

cells of mature erect filaments containing 2- to 4- (-7) lobed parietal forms, each with a central pyrenoid. Monosporangia born secundly on adaxial side of branches or on short branches, sessile or on one-celled stalks, solitary or in pairs, ellipsoid, 15-20 μm wide, 15-25 μm long. Regeneration occurring in empty monosporangia. Tetrasporangia or sexual reproductive structures unknown. Nemaliophycidae: Colaconematales: Colaconemataceae 41

ABC

DEFG

Fig. 28. Colaconema codicola. A-G. Room cultures; A. Habit of branching and rhizoidal cell (arrow); B, C. Monosporangia on adaxial side of laterals (arrow head) or short laterals (arrow); D. Mono- sporangia sessile or on one-celled pedicel; E. Regeneration of monosporangia (arrow); F, G. Detail of plastids and pyrenoids.

TYPE: Canary Islands. C (Botanical Museum and Herbarium, Copenhagen, Denmark). SEASONALITY: Observed June to October. DISTRIBUTION: Cosmopolitan. KOREA: Around all coasts of Korea. 42 Algal Flora of Korea·Nemalian Red Algae

SPECIMEN EXAMINED: KN1104 (Jukdo, Ulleung Island: 29.vii.2009). Partly endophytic in Codium. REMARKS: Colaconema codicola is partly endophytic in the lower part of thallus of Codium fragile and was reported earlier than other acrochaetioid in Korea since Kang (1962). This species is easily recognized by the relatively larger thalli as one of the epi-endo- phytes on Codium fragile. This species is very similar with Colaconema codii (=Callithamnion codii Crouan et Crouan), so Levring (1974) suggested that C. codicola and C. codii are conspecific. However, both species are distinguished by pyrenoid (see C. codii in this work). It is regarded that the peculiar type of plastid of this species, which contains 2 to 4 (-7) conspicuous pyrenoids per mature cell of erect filaments, is one of the main character to separate this species from other acrochaetioids. The conspicuous multi-pyrenoids per cell of this species are well illustrated by Stegenga et al. (1997, pl. 76, f. 1, 2), which is same with our specimens. Lee and Lee (1974) distinguished C. codicola (as name Rhodochorton codicola) and Rhodochorton rhizoideum based on the characteristic that the basal portions of the former species are hyaline, whereas they are Fig. 29. Distribution of Colaconema codi- lightly pigmented in the latter. However, the degree cola. of pigments in cell of basal filament is affected by environmental conditions and also we found no reliable morphological difference to separate both Korean specimens comparing to illustrations given by Lee and Lee (1974, f. 8, 9). This species seems to be a rather common epi-endophyte upon various acrochaetioidal species of Codium in Korea.

14. Colaconema codii (Crouan et Crouan) Hwang et Kim, comb. nov. (Figs. 30, 31) Keun-hong-da-bal-sol (큰홍다발솔: 개칭)

BASIONYM: Callithamnion codii Crouan and Crouan 1860: 368. SYNONYM: Acrochaetium codii (Crouan and Crouan) Bornet 1904: xx. Audouinella codii (Crouan and Crouan) Garbary 1979: 490. Lee 1987: 8. f. 2. Chantransia codii (Crouan and Crouan) Levring 1937: 89. Rhodochorton codii (Crouan and Crouan) Nakamura 1944: 112. Rhodothamniella codii (Crouan and Crouan) Feldmann 1981: 431.

Thalli epi-endophytic on Codium sp., caespitose, composed of endophytic basal filaments and erect filaments, 2-10 mm high, bright red colour. Endophytic basal filaments running downwards and horizontally between utricles of host, usually forming a cluster. Cells of endophytic filaments contorted, cylindrical, including one or two parietal laminate or patch-like plastids, 15-25 μm wide, Nemaliophycidae: Colaconematales: Colaconemataceae 43

A B

CDEF

Fig. 30. Colaconema codii. A. Detached thallus from Codium fragile; B. Monosporangia and erect filamentous cells contain 1-2 pyrenoids (arrow); C. Detail of erect and prostrate filaments; D. Detail of monosporangia; E, F. Detail of plastids and one or two pyrenoids (arrow).

55-85 μm long. Erect filaments arisng in a cluster, somewhat rigid, issuing branches subdichotomously. Cells of erect filaments cylindrical, not constricted at septum, 15-20 μm wide, 35-50 μm long. Plastids of erect filaments parietal laminate with one or two pyrenoids. Monosporangia born laterally, solitary on single-celled stalks, oblong to ellipsoid, 13-18 μm wide, 20-30 μm long. Tetra- sporangia or sexual reproductive structures unknown. 44 Algal Flora of Korea·Nemalian Red Algae

TYPE: Délec, Brest, France (see Hamel 1927: 31). SEASONALITY: Summer. DISTRIBUTION: Worldwide. KOREA: All around the Korean coast. SPECIMEN EXAMINED: KN1106 (Sinji islets, Wando: 18.viii.2009, epi-endophytic on Codium fragile). REMARKS: Colaconema codii has been confused to identify based on literatures because there is no de- pendable descriptions or illustrations for this species at present. After short original discription given by Crouan and Crouan (1960) as “Sur le Codium elongatum, sur lequel il forme des gazons serrés” additional descriptions were provided by Bornet (1904), and later by Hamel (1927) who given illustrations of this species at first. Thus, the description and illusration given by Hamel (1927) were mainly applied for identification of this species at present. However, the plastidal features of this species were not described by the French- ish phycologists. The morphological boundary of this species was configured in detail by Børgesen (1927), who established a new species C. codicola based on eip-endo- Fig. 31. Distribution of Colaconema codii. phyte on Codium sp.. According to Børgesen (1927), C. codii is distinguished from C. codicola by the following features; the uppermost ends of the branches are very attenuated as subhyline, the plastids (two or more in each cell) contain only 1-2 pyrenoids, the sporangia are always pedicellate, and the ramification as a rule alternate (rarely opposite). Thus, the features of 1-2 pyrenoids per cells and pedicellate monosporangia were applied for identification of this species in this work. How- ever, Garbary (1987) mentioned that C. codii (as Audouinella codii) is a part of a species complex that includes Audouinella botryocarpa, Audouinella phacelorhiza, Audouinella rhizophora; some or all of these may be conspecific. Further morphological examination, culture observations and molecular analyses may prove the related speceis to be conspecific with Audouinella codii. The overall morphological characteristics of this species, especially the parietal laminate plastid with a conspicuous pyrenoid and the monosporangial feature, agree well with the characters of Colaconema species. Thus, we proposed a new combination for this species, as Colaconema codii (Crouan and Crouan) Hwang et Kim, comb. nov., in this work.

15. Colaconema compta (Børgesen) Hwang et Kim, comb. nov.(Figs. 32, 33) Pyeon-saeng-hong-da-bal-sol (편생홍다발솔, 신칭)

BASIONYM: Acrochaetium comptum Børgesen 1915: 46. f. 44-46. SYNONYM: Audouinella compta (Børgesen) Garbary 1979: 490. Chantransia compta (Børgesen) De Toni 1924: 40. Nemaliophycidae: Colaconematales: Colaconemataceae 45

ABC

DEF

Fig. 32. Colaconema compta. A. Habit of epiphytic thallus on blade of Ecklonia cava; B. Detached single erect filament; C, D. Secundly arranged branches and monosporangia; E. Detail of pedicellate monosporangia; F. Detail of monosporangia and plastids.

Thalli epiphytic on Ecklonia cava, bright to purple red, caespitose, composed of prostrate and erect filaments, 1.0-1.2 mm high. Prostrate filaments partly penetrate into host epidermis, branched irregularly and entangled, forming a cluster. Cells of basal filaments contorted, cylindrical, including one parietal laminate plastids, 5-6 μm wide, 3-7 μm long. Erect filaments straight, simple below, 46 Algal Flora of Korea·Nemalian Red Algae but much branched above subdichotomously, slightly tapered to apex but not developing pseudohairs. Lat- eral branches distichious, arranged secundly, reached almost same height. Cells of erect filaments cylindrical, not constricted at septum, 6-8 μm wide, 15-20 μm long. Plastids parietal laminate, with one conspicuous pyrenoid. Monosporangia born series on adaxial side of laterals or rarely clustered on 2-3 celled short laterals, solitary on single- or rarely two-celled stalks, oblong to ellipsoid, 6-8 μm wide, 12-14 μm long. Tetrasporangia or sexual reproductive structures unknown.

TYPE: White Bay, St. Croix, Virgin Island (C, Botani- cal Museum and Herbarium, Copenhagen, Denmark; Epiphytic Liagora pinnata). SEASONALITY: Summer. DISTRIBUTION: Virgin Island, Korea (this work). KOREA: Jeju Islands. SPECIMEN EXAMINED: KN1108 (Moon islet, Jeju Island: 16.viii.2009, epiphytic on Ecklonia cava). REMARKS: Colaconema compta has been rarely reported in Virgin Island (Børgesen 1915; Taylor 1960) as an Fig. 33. Distribution of Colaconema comp- epiphyte on Liagora pinnata. Thus, we are in two minds ta. about determination of this species based on Korean materials. However, the secundly-seriately arranged lateral branches and the seriately-singly born monosporangia on one-celled stalks at adaxial side of laterals are well agreed between both specimens. According to Børgesen (1915), this species was characterized by the fact that the germinating spore is divided by a transverse wall into two cells from the upper of which an ererct filament grows upwardly, while from the lower endophytic filaments growing downwards are produced. Although we could not detect the spore germination mode in our Korean specimens, but the germination pattern is regarded as a popular manner in Colaconema codicola and others. Thus the diagnostic characters of this species are considered the secundly-seriately arranged branches and seriately-adaxially born monosporangia. The secundly-seriately arranged branches and seriately- adaxially born monosporangia are also found in Acrochaetium seriatum Børgesen (1915, Fig. 32). However, Acrochaetium seriatum distinguished by the sessile type of monosporangia. Therefore, we provisionally identified the Korean specimens to this species based on the above mentioned diagnostic characters. The overall morphological characteristics of this species, especially the parietal laminate plastid with a conspicuous pyrenoid and the monosporangial feature, agree well with the characters of Colaconema species. Thus, we proposed a new combination for this species, as Colaconema compta (Børgesen) Hwang et Kim, comb. nov., in this work. Nemaliophycidae: Colaconematales: Colaconemataceae 47

16. Colaconema daviesii (Dillwyn) Stegenga 1985: 317(Figs. 34-37). Song-i-hong-da-bal-sol (송이홍다발솔: 개칭)

Stegenga 1985: 317, 320. f. 20. Harper and Saunders 2002: 473.

BASIONYM: Conferva daviesii Dillwyn 1809: 73. suppl. F. SYNONYM: Acrochaetium daviesii (Dillwyn) Nägeli 1861: 405. Audouinella daviesii (Dillwyn) Woelkerling 1971: 28. Lee 1987: 10. f. 3. Callithamnion daviesii (Dillwyn) Lyngbye 1819: 129. tab. 41B Ceramium daviesii (Dillwyn) C. Agardh 1824: 132. Rhodochorton daviesii (Dillwyn) Drew 1928: 172. Lee and Lee 1974: 41. f. 4. Trentepohlia daviesii (Dillwyn) Harvey in Mackay 1836: 219.

Thalli epiphytic on various algae or epilithic, endophytic, forming a spherical to hemispherical fringe, composed of basal creeping and erect filaments, up to 1.5 mm high, purple red colour. Basal filaments creeping, branching, entangled, forming a rather massive basal structure. Cells of basal filaments contorted cylindrical with parietal laminate rhodoplasts, 7-8 μm wide and 10-15 μm long. Erect filaments more or less rigid, issuing branches at various intervals, branches reaching the upper level of main axes. Cells of erect filaments cylindrical, 8-12 μm wide, 10-35 μm long. Plastids of erect filaments parietal laminate with a pyrenoid. Monosporangia born in clusters on short branchlets, solitary or pairs on stalks of a single obtrian- gular cell, ovoid to ellipsoid, 8-11 μm wide, 13-19 μm long. Tetrasporangia or sexual reproductive structures not observed.

TYPE: Bantry Bay, Ireland (NMW: Department of Botany, National Museum of Wales, Great Britain). SEASONALITY: Rarely observed summer season (June to September). DISTRIBUTION: Worldwide cosmopolitan. KOREA: All around coasts of Korea. SPECIMEN EXAMINED: KN1010 (Gangneung: 5.vi.2009), KN1011 (Tongumi, Ulleung Island: 28-29.vii.2009), KN1013 (Incheon: 9.i.2009). REMARKS: Colaconema daviesii is one of the common- er species of Colaconema in Korea and reported all over coast of Korean (see Lee and Kang 2001). Futhermore, this species occurs on diverse hosts such as hydroids, various algae, ghost net or fishing wire. The main character for identification of this species is the peculiar type of monosporangia which formed as lateral groups of monosporangia on branched stalks. Korean specimens in our observations agree well with the pre- vious descriptions in worldwide (Kylin 1909, f. 27; Fig. 34. Distribution of Colaconema davi- Rosenvinge 1909, f. 34A-F, excluding f. C; Kornmann esii. 48 Algal Flora of Korea·Nemalian Red Algae

AB C

DEF

Fig. 35. Colaconema daviesii. A-F. Cultured plants isolated from Gangneung; A, B. Sparsely branches with fascicles of monosporangia; C. Detail of parietal laminate plastid with single pyrenoid; D-F. Detail of the position and morphology of monosporangia. Nemaliophycidae: Colaconematales: Colaconemataceae 49

AB C

DE F

Fig. 36. Colaconema daviesii. A-F. Cultured plants isolated from Jumunjin, Gangneung; A. Habit of thallus; B, C. Subdichotomous laterals with short sporangial branchlets; D-F. Detail of the position and morphology of monosporangia. 50 Algal Flora of Korea·Nemalian Red Algae

ABC

DE F G

Fig. 37. Colaconema daviesii (?) A-G. Field plants isolated from Incheon harbor; A. Habit of thallus detached from wasted fishing wire; B, C. Subdichotomous branches and secundly arranged monosporangia; D-F. Detail of the position and morphology of monosporangia; G. Detail of plastid. Nemaliophycidae: Colaconematales: Colaconemataceae 51 and Sahling 1977, f. 97A-C; Garbary et al. 1982, f. 6a-j; Woelkerling and Womersley 1994, f. 9A-H; Stegenga et al. 1997, pl. 78. f. 2). There are some variations of Korean plants in cell dimensions and monosporangial features as shown in Fig. 35-37. However, all specimens in our observations had the peculiar type of monosporangia to determine this species.

17. Colaconema dictyotae (Collins) Hwang et Kim, comb. nov.(Figs. 38-40) Ga-neun-hong-da-bal-sol (가는홍다발솔: 개칭)

BASIONYM: Acrochaetium dictyotae Collins 1906: 193. SYNONYM: Audouinella dictyotae (Collins) Woelkerling 1971: 38. f. 13, 25. Lee 1987: 12. f. 4. Woel- kerling and Womersley 1994: 53. f. 10A-H. Chantransia dictyotae (Collins) Collins 1911: 186. Rhodochorton dictyotae (Collins) Drew 1928: 190. pl. 47. f. 79, 80. Rhodochorton magnificum sensu Kang 1962: 75. Lee and Lee 1974: 72. f. 5.

Thalli usually epi-endophytic on Dictyota or Pachydictyon sp, caespitose, composed of basal and erect filaments, to 3 mm high, bright red colour. Basal filaments irregular in size and shape, penetrating between cortical layer and medullary cells of the host, single cell state as larger globular cell at least earlier, later issuing downwardly to horizonitally endo-creeping filaments from the basal cell. Cells of basal filaments various in shape with round margin, with a parietal laminate plastids, extremely constricted at the septum, 10-25 μm wide, 20-45 μm long, issuing erect filaments at right angles. Erect filaments much branched subdichotomously to alternately throughout, tapering slightly toward apices. Cells of erect filaments cylindrical, not constricted at the septum, 8-11 μm wide, 15-35 μm long. Plastids of the cells of erect filaments parietal laminate with one or rarely two conspicuous pyrenoids. Monosporangia singly on 1- 4 celled laterals or short clusters, usually on the proxi- mal cells of the laterals, ovoid to oblong, 15-18 μm wide, 18-22 μm long. Tetrasporangia or sexual reproductive structures not observed in this work.

TYPE: La Jolla, California, USA (Farlow Library and Herbarium of Cryptogmic Botany, Harvard Univer- sity, Cambridge, Massachustts, USA). SEASONALITY: Rarely observed summer season (June to September). Fig. 38. Distribution of Colaconema dic- DISTRIBUTION: California, Australia and Korea. tyotae. 52 Algal Flora of Korea·Nemalian Red Algae

ABC

DEF

Fig. 39. Colaconema dictyotae. A, B. Habit of epiphytic thallus on Dictyota sp. (Moon islet, Jeju Island); C. Detail of single globular basal cell (arrow); D. Subdichotomously to alternately arranged branches; E. Single monosporangium on pedicel; F. Detail of laminate plastid with a conspicuous pyrenoid. Nemaliophycidae: Colaconematales: Colaconemataceae 53

ABC

D E F

Fig. 40. Colaconema dictyotae. A. Habit of epiphytic thallus on Dictyota coriacea (Anin, Gangneung); B. Detail of single-celled basal cell (arrows); C. Subdichotomously to alternately arranged branches; D, E. Monosporangium clusters on short laterals; F. Detail of laminate plastid with a conspicuous pyrenoid. 54 Algal Flora of Korea·Nemalian Red Algae

KOREA: All around coast of Korea. SPECIMEN EXAMINED: KN1012 (Moon islet, Jeju Island: 5.vi.2009), KN1014 (Tongumi, Hakpo, Ulleung Island: 28-29.vii.2009), KN1016 (Anin, Gangneung: 23.viii.2009). Epi-or endophytic on/in Dictyota or Dictyopteris. REMARKS: Colaconema dictyotae was usually growing on the thallus of Dictyota or of the related genera Spatoglosum, Pachydictyon, Dictyopteris. This species was reported as Rhodochorton magnificum Drew by Kang (1962) and later by Lee and Lee (1974) in Korea, however, Lee (1987) noticed an iden- tificational errors it. Furthermore, Lee (1987) found the tetrasporic and sexual gametophytic plants of this species from Korean materials. However, we failed to collect the sexual plants in this work. Korean plants agree well with the descriptions and illustrations given by Drew (1928, pl. 47, f. 79, 80). The tetrasporangial plants of C. dictyotae share several characters with C. daviesii such as the laminate parietal plastid with a conspicuous pyrenoid, epi-endophytic habits, and mode of monosporangium formation. However, both species are clearly distinguished by basal endophytic filaments. The overall morphological characteristics of this species, especially the laminate plastid with a conspicuous pyrenoid and the monosporangial features, is acceptable to the member of genus Colaconema. Therefore, we proposed a new combination for this species, as Colaconema dictyotae (Børgesen) Hwang et Kim, comb. nov., in this work.

18. Colaconema elegans (Drew) Hwang et Kim, comb. nov.(Figs. 41, 42) Go-bi-hong-da-bal-sol (고비홍다발솔: 개칭)

BASIONYM: Rhodochorton elegans Drew 1928: 183. SYNONYM: Acrochaetium elegans (Drew) Papenfuss 1945: 314. Audouinella elegans (Drew) Lee 1987: 15. f. 6A-E.

Thalli epi-endophytic usually on Sargassum coreanum, caespitose, composed of endophytic prostrate and erect filaments, 2-3 mm high, purple fire-red colour. Monospores germinating in unipolar mode, original spore persistent when germination. Endophytic filaments penetrating downwards into cortical layer of host, branching up- and downwards. Cells of endophytic filaments contortedly cylindrical with a single parietal laminate plastid with a single pyrenoid, 10- 14 μm wide, 22-36 μm long. Erect filaments developing upwards from endophytic filaments, simple in lower portion but issuing longer or shorter branches from the upper portion, not tapered to apex or not formed pseudo-hairs. Cells of erect filaments cylin- Fig. 41. Distribution of Colaconema ele- drical, not constricted at septum, 8-10 μm wide, 13- gans. Nemaliophycidae: Colaconematales: Colaconemataceae 55

ABC

DEF

Fig. 42. Colaconema elegans. A-F. Room cultured plants (isolated from Udo islet, Jeju Island); A, B. Branches and monosporangia formed in upper portion; C. Detail of upper portion of thallus; D, E. Detail of monosporangia; F. Detail of plastid.

30 μm long. Plastid of erect filaments parietal laminate with a pyrenoid. Monosporangia born solitary or in pairs on one-or two-celled stalks, in secund arrangement on adaxial side of branches on erect filaments, ellipsoid to ovoid, 10-12 μm wide, 16-20 μm long. Tetrasporangia or sexual reproductive structures unknown. 56 Algal Flora of Korea·Nemalian Red Algae

TYPE: La Jolla, California, USA (UC 294555, Herbarium of the UC Berkeley, California). SEASONALITY: Rarely observed in summer season (June to September). DISTRIBUTION: North Pacific Ocean, Korea. KOREA: Jeju Island. SPECIMEN EXAMINED: KN1018 (Udo islet, Jeju Island: 2.vi.2009, epi-endophytic on Sargassum coreanum). REMARKS: This species was reported from Korea as named Audouinella elegans (Drew) by Lee (1987). Korean plants of C. elegans are epi-endophytic in cortical layer of the lower part of the stipe of Sargassum coreanum. Drew (1928) described Californian plants parasitic in the stipe of Eisenia arborea. However, it is difficult for biological determination whether the plants at hand are parasitic or merely endophytic, although the infected cortical layer is detached rather easily from the medullary layer of the host. Woelkerling (1971) regarded C. elegans to be closely related with C. daviesii. Abbott and Hollenberg (1976) synonymized the former with C. pacificum (Kylin) Woelkerling. The morphological characteristics of this species, especially the laminate plastid with a conspicuous pyrenoid and the monosporangial features, is acceptable to the member of genus Colaconema. Therefore, we proposed a new combination for this species, as Colaconema elegans (Børgesen) Hwang et Kim, comb. nov., in this work.

19. Colaconema hyalosiphoniae (Nakamura) Hwang et Kim, comb. nov. (Figs. 43, 44) Ga-si-hong-da-bal-sol (가시홍다발솔: 개칭)

BASIONYM: Rhodochorton hyalosiphoniae Nakamura 1941: 287. f. 14-16. SYNONYM: Acrochaetium hyalosiphoniae (Nakamura) Papenfuss 1945: 314. Audouinella hyalosiphoniae (Nakamura) Garbary 1979: 490. Rhodochorton hyalosiphoniae Nakamura senusu Lee and Lee 1974: 45. f. 7.

Thalli epiphytic, caespitose, heterotrichous compos- ing of prostrate and erect filaments, 1-1.5 mm high, bright red colour. Basal prostrate filaments comprised of branched filaments, often forming compact con- flent layer on the substratum. Erect filaments subdichotomously to secundly branched, often producing prominent primary axes from which numerous short branchlets formed. Short branchlets 4-7 celled long, tapered gradually, terminated with long hyaline pseudo-hair, producing monosporangia secundly. Cells of erect filaments cylindrical, 10-14 μm wide, 14-27 μm long. Plastids single per cells, parietal lam- Fig. 43. Distribution of Colaconema hyalo- inate with a conspicuous pyrenoid. Hayline pseudo- siphoniae. Nemaliophycidae: Colaconematales: Colaconemataceae 57

ABCD

EF G H

Fig. 44. Colaconema hyalosiphoniae. A, B. Detached thalli from Sargassum fusiforme (Anmok, Gang- neung); C. Habit of erect filaments and basal portion; D. Erect filaments with short monosporangial branchlets; E. Branching mode; F. Detail of hyaline hairs on terminal of branches and monosporangia; G, H. Detail of sigle laminate plastid with a conspicuous pyrenoid. 58 Algal Flora of Korea·Nemalian Red Algae hairs common, formed on terminal of branches or short branchlets. Monosporangia produced secundly on short branchlets, forming clusters, usually arose on one- or two-celled stalks, oblong in shaped, 8-10 μm wide, 10-15 μm long. Tetrasporangia or sexual reproductive organs not observed.

TYPE: Murran, Hokkaido, Japan. SEASONALITY: Rarely observed summer season (June to September). DISTRIBUTION: West Indian Sea, Florida and Gulf of Mexico, Korea. KOREA: East coast. SPECIMEN EXAMINED: KN3406 (Anmak, Gangneung: 13.viii.2010, epiphytic on Sargassum sp.). REMARKS: This species was reported as named Rhodochorton hyalosiphoniae Nakamura in Korea by Lee and Lee (1974), however, it was regarded to be conspecific with Colaconema daviesii (Yoshida 1998; Lee and Kang 2001). Although, C. daviesii has been described broadly including hairless to haired types (eg. Lee 1987, f. 3A-E; Lee and Yoshida 1997, f. 1A-F), most of descriptions induding protologue (Dillwyn 1809) excluded the haired type. Furthermore, this species is clearly distinguished by rbcL sequencing data from C. daviesii (unpublished data). Thus, we separated this species as a distinct species in this work. The morphological characteristics of this species, especially the laminate plastid with a conspicuous pyrenoid and the monosporangial features, is acceptable to the member of genus Colaconema. Therefore, we proposed a new combination for this species, as Colaconema hyalosiphoniae (Nakamura) Hwang et Kim, comb. nov., in this work.

20. Colaconema infestans (Howe et Hoyt) Woelkerling 1973: 89 (Figs. 45, 46). Hae-myeon-son-hong-da-bal-sol (해면손홍다발솔: 개칭)

Woelkerling 1973a: 89. f. 5, 6.

BASIONYM: Acrochaetium infestans Howe et Hoyt 1916: 116. SYNONYM: Audouinella infestans (Howe et Hoyt) Dixon in Parke and Dixon 1976: 590. Dixon and Irvine 1977: 99. f. 28. Lee 1987: 17. f. 7. Chantransia infestans (Howe et Hoyt) De Toni 1924: 64. Kylinia infestans (Howe et Hoyt) Papenfuss 1947: 438. Rhodochorton infestans (Howe et Hoyt) Drew 1928: 151. Nakamura 1944: 118. f. 13.

Thalli endozoic usually on the fiber of Callyspongia sp., composed of endozoic creeping and erect filaments, purple fire-red colour. Endozoic filaments tightly adhering and extending tortuously or parallelly along the fiber, irregularly branched, forming compacted cortication on the surface of inner fiber of sponge. Cells of creeping filaments cylindrical to globose or polygonal, 8-10 μm wide, 15-35 μm long. Erect filaments protruding beyond the host surface here and there, branching secundly, shorter less than 60-80 μm high, slightly or not tapered toward apex. Cells of erect filaments cylindrical, 6-7 μm wide, 6-12 μm long. Plastids single per cell, parietal laminate with a pyrenoid. Hairs not formed. Monosporangia arising laterally to seriately on erect filaments, secundly arranged, sessile or stalked, ovoid, 6-7 μm wide, 8-9 μm long. Tetrasporangia not observed. Nemaliophycidae: Colaconematales: Colaconemataceae 59

AB C

DEFG

Fig. 45. Colaconema infestans. A. Habits of endozoic thalli on inner fiber of Callyspongia elegans; B. Detail of endozoic creeping and short erect filaments; C. Seriately-laterally developed monosporangia on erect filaments; D-G. Cultured plants (isolated from Sagye, Jeju); D. Filamentous thalli grow free of host in laboratory culture; E. Detail of mature lower filament; F. Series of lateral monosporangia on erect filament; G. Detail of plastid contain single pyrenoid. 60 Algal Flora of Korea·Nemalian Red Algae

TYPE: Beaufort, North Carolina, USA (NA: United States National Herbarium, Washington D.C.). SEASONALITY: Rarely observed summer season (June to September). DISTRIBUTION: North Atlantic Ocean, Sargasso Sea, Africa (Algeria; Báez et al. 2005), Japan, Korea. KOREA: Jeju and Dokdo Island. SPECIMEN EXAMINED: KN5342 (Sagye, Jeju Island: 3.vi.2009, endozoic on inner fiber of Callyspongia elegans). REMARKS: Colaconema infestans was reported from Jeju and Dokdo Island in Korea (see Lee and Kang 2001), as endozoic plants in sponges. This species is easily observed from the cast ashores of sponge (eg. Callyspongia elegans) in Jeju Island from June to August. Korean plants of C. infestans agree quite well with the original description and figures (Howe and Hoyt 1916: 116, pl. 14., f. 1-12) and with the later description given by Woelkerling (1973a, f. 5, 6). However, there is some differences between British specimens described by Dixon and Irvine (1977, p. 99, f. 28). Our specimens grow on inner fiber of spongia, but most of described specimens of this species living in the inner Fig. 46. Distribution of Colaconema infestans. layers of the perisarc of hydroids (Woelkerling 1973a; Dixon and Irvine 1977). Our specimens grow well free from host in laboratory culture condition. The morphology and development of endozoic filaments of C. infestans appear to be some of those of Acrochaetium spongicolum Weber van Bosse (1921). However, the plants of the latter species do not exsert the erect filaments from the host surface (Stegenga 1985; Woelkerling and Womersley 1994). Woelkerling (1973a) discussed several species related to C. infestans. According to Woelker- ling (1973a), Acrochaetium effusum Levring (1953) and Rhodochorton penetrale Drew (1928) are conspecific with this species. C. infestans is also closely related to Audouinella endozoica (Darbishire) Dixon (Darbishire 1899; as Chantransia endozoica). Howe and Hoyt (1916) indicated several characters distinguishing the two taxa, A. endozoica and C. infestans: the width of both immersed and exserted filaments, tortuosity and branching mode of immersed filaments, and the size of monosporangia and their stalk cell. Unfortunately, the holotype of A. endozoica is not available so the original illustration given by Darbishire (1899) is referred to for identification (Dixon and Irvine 1977; Woelker- ling 1973a). Further study is needed to clarify the relationship between the two species, C. infestans and A. endozoica (see also Lee 1987). Nemaliophycidae: Colaconematales: Colaconemataceae 61

21. Colaconema pacificum (Kylin) Woelkerling 1971: 47 (Figs. 47-49). Dae-yang-hong-da-bal-sol (대양홍다발솔: 신칭)

Woelkerling 1971: 47. f. 17A-D, 26A.

BASIONYM: Acrochaetium pacificum Kylin 1925: 11. f. 4g-i. SYNONYM: Audouinella pacifica (Kylin) Garbary 1979: 490. Garbary et al. 1982: 37. f. 15. Woelkerling and Womersley 1994: 58. f. 11E-I. Chantransia pacifica (Kylin) Levring 1941: 631. Rhodochorton pacificum (Kylin) Drew 1928: 169. f. 25.

Thalli epiphytic and epizoic, caespitose, heterotrichous with prostrate and erect filaments, 5-10 mm high, purple fire-red colour. Basal prostrate filaments creeping on substrate, forming unistratose layer over the surface, giving rise to erect filaments. Erect filaments rarely branching, producing numerous monosporangia from almost cells in the middle to upper portion of the cells. Cells of erect filaments cylindrical, 7-8 μm wide, 12-18 μm long. Plastids single per cell, parietal laminate with a pyrenoid. Hairs not formed. Monosporangia unilateral, on short one- or three celled stalks, oblong in shaped, 5-7 μm wide, 8-11 μm long. Tetrasporangia not observed.

TYPE: San Juan Island, Washington, USA (Not designated, see Garbary et al. 1982). SEASONALITY: Rarely observed summer season (June to September). DISTRIBUTION: Pacific Ocean (USA, Japan, Austra- lia). KOREA: Jeju Island. SPECIMEN EXAMINED: KN6213 (Udo islet, Jeju Island: 6.vi.2010, epizonic on hydroids), KN-CJU003 (Jukdo islet, Ulleung Island: 23.vi.2009). REMARKS: This is a new record of C. pacificum from Korea. Korean C. pacificum is almost identical with description of Kylin (1925) except the branching mode. Kylin described the branching as irregular and more or less rich in Califonia plant, but those are rare in Korean plants. Thus, Korean specimens require more investigations to certify the taxonomic entity. Abbott and Hollenberg (1976) used a wide species concept for this species and considered Rhodochorton elegans, R. magnificum, R. plumosum and R. variabile of Drew (1928) to be conspecific with C. pacificum. Garbary et al. (1982) considered Acrochaetium iyengarii Børgesen as Fig. 47. Distribution of Colaconema paci- closely related, if not conspecific with this species. ficum. 62 Algal Flora of Korea·Nemalian Red Algae

AB C

DE F G

Fig. 48. Colaconema pacificum. A. Habits of epizoic thalli on hydroid (Udo islet, Jeju Isalnd); B. Heterotrichous organization with prostrate and erect filaments; C. Detail of prostrate basal filaments; D. Detail of plastid; E-H. Cultured plants; E, F. Nemerous pedicellate monosporangia developed unilaterally from the middle to upper portion of erect filaments; G. Germling of monospore; H. Detail of monosporangia and plastid. Nemaliophycidae: Colaconematales: Colaconemataceae 63

ABC

DEFG

Fig. 49. Colaconema pacificum (KU-CJU003). A-G. Cultured plants (isolated from Jukdo islet, Ulleung Island); A, B. Heterotrichous organization with prostrate and erect filaments; C-E. Nemerous pedicellate monosporangia developed unilaterally from the middle to upper portion of erect filaments; F, G. Detail of plastid. 64 Algal Flora of Korea·Nemalian Red Algae

22. Colaconema thuretii (Bornet) Gabrielson in Gabrielson, Widdowson, Lindstrom, Hawkes et Scagel 2000: 40 (Figs. 50-52). Wang-hong-da-bal-sol (왕홍다발솔: 개칭)

Gabrielson et al. 2000: 40.

BASIONYM: Chantransia efflorescens var. thuretii Bornet 1904: XVI. pl. 1. f. 1-6. SYNONYM: Acrochaetium thuretii (Bornet) Collins and Harvey 1917: 98. Audouinella thuretii (Bornet) Woelkerling 1971: 9. f. 9. Lee 1987: 21. f. 9. Chantransia thuretii (Bornet) Kylin 1907: 119. f. 28. Rhodochorton thuretii (Bornet) Drew 1928: 152.

Thalli epiphytic, forming hemispherical to spherical fringes, composed of prostrate and erect filaments, up to 5 mm high, bright red colour. Prostrate filaments branching, entangled, coherent together, forming a massive basal structure. Cells of creeping filaments contortedly cylindrical with ca. 3 μm thick walls, 9-11 μm wide, 20-28 μm long. Erect filaments straight, rigid, tapering gradually to tips, branching more frequent towards apices alternate or secund at shorter or longer intervals. Cells of erect filaments cylindrical, not constricted at junction, 10-11 μm wide, 30-50 μm long. Plastids single per cell, parietal laminate with a pyrenoid. Monosporangia solitary or in pairs on single-celled stalks, on adaxial side of the lower region of branches or on short laterals, long, ellipsoid in shape, 11-13 μm wide, 19-22 μm long. Tetrasporangia or sexual reproductive organs not observed.

TYPE: Cherbourg, France (?). SEASONALITY: Rarely observed summer season (June to September). DISTRIBUTION: World cosmopolitan. KOREA: South and East coasts. SPECIMEN EXAMINED: KU654 (Deoksan harbor, Sam- cheok), KU742 (Salpido islet, Yeosu: 22.vi.2009), KN6400 (Jumunjin, Gangneung: 8.vii.2009, on rock). REMARKS: The Korean plants of Colaconema thuretii has characteristic hemispherical to spherical fringes here and there on the host thallus, having straight and somewhat rigid erect filaments, and a single parietal laminate plastid with a pyrenoid in long cylindrical cells of erect filaments (Lee 1987). These charcters of Korean plants agree well with the descriptions and illustrations of Bornet (1904: XVI. pl. 1. f. 1-6), who examined Thuret’s collections under the name of Chantransia corymbifera Thuret and found the collections to be a mixture of two elements; one epiphytic on Helminthocladia and the other on Ceramium. Bor- net (1904), therefore, identified the former as Chantran- sia corymbifera and described the latter as Chantransia Fig. 50. Distribution of Colaconema thu- efflorescens var. thuretii Bornet. Rosenvinge (1909) retii. Nemaliophycidae: Colaconematales: Colaconemataceae 65

ABC

DEF

Fig. 51. Colaconema thuretii. A-F. Cultured plants (KU654; Deoksan, Samcheok); A. Habit of caspi- tose thallus; B. Features of prostate filaments and subdichotomous-alternate branching pattern of erect filaments; C. Monosporangia born on short laterals or secundly on branches; D. Numerous branches on middle to upper portion of erect filament; E. Detail of pedicellate monosporangia; F. Detail of plastid. 66 Algal Flora of Korea·Nemalian Red Algae

ABC D E

FG HI

Fig. 52. Colaconema thuretii (KU742; Salpido islet, Yeosu). A-E. Germination and morphogenesis in culture (monosporangia usually bleached; arrows); F. Numerously branched on erect filaments with monosporangia clusters; G, H. Monosporangia; I. Detail of monosporangia and plastid. Nemaliophycidae: Palmariales: Rhodophysemataceae 67 separated Danish specimens of Chantransia thuretii into two varieties; C. thuretii var. amphicarpa and C. thuretii var. agama. However, it was not generally accepted by later workers (Abbott and Hollen- berg, 1976; Garbary et al. 1982; Lee 1987). Woelkerling (1973b) regarded C. thuretii to be synonymous with Audouinella saviana (Meneghini) Woelkerling after comparing specimens from the type collections of the two taxa.

Order Palmariales Guiry et Irvine in Guiry 1978: 138. Pal-son-i-pul-mok (팔손이풀목)

Thalli various, from free filamentous, pseudo-parenchymatous, or parenchymatous. Cell fusion occurred in pseudo-parenchymatous construction. Tetrasporophytes grow directly on the fertilized carpogonium, simple or mutlicelluar, producing cruciate tetrasporangia. Plastid variable, one to several per cells, parietal lobed to discoid with or without pyrenoid. Asexual reproduction by regenerative tetrasporangia, no monosporangia. Sexual life history unusual, lacking carposporophyte generation, no carpogonial branch or auxiliary cell.

REMARKS: Originally, Guiry and Irvine (1978) established the order Palmariales for the family Palmariaceae which includes the genera Palmaria, Halossaccion and Leptosarca formerly included in the order Rhodymeniales based on the characteristics of the absence of carposporophyte and the presence of a generative tetrasporangial mother cell. The evidences for the ordinal level disctinc- tiveness of Palmariales were provided by life history investigations (Van der Meer and Todd 1980; DeCew and West 1982) and ultrastructural survey of pit plugs (Pueschel and Cole 1982). Surprisingly, the order Palmariales was reveled that it is closely related with Acrochaetiales and Nemaliales based on molecular phylogenetic investigations by Saunders’ group (Saunders and McLachlan 1991; Saunders et al. 1995; Harper and Saunders 1998, 2001, 2002; Saunders and Hommersand 2004; Clayden and Saunders 2010). Now, Palmariales is classified into 4 families; Palmariaceae, Rhodophysemataceae, Meiodiscaceae and Rhodothamniellaceae. In Korea, three species has been listed up; Palmaria palmata (Palmaria- ceae), Rhodophysema georgii (Rhodophysemataceae) and Rhodonematella subimmersa (as named Rho- dochorton subimmersum; Rhodophysemataceae). However, the taxonomic identities of the former two species are ambiguous at present in Korea.

Family Rhodophysemataceae Saunders et McLachlan 1989: 20. Beo-jim-deo-bu-sal-i-gwa (버짐더부살이과)

Thalli free filamentous or pseudo-parenchymatous, crustose on rock or other algae, or endophytic in other algae. Cellular fusion occurred in pseudo-parenchymatous construction. Tetrasporophytes reduced to diminutive filamentous, producing cruciate tetrasporangia directly on one-celld stalk cell derived from fertilized carpogonium. Plastids variable, one to several per cells, parietal lobed 68 Algal Flora of Korea·Nemalian Red Algae to discoid with or without pyrenoid. Asexual reproduction by regenerative tetrasporangia, no monosporangia. Sexual life history lacking of carposporophyte generation, and absent of carpogonial branch and auxiliary cell. Type genus: Rhodophysema Batters 1900: 377.

GENERA AND SPECIES: 4 genera at present (5). DISTRIBUTION: Worldwide in marine. KEY REFERENCES: Saunders and McLachlan (1989), Clayden and Saunders (2010). REMARKS: Saunders and McLachlan (1989) recognized the taxonomic affinities of Rhodophysema with Acrochaetiales-Palmariales complex based on molecular and morphological data, and thus they proposed the second family of Palmariales, Rhodophysemataceae, for receiving this genus. They suggested the provisional inclusion of this family to the Palmariales by the following characteristics; 1) having pit plugs with two membranes, 2) lacking of carposporophyte and monosporangia in the life history, 3) lacking of carpogonial branch, 4) presence of regenerative stalk cell associated with tetrasporangia, 5) occurrence of celluar fusions in pseudo-parenchymatous thallus. And also, they argued that the absence of tetrasporophyte in the life history and occurrence of B- phycoerythrin rather than R-phycoerythrin, were main characters for separating the second family from Palamiaceae. The familian concept was guaranteed more strongly and extended somewhat broadly for receving a new genus Rhodonematella based on molecular phylogenetic analysis and morphological reproductive characteristics (Clayden and Saunders 2010).

Key to the gerea of family Rhodophysemataceae

1. Thalli pseudo-parenchymatous, crustose on other algae·······································Rhodophysema - Thalli free filamentous, endophytic usually on Gratelopia sp.·······························Rhodonematella

Genus Rhodonematella Clayden et Saunders 2010: 297. Sum-gin-bul-geun-teol-sok (숨긴붉은털속: 신칭)

Thalli gametophytic stage, endophytic, consisting of basal and erect systems. Basal system composed of endophytic filamentous cells, immersed in the host cotex, but not penetrating host cells. Erect system consisting of free filaments. 1-5 celled, arising directly from basal cells. Game- tophytes monoecious with sessile carpogonia and spermatangia on 1-2 celled erecte filaments. Tetrasporophytes diminutive, surrounded by presumed gametophytic multi-celled filaments (paraphyses), developed directly from fertilized capogonia, bearing cruciate tetrasporangia. Type species: Rhodonematella subimmersa (Setchell et Gardner) Clayden et Saunders 2010: 297.

SPECIES only one at present. DISTRIBUTION: Cosmopolitan, widely distributed in temperate and tropical waters. KEY REFERENCES: Clayden and Saunders (2010). REMARKS: The genus Rhodonematella was established recently based on Rhodochorton subimmersum by Clayden and Saunders (2010). They certified the taxonomic postion of this genus based on phylogenetic analyses of LSU-rDNA and coxI mitDNA sequencing data. Nemaliophycidae: Palmariales: Rhodophysemataceae 69

23. Rhodonematella subimmersa (Setchell et Gardner) Clayden et Saunders 2010: 297 (Figs. 53, 54). Sum-gin-bul-geun-teol (숨긴붉은털: 개칭)

BASIONYM: Rhodochorton subimmersum Setchell et Gardner 1903: 347. pl. 17. f. 12. SYNONYM: Acrochaetium subimmersum (Setchell et Gardner) Papenfuss 1945: 318. Audouinella subimmersa (Setchell et Gardner) Garbary and Rueness 1980: 22. Rhodochorton subimmersum Setchell et Gardner sensu Lee and Kurogi 1978: 115. f. 1-10. Lee 1987: 45. f. 22.

Thalli (gametophytic) partly endophytic usually on Grateloupia turuturu, forming irregular-shaped dark red spots on host, composed of endophytic basal and erect filaments, up to 50 μm high. Endophytic filaments developing in host tissue, runnig along the outer medulla layer of the host and forming networks in cortical layers. Cells of endophytic filaments conspicuously longer and broader than erect filaments, cylindrical, 8-10 μm, 35-55 long. Erect filaments projecting out, straight, 3-5 celled high, simple or sometimes forked at base, bearing reproductive structures, slightly tapered toward apices. Cells of erect filaments oblong, 3-5 μm wide, 5-7 μm long. Hairs absent. Tetrasporangia born terminally, directly on fertilized carpogonium (probable) with forming a generative tetrasporangial mother cell, cruciately dividing, ellipsoid to obovoid, 10-13 μm, 14- 16 μm long. Regeneration occurring in empty sporangia. Tetrasporophytes absent. Thalli monoecious; male and female reproductive structures usually born on one- to three-celled stalks or directly on the cell of endophytic filaments near host surface. Spermatangia solitary or in pairs on spermatangial mother cells, globose to ellipsoid, 3-4 μm in diameter. Carpogonia conical to bottle-shaped with terminal trichogynes, 2-3 μm wide, 6-8 μm long. Trichogynes constricted at junction with inflated tips, 1-2 μm wide, 5-4 μm long.

TYPE: Whidey Island, Washington, USA (UC no. 96094; Herb. UC, Berkeley, California) SEASONALITY: Winter to summer (January to Sep- tember). DISTRIBUTION: Cosmopolitan. KOREA: South and East coasts. SPECIMEN EXAMINED: KN1109 (Jumunjin, Gangneung: 18.vii.2010). REMARKS: Korean plants of R. subimmersa endophytic in Grateloupia turuturu which are almost similar with descriptions and illustrations provided by Clayden and Saunders (2010). Lee and Kurogi (1978) reported that R. subimmersa has the sexual cycle with reduced carposporophytic phase, which was the pattern simi- Fig. 53. Distribution of Rhodonematella lar to those of Rhodochorton purpureum and R. floridulum subimmersa. 70 Algal Flora of Korea·Nemalian Red Algae

ABC

DE F

Fig. 54. Rhodonematella subimmersa. A. Habit of endophytic thalli on Grateloupia turuturu (arrows); B, C. Cross section view large basal filaments and erect filaments; D, E. Representative of the Palma- riales type of tetrasporangial mother cell (arrows); F. Carpogium with trichogyne (arrow).

(West 1969, 1970; Stegenga 1979). However, R. subimmersa has a much reduced the tetrasporophytic phase as like in Palmarian taxa (Clayden and Saunders 2010). Accordiing to the Lee (1987), the plants of R. subimmersa from Korea, Japan and British Columbia show neither spore germination nor growth under host free culture conditions. Nemaliophycidae: Nemaliales: Galaxauraceae 71

Order Nemaliales Schmitz in Engler 1892: 17. Guk-su-na-mul-mok (국수나물목)

Thalli erect or decumbent, terete or compressed, usually subdichotomously to irregularly branched, calcified or non-calcified, multiaxially organized with a central filamentous medulla and anticlinal filaments forming a loose or coherent cortex. Secondary cell fusion present in some taxa, but secondary pit-connections absent. Post-fertilization development simple, producing gonimoblast directly from fertilized carpogonium without specialized auxiliary cell. Carposporophyte immersed in the thallus or bursting out, naked or with a distinct pericarp or loose involucres of subsidiary filaments. Life history triphasic, with isomorphic gametophytes and tetrasporophytes, or hetermor- phic with a conspicuous gametophyte and a minute filamentous or crustose tetrasporophytes.

Key to the families of order Nemaliales (Huisman 2006d)

1. Thallus with terete or compressed branches with a cortex of loosely arranged filaments; carpogonial branches various from 3- to many celled; hypogynous cell generally naked or rarely short filaments; carposporopyte naked or with a loose involucres····································Liagoraceae - Thallus with flattened or terete branches with a distinct cortex of laterally coherent or fused cells or utricles; carpogonial branches 3-celled; hypogynous cell bearing nutritive cells; carposporopyte with a distinct pericarp and ostiole ··························································································2 2. Thallus calcified with distinctly flattened or terete axes; life history isomorphic or differing cortical structures between gametophytes and tetrasporophytes or markedly heteromorphic············ ····························································································································Galaxauraceae - Thallus noncalcified with terete axes or slightly compessed; life history markedly heteromorphic with a filamentous or crustose tetrasporophyte························································Scinaiaceae

REMARKS: Nemaliales was proposed by Schmitz (1892) as one of the four orders of the subclass Florideophycidae at that time. Originally, the order was characterized by simple post-fertilization development in which the gonimoblastis developed directly from the fertilized carpogonium, without transferring the zygote nucleus to a specialized auxiliary cell. Base on the order’s circumscription, the Nemaliales included both uniaxial and multiaxial red algal taxa; Helminthocladiaceae, Chaetangiaceae (=Galaxauraceae), Acrochaetiaceae, Batrachospermaceae, Lemanaceae, Thoreaceae, Bonnemaisoniaceae, Naccariaceae (Fritsch 1945; Kylin 1956). However, Nemaliales is restricted to multiaxial taxa at present, and thus the uniaxial taxa separated into new several orders as Acrochaetiales, Batrachospermales (including Lemanaceae, Thoreaceae), and Bonnemaisoniales (including Naccariaceae). The systematics of Nemaliales is currently revised at various taxonomic ranks base on DNA sequence analysis. As a result, the order was subdivided into three family currently as Liagoraceae (including Nemaliaceae, Dermonenmataceae), Galaxaur- aceae, and Scinaiaceae (Huisman et al. 2004; Schneider and Wynne 2007).

Family Galaxauraceae Parkinson 1983: 608. Ga-wi-son-mal-gwa (가위손말과)

Thalli erect or partially decumbent, terete or compressed, calcified, subdichotomously branched, 72 Algal Flora of Korea·Nemalian Red Algae multiaxially organized with a central filamentous medulla and anticlinal sudichotomously branched assimilatory filaments forming a cortex. Cortex filamentous or becoming pseudoparenchymatous, often with fusions between adjacent cells. Cells cylindrical or ovoid. Spermatangia borne on inner cortical filaments, hypodermal cells, or within cavities. Gonimoblast developed directly from the fertilized carpogonium. Carposporophytes within the outer medulla without protruding, having an ostiole with branched filaments and terminal carposporangia produced single or in short chains, surrounded by a distinct involucral filaments arising from the basal cell of the carpogonial branch or derived from gonimoblast filaments. Tetrasporophytes either filamentous, crustose, or of similar form to gametophytes, producing cruciately divided tetrasporangia. Type genus: Galaxaura Lamouroux 1812: 185.

GENERA AND SPECIES: 4 genera (38). DISTRIBUTION: Worldwide in tropical to subtropical sea. KEY REFERENCES: Huisman and Womersley (1994), Wang et al. (2005).

Key to the genera of family Galaxauraceae

1. Tetrasporangia produced terminally on elongate assimilatory filaments ··································2 - Tetrasporangia produced on epidermal cells of cortex, subcortical cells enlarged by fusion of neighboring cells ····································································································Dichotomaria 2. Tetrasporophytes and gametophytes distinctly heteromorphic, the former minute and filamentous ··························································································································Tricleocarpa - Tetrasporophytes and gametophytes isomorphic or dimorphic················································3 3. Tetrasporophytes and gametophytes isomorphic, axes bearing whorled emergent hairs of regular invervals, subcortical cells uniformal without enlarging by fusion of neighboring cells ········· ································································································································Actinotrichia - Tetrasporophytes and gametophytes dimorphic, axes bearing hairs but do not arranged whorled with regular intervals, subcortical cells enlarged by fusion of neighboring cells··········Galaxaura

REMARKS: Galaxauraceae is characterized by a three-celled carpogonial branch, absence of an auxiliary cell, direct development of gonimoblasts from the fertilizied carpogonium, two to four sterile branches borne on the hypogynous cell that become modified with enlarged nuclei after fertilization, basal cell of the carpogonial branch bearing prefertilization lateral sterile filaments that develop extensively after fertilization, and cystocarps that are deeply immersed in the fronds and prominently ostiolate (Huisman and Womersley 1994; Wang et al. 2005). Huisman et al. (2004) reduced the Galaxauraceae to four calcified genera, Actinotrichia Decaisne (1842), Galaxaura Lam- ouroux (1812), Tricleocarpa Huisman et Borowitzka (1990), and Dichotomaria Lamarck (1816), and transferred four noncalcified genera (Scinaia, Gloiophloea, Nothogenia, Whidbeyella) into their newly established family, the Scinaiaceae, based on molecular phylogenetic analysis of nuclear LSU- rDNA sequences. Owing to the similarilty of the anatomical structures of Galaxaura and Dichotomaria in gametophytic phase, it is very difficult to distinguish both genera without observations of their tetrasporophytic phases. Ten species names of Galaxaura were listed in Korean marine algal floristic list (Lee and Kang 2001; Lee 2008) as follows: G. apiculata Kjellman, G. clavigera Kjellman, G. falcata Kjellman, G. filamentosa Chou ex Taylor, G. hystrix Kjellman (=G. apiculata), G. marginata (Ellis et Solander) Lamouroux, G. rugosa (Ellis et Sonder) Lamouroux, G. rudis Kjellamn (=G. rugosa), G. pilosula Kjell- Nemaliophycidae: Nemaliales: Galaxauraceae 73 man (=G. marginata), and G. subverticillata Kjellman (=G. rugosa) (Lee and Kang 1986; Lee and Kang 2001; Lee 2008). However, most of the reported species in Korea has been ambiguous of their taxonomic entities and needed to their comparative morphological works including DNA sequences of phylogenetic studies. Recently, Lee (2008) recognized six species of Galaxaura; G. apiculata, G. hystrix, G. falcata, G. marginata, G. pilosula, and G. rugosa from Jeju Island. Among them, only one species G. rugosa (=G. pilosula) is remained in the genus Galaxaura and the others has been transferred to the resurrected genus Dichotomaria Lamarck (1816: 145) base on anatomy and molecular data (Huisman et al. 2004).

Genus Actinotrichia Decaisne 1842: 118. Go-ri-bang-sa-teol-sok (고리방사털속)

Thalli partially decumbent, subdichotomously to irregularly branched, attached by discoid holdfasts and by several secondary holdfasts. Erect axes multiaxial, terete mostly of uniformal diameter, heavily calcified, often bearing whorled emergent filaments on subepidermal cells. Central axial filaments bearing anticlinal dichotomously branched assimilatory filaments, forming a pseudoparenchymatous cortex. Cortical cells decrasing in size towards the periphery, spherical or wedge- shaped in its outmost cells. Gametophytes monecious, isomorphic with tetraspophytes. Sperma- tangia in sunken cavitie, terminal on free filaments projecting into the cavity from all faces of the conceptacle wall. Carpogonial branches accessory or arising in place of normal vegetative filaments. Post-fertilisation hypogynous cell with 3 nutritive branches, basal cell with 3 or 4 small- celled sterile branches. Gonimoblast initials arising directly from the fertilized carpogonium, forming a pseudoparenchymatous, with carposporangia produced singly from the apices of gonimoblast filaments. Pericarp formed from the basal cell of the carpogonial branch. Tetrasporangia cruciately divided, borne laterally and pedicellate, or terminally on the whorled ermergent filaments. Type species: Actinotrichia rigida (Lamouroux) Decaisne 1842: 118 [=Actinotrichia fragilis (Forsskål) Børgesen].

SPECIES 4 (1 in Korea). DISTRIBUTION: Widely distributed in temperate and tropical waters in West Pacific, Australia and Newzealand, and Indian Ocean. KEY REFERENCES: Huisman (2006a), Liu and Wang (2009). REMARKS: Although the genus Actinotrichia was established in early by Decaisne (1842), but its phylogenetic position base on molecular data and the anatomical features of reproductive struc- trures were resolved clearly by recent works (Wang and Chiang 2001; Huisman et al. 2004, Wang et al. 2005, Liu and Wang 2009). In Korea, only one species A. fragilis was reported mostly in Jeju Island by several authors (Lee and Kang 2001), and excellent illustrations of Korean materials were provided recently by Lee (2008). Despite a few attempt had fail to collect this species in this work. Thus, we describe this genus and A. fragilis based on the related references and illustrations of Lee (2008) without observation of specimens. 74 Algal Flora of Korea·Nemalian Red Algae

24. Actinotrichia fragilis (Forsskål) Børgesen 1932: 6. Go-ri-bang-sa-teol (고리방사털)

Børgesen 1932: 6. pl. 1. f. 4. Kang 1968: 195. f. 65. Lee 2008: 124 with 3 figures. Wang and Chiang 2001: 377. f. 1-16

BASIONYM: Fucus fragilis Forsskål 1775: 190. SYNONYM: Actinotrichia rigida (Lamouroux) Decaisne 1842: 118. Okamura 1916: 30. pl. 158. f. 17- 19. Galaxaura rigida Lamouroux 1816: 265. pl. VIII. f. 4. p. 318.

Thalli to 5 cm high, highly calcified, grey-pink to pale greenish, subdichotomously branched more or less regular interval, forming loosly entangled globular mass. Axes multiaxial, terete and rigid, not segmented, to 500 (-800) μm in diameter, bearing whorled emergent filaments, ending with blunt apices. Whorled filaments uniformal, 150-200 μm long, arising from subepidermal cells, regular intervals of 200-220 μm, forming densely clustered narrow ring, conspicuous in upper portion, but deciduous in older branches leaving annular scars. Medullary filaments arranged axially, composed of elongated cylindrical cell row, 5-12 μm in diameter, producing peripherally a pseudoparenchymatous cortex. Cortex 3 or 4 celled, to 75 μm thick, composed of inner cylindrical cells of 12-16 μm broad, 18-23 μm long, and outer turbinate epidermal cells of 15-18 μm diameter. Teterasporangia and sexual reproductive structures unknown in Korean materials. -Described based on illustrations given by Lee (2008)-

TYPE: Mokha, Yemen (C Herbarium, Botanical Museum, Univ. of Copenhagen, Denmark) SEASONALITY: All season (see Lee 2008). DISTRIBUTION: Widely distributed in temperate and tropical waters in West Pacific, Australia and Newzealand, and Indian Ocean (Guiry and Guiry 2010). KOREA: Jeju Island. SPECIMEN EXAMINED: Not observed in this work. REMARKS: The anatomical features of this species was described and illustrated in detail by Oka- mura (1916; pl. 158, f. 17-19) based on Japanese materials. Korean materials, rarely collected in Jeju Island, were illustrated by photographs in detail (Lee 2008) which are identical with Japanese plants. The reproductive structures of Taiwan materials were investigated in detail by Wang and Chiang (2001) and Liu and Wang (2009).

Genus Dichotomaria Lamarck 1816: 143. gen. emend. Wang et al. 2005: 688. Du-ga-dak-ba-dat-mal-sok (두가닥바닷말속: 신칭)

Thalli ererct, calcified, somewhat rigid, subdichotomously branched, with or without distinct segmenets, attached by discoid holdfasts. Axes multiaxial, terete or flattened, consisted of filamentous medulla and pseudoparenchymatous cortex, occasionally with a terete hirsute basal portion. Cortex mostly 3-layered in gametophyte, with an inner 1-2 layers of large hyaline fusion cells that Nemaliophycidae: Nemaliales: Galaxauraceae 75 are closely packed and an outer layer of small, pigmented cells; 3-6-layered in tetrasporophytes, with an inner 1-3 layers of large hyaline cells and outer layer of ellipsoidal, subglose or wedge- shaped cells borne on pigmented stalk cells. Medulla filamentous. Spermatangia arising in conceptacles; the primary spermmatangial filaments cutting off laterally and transversely at first, later growing distally to form a hemispherical conceptacles and progressively sunken in the cortex; the secondary spermatangial filaments arising from the inner cells of the conceptacles, bearing terminal spermatangia mother cells, each of which producing terminally or subterminally 1-3 spermatia. Carpogonial branches 3-celled with a trichogyne, consisting of a carpogonium, hypogynous cell, and a basal cell; the hypogynous cell bearing 3-4 sterile branches and the basal cell bearing 4-5 involucral filaments. Gonimoblast initials developed from the lower cells of transversely dividied fertilized carpogonuim, growing distally and branched laterally to radially to form gonimoblast filaments. Cystocarps hemispherical in mature, ostiolate, sunken deeply in the cortex/medulla, bearing oval to obovate carposporangia. Tetrasporangia cruciately divided, arising from epidermal cells of cortex, solitary or aggregated in sori on the thallus surface. Type species: Dichotomaria marginata (J. Ellis et Solander) Lamarck.

SPECIES ca. 22 (3 in Korea). DISTRIBUTION: Worldwide from subtropical to tropical sea, usually subtidal. KEY REFERENCES: Huisman et al. (2004), Wang et al. (2005), Kurihara et al. (2005).

Key to the species of genus Dichotomaria

1. Outer cortical cells obovoid with an apiculate apex ··································Dichotomaria apiculata - Outer cortical cell ellipsoidal without an apiculate apex·······························Dichotomaria falcata

REMARKS: The genus Dichotomaria Lamarck (1816: 143) was resurrected by Huisman et al. (2004) and later the generic concept was emended by Wang et al. (2005) as follows; 1) gametophytes and tetrasporophytes that are isomorphic in habit but differ in cortical-cell features, 2) a three-celled corpogonial branch in which, following fertilization, the basal cell produces involucral filaments that ultimately form the pericarp, 3) a multinucleate fusion cell that incorporates the gonimoblast initial and the three to four inner gonimoblast cells, 4) cruciate tetrasporangia produced on stalk cells initiated from cortical or subcortical cells. According to Huisman et al. (2004), Dichotomaria belongs to a separate molecular clade from Galaxaura (gereic type: G. rugosa) is morphologically characterised by tetrasporangia formed laterally or terminally on stalk cells of epidermal cells and has two-celled outer layer in cortex of tetrasporophyte. On the other hand, Galaxaura is characterised by the production of tetrasporangia on elongate assimilatory filaments and has a filamentous cortex in tetrasporophyte. Galaxaura apiculata, G. falcata, and G. papillata are included in a monophyletic clade along with the generic type species, Dichotomaria marginata, so that the three species in Japan were transferred into genus Dichotomaria (Kurihara et al. 2005). In Korea, three species were reported as named Galaxaura apiculata (conspecific with G. hystrix; Kurihara et al. 2005), G. falcata and G. marginata (conspecific with G. pilosula; Huisman and Borowizka 1990) without observations of their reproductive structures (Lee 2008). Even so, Lee (2008) reported Dichotomaria marginata (as named G. marginata) in Jeju Island, the being of it in coast of Korea is in doubt. Based on the illustrations given by (Lee 2008), the Korean D. marginata is never similar with those of Australian in morphology of outer cortical cell and gross habit of thallus (Huisman and Borowitzka 1990, f. 14-27). We considered the reporting of D. marginata in Jeju Island is resulted from misidentifying of D. falcata. Moreover, we could not find the distinction between Galaxaura 76 Algal Flora of Korea·Nemalian Red Algae pilosula sensu Lee (2008) and Dichotomaria falcata except the hair-like filaments on upper thallus in Galaxaura pilosula. However, the hair-like filaments are frequently observed in D. falcata in this work. Thus, we excluded D. marginata and Galaxaura pilosula as Korean members of Dichotomaria in this work.

25. Dichotomaria apiculata (Kjellman) Kurihara et Masuda 2005: 52 (Figs. 55-57). Ppyo-jok-du-ga-dak-ba-dat-mal (뾰족두가닥바닷말: 개칭)

Kurihara et al. 2005: 52. f. 15-24.

BASIONYM: Galaxaura apiculata Kjellman 1900: 74. pl. 12. f. 13-26. pl. 20. f. 36 (tetrasporphytic phase). Tanaka 1936: 162. f. 26, 27. pl. XLI. f. 2. Lee 2008: 215 with 3 figures. SYNONYM: Galaxaura hystrix Kjellman: 1900: 79. pl. 16. f. 1-10. pl. 20. f. 34 (gametophytic phase). Tanaka 1936: 166. f. 32, 33. pl. XLIII. f. 2. Lee and Lee 1989: 6. f. 4. Lee 2008: 217 with 3 figures.

Thalli tufted with short stipe, dark or dirty red, up to 10 cm high, dichotomously branched several times on same plane, forming flabellate habit, calcified at intercellular space of cortex with faintly transverse striation, attached by a holdfast. Stipe terate, villose, to 1.5-3 mm diameter. Axes flattened except basal stipe, 2-4 mm wide, consisted of medulla and cortex. Medulla composed of periclinally arranged and intertwined colourless filaments, 9-12 μm in diameter. Cortex two layers with inner and outer cortices, composed of retangular to irregular inner cells and pedicellate epidemal cells, dimorphic between tetrasporophytes and gametophytes of their structure of epidermal cells. Inner cortex almost similar in tetrasporophytes/gametophytes, two or three layers of tightly (tetrasporophytes) and loosely (gametophytes) packed cells, 80-140 μm thick, consisted of hyaline retanglar or irregular shaped cells of 30-100 μm wide and 25-70 μm high in cross section view. Epidermal cells obovoid or clavate, with an apiculate apex (rarely with a round apex among them) in cross section, arising on 1- or 2-celled cylindrical pedicel, 15-25 broad, 35-43 μm long. Cystocarp spherical to flattened, 500- 550 μm broad, 300-350 high in cross section view. Tetrasporangia and spermatangia not observed in this work.

TYPE: Unknown locality in Japan (D. apiculata), Goto Fig. 55. Distribution of Dichotomaria api- Islands, Nagasaki Pref. Japan (Galaxaura hystrix). culata. Nemaliophycidae: Nemaliales: Galaxauraceae 77

ABC

DEF

Fig. 56. Dichotomaria apiculata (KN7021; Dokdo Island). A. Habit of gametophytic thallus; B, C. Cross section view in middle portion of branch; D, E. Detail of apiculate apex of outer cortical cells; F. Detail of the organization of outer and inner cortex and medulla. 78 Algal Flora of Korea·Nemalian Red Algae

A B C

DE F

Fig. 57. Dichotomaria apiculata (KN7011; Pyoseon, Jeju Island). A. Habit of gametophytic thallus; B. Detail of base; C. Cross section including a cystocarp; D. Cross section a flattened branch show a calcified layer (arrow); E, F. Detail of outer cortical cells with a mucronate apex (rarely some of them with round apex) on cylindrical pedicels. Nemaliophycidae: Nemaliales: Galaxauraceae 79

SEASONALITY: All season (see Lee 2008). DISTRIBUTION: Japan and Korea (see Kurihara et al. 2005). KOREA: Jeju and Dokdo Island. SPECIMEN EXAMINED: KN7021 (Dokdo Island: 6.vii.2010), KN7011 (Pyoseon, Jeju Island: 5.vi.2009). REMARKS: The taxonomic identity of Dichotomaria apiculata was newly emended by Kurihara et al. (2005), and they discovered the conspecificity of this species with Galaxaura hystrix based on DNA sequencing phylogeny of rbcL and ITS1. The tetrasporophytes (=Galaxaura hystrix) of Korean materials was well described and illustrated by Lee and Lee (1989: 6. f. 4). Korean plants agree well with the description and illustrations provided by Kurihara et al. (2005).

26. Dichotomaria falcata (Kjellman) Kurihara et Masuda 2005: 52 (Figs. 59, 60). Yeo-rin-du-ga-dak-ba-dat-mal (여린두가닥바닷말: 개칭)

Kurihara et al. 2005: 52.

BASIONYM: Galaxaura falcata Kjellman: 1900: 73. pl. 11. f. 12-21. pl. 12. Kang 1968: 195. pl. 38. f. 140 with text f. 66. Lee 2008: 216 with 3 figures.

Thalli tufted with short stipe, dark or dirty red, up to 12 cm high, dichotomously branched several times on same plane, forming flabellate habit, calcified at intercellular space of cortex with faintly transverse striation, attached by a holdfast. Stipe terate, villose, to 1.5-2.5 mm diameter. Axes flattened except basal stipe, 2.0-3.5 mm wide, consisted of medulla and cortex. Medulla composed of periclinally arranged and intertwined colourless filaments, 10-12 μm in diameter. Cortex two layers with inner and outer cortices, composed of retangular or irregular inner cells and pedicellate epidemal cells, dimorphic between tetrasporophytes and gametophytes of their structure of epidermal cells. Inner cortex almost similar in tetrasporophytes/gametophytes, two or three layers of tightly (tetrasporophytes) and loosely (gametophytes) packed cells, 200-450 μm thick, consisted of hyaline retanglar or irregular shaped cells of 30-120 μm wide and 25-100 μm high in cross section view. Epidermal cells ellipsoid, without an apiculate apex (always) in cross section view, arising on 1-celled cylindrical pedicel (gametophyte) or on retangular pedicel (tetrasporophyte), 20-30 μm broad, 35-40 μm high. Reproduc- Fig. 58. Distribution of Dichotomaria fal- tive structure not observed in this work. cata. 80 Algal Flora of Korea·Nemalian Red Algae

CD E

Fig. 59. Dichotomaria falcata (Udo islet, Jeju Island). A. Habit of thallus; B. Detail of longitudinal section views; C. Cross section view; D, E. Detail of ellipsoidal-shaped outer cortical cells without mucronate apex on a cylindrical pedicel (gametophytic type). Nemaliophycidae: Nemaliales: Galaxauraceae 81

A BC

DEF

Fig. 60. Dichotomaria falcata (Pyoseon, Jeju Island). A. Habit of thallus; B. Detail of base producing young proliferations (arrow); C. Cross section view of flattened branch show calcified layer (arrow); D. Inner and outer cortical cells; E, F. Detail of outer cortical cells on a rectangular pedicel (arrow; tetrasporophytic type). 82 Algal Flora of Korea·Nemalian Red Algae

TYPE: Yenosima, Japan. SEASONALITY: All around year. DISTRIBUTION: Japan and Korea. KOREA: South coast including Jeju Island. SELECTED SPECIMENS: KN7014 (Udo islet, Jeju Island: 6.vi.2010), KN7008 (Pyoseon, Jeju Island: 5.vi.2009). REMARKS: This species is more plentiful rather than D. apiculata in coastal castashores of Jeju Island at April to June. It is very difficult to identify both species in field owing to their almost similar gross morphology. Both species, however, are clarly distinguished by the features of outer cortical cells (Kurihara et al. 2005), ellipsoidal without mucronate apex in this species but obovoid- shaped with mucronate apex in D. apiculata.

Genus Galaxaura Lamouroux 1812: 185. Ga-wi-son-mal-sok (가위손말속)

Thalli erect, calcified, subdichomously branched, attached by discoid holdfasts. Axes multiaxial, terete, consisted of filamentous medulla and pseudoparenchymatous or filamentous cortex, occasionally with a hirsute in basal portion. Cortical structure of gametophytes and tetrasporophytes is different; gametophyte cortex three layered, inner one or two layers of large colorless cells often fused laterally; tetrasporophyte cortex filamentous, with or without inflated basal cells supporting assimilatory filaments. Gametophytes usually monoecious. Spermatangia formed in subcortical cavities. Carpogonial branches 3-celled, arising in place of normal vegetative filaments, consisting of a carpogonium, hypogynous cell, and a basal cell; the hypogynous cell bearing 3-4 sterile branches and the basal cell bearing 4-5 involucral filaments before fertilization. Gonimoblast initials arising from the fertilized carpogonuim and radiating outwards, some grow laterally then distally, contributing wall of cystocarp. Cystocarps terminal on gonimoblast filaments, projecting toward the center of the cavity. Tetrasporangia cruciately divided, lateral or terminal on assimilatory filaments. Type species: Galaxaura rugosa (J. Ellis et Solander) J.V. Lamouroux.

SPECIES ca. 22 (1 in Korea; Guiry and Guiry 2010). DISTRIBUTION: Worldwide temperate to subtropical seas, usually subtidal. KEY REFERENCES: Huisman (2006b), Wang et al. (2005). REMARKS: The genus Galaxaura, generic type G. rugosa (Ellis et Solnder) Lamouroux, was erected by Lamouroux in 1812. Numerous species, ca. 99 names, have been described in this genus. How- ever, most species of Galaxaura has been described based on the minor differences of cortical-cell structures as in case of Kjellman (1900) who recognized 62 species including 47 new species. Owing to the discoving of the dimorphism in structure of cortex between gametophytes and tetrasporophytes of Galaxaura, most of Kjellman’s species have been reduced to synonymy (Howe 1917; Papanfuss et al. 1982). Moreover, the molecular phylogenetic studies in recent years (Huisman et al. 2004; Wang et al. 2005; Kurihara et al. 2005) reveal that some of the Galaxaura species must be transferred into the resurrected genus Dichotomaria. Only one species, G. rugosa, was reported in Korea (Lee 2008). Nemaliophycidae: Nemaliales: Galaxauraceae 83

27. Galaxaura rugosa (J. Ellis et Solander) J.V. Lamouroux 1816: 263. Ae-gi-ga-wi-son-mal (애기가위손말)

Lamouroux 1816: 263. Lee and Lee 1989: 2. f. 1. Huisman and Borowitzka 1990: 153. f. 1-13. Wang et al. 2005: 693. f. 3d-g, 4, 7. Dawes and Mathieson 2008: 209. pl. 27. f. 13-16. Lee 2008: 220.

BASIONYM: Corallina rugosa J. Ellis et Solander 1786: 115. pl. 22. f. 3. SYNONYM: Galaxaura collabens J. Agardh 1885: 74. Galaxaura elongata J. Agardh 1876: 529. Galaxaura glabriusculosa Kjellman 1900: 56. Galaxaura lapidescens (J. Ellis et Solander) Lamouroux 1816: 264 Galaxaura pacifica Tanaka 1935: 55. Galaxaura rudis Kjellman 1900: 43. Galaxaura squalida Kjellman 1900: 55. Galaxaura subfruticulosa Chou 1945: 41.

Thallus solitary, 5-8 cm high, dichotomous branching more frequent at upper portion, resulting in an umbellate habit, heavily calcified in intercellular spaces, sparsely villose on whole surface except for terminal segments. Internodal segments terete, more or less rugose, terminal segments club shaped with obtuse or slightly inflated apices. Medullary filaments straight, ending in various directions, more dense near cortex than central portions, ca. 14 μm wide with 5-6 μm thick wall and fine cytoplasm, branching dichotomous to subdichotomous, issuing 4-5 μm wide thin filaments with less than 2 μm thick wall and much contorted terminals showing rhizoid-like form. Cortex 2-3 layer with compacted parenchymatous construction; inner cortical cells largest, connected with medullary filaments, ellipsoid, periclinally elongate, frequently lobed, 20-30 μm wide and 30-50 μm long; outer cortical cells globose to ellipsoid, anticlinally elongate, 10-15 μm wide and 10-20 μm long. Epidermis monostromatic or with long simple filaments; epidermal cells polygonal in surface view, discoid to turbinate in cross section, 12-18 μm wide, 10-12 μm high; epidermal filaments arising on outer cortical cells, with tumid basal cells, simple, 10-13 μm wide, 75-100 μm long, composed of 7-15 cells. Gametophytes monoecious. Spermatangial conceptacles globose with compactly intertwined filamentous pericarps, in outer medullary portions. Sperma- tangia terminal on 2-3 celled branches, solitary, obovoid, 6-8 μm wide, 9-10 μm long. Cystocarps in outer medullary portions, with compactedly intertwined filamentous pericarps. Carposporangia terminal on one to four celled stalks, ellipsoid to pyriform, 22-24 wide, 44-46 μm long. Tetraspo- rophytes not observed in Korea. -Modified the description of Lee and Lee (1989)-

TYPE: Jamaica (the illustration in J. Ellis and D. Solander 1786) SEASONALITY: April (Lee 2008), August (Lee and Lee 1989). DISTRIBUTION: Widespread from temperate to tropical seas, usually subtidal. KOREA: Wimi, Jeju (Lee and Lee 1989). SPECIMEN EXAMINED: Not observed in this study. REMARKS: The description and illustrations of Korean plant provided by Lee and Lee (1989, f. 1) agree well with the Taiwan plants (Wang et al. 2005, f. 3d-g, 4, 7). It is considered that this species is very rare in Jeju Island with 2 times collections by Lee and Lee (1989) and Lee (2008). Despite a few attempts had fail to collect this species in this work. 84 Algal Flora of Korea·Nemalian Red Algae

Genus Tricleocarpa Huisman et Borowitzka 1990: 164. Ga-rak-mal-sok (가락말속)

Thalli (gametophytes) erect, 3-7 cm high, heavily calcified, terete, firm to brittle, dichotomously branched, composed of medulla and cortex, attached by a discoid holdfast. Medulla filamentous. Cortex 3-4 cell layered, with the inner ones inflated and grading into the outer small, pigmented ones, lack secondary fusions in inner cortical cell. Gametophyte monoecious or dioecious. Sper- matangia formed in subcortical and spherical cavities. Carpogonial branches are 3-celled, arising in place of normal vegetative filaments, consisting of a carpogonium, hypogynous cell, and a basal cell; the hypogynous cell bearing 3-4 nutritive branches and the basal cell bearing 3-5 small celled sterile branches that form the pericarp. Pericarps arise from basal cells of carpogonial branches before the formation of gonimoblasts and vegetative filaments projecting into cystocarp cavities and mix with gonimoblastic ones, with paraphyses projecting the cavity. Gonimoblast initials arising from the fertilized carpogonuim, growing laterally along the base and, sometime, the walls of the pericarp. Carposporangia obovoid, terminal on gonimoblast filaments that project toward the central of cavity. Tetrasporophyte filamentous, with cruciate tetrasporagia. Type species: Tricleocarpa cylindrica (J. Ellis et Solander) Huisman et Borowitzka

SPECIES 3 (1 in Korea). DISTRIBUTION: Worldwide subtropical to tropical seas, intertidal and subtidal. KEY REFERENCES: Huisman and Borowitzka (1990). REMARKS: The genus Tricleocarpa was segregated from Galaxaura by Huisman and Borowitzka (1990), based on the distinct, sterile pericarp formed from the basal cell of the carpogonial branch. In Galaxaura the walls of the cystocarp are formed from the gonimoblast filaments themselves. Sterile paraphyese are produced from the pericarp in Tricleocarpa and project into the cystocarp cavity, intermingling with the gonimoblast filaments. The genus also differs in producing a filamentous tetrasporophyte (Magruder 1984), as opposed to essentially isomorphic tetrasporophytes of Galax- aura. Vegetatively, the lack of lateral fusions between cortical cells in the gametophyte also serves to separate the genera.

28. Tricleocarpa cylindrica (J. Ellis et Solander) Husiman et Borowitzka 1990: 164 (Figs. 61, 62). Min-ga-rak-mal (민가락말)

Husiman and Borowitzka 1990: 164. f. 40-45, 50-52. Millar 1990: 306. f. 5A-C. Huisman and Womersley 1994: 116. f. 32G-I, 33E, F. Huisman 2006e: 25. f. 9. Dawes and Mathieson 2008: 211. pl. 27. f. 17, 18. Lee 2008: 221 with 5 figures.

BASIONYM: Corallina cylindrica J. Ellis et Solander 1786: 114. pl. 22. f. 4. SYNONYM: Galaxaura conglutinata Kjellman 1900: 62. pl. 9. f. 14-17. Galaxaura cylindrica (J. Ellis et Solander) J.V. Lamouroux 1821: 22. pl. 22. f. 4. Galaxaura fastigiata Decaisne 1842: 116. Nemaliophycidae: Nemaliales: Galaxauraceae 85

CDE

Fig. 61. Tricleocarpa cylindrica (Munseom islet, Jeju Island). A. Habit of thallus; B. Detail of branches and apex; C. Detail of calcified thallus showing trichotomous branching; D, E. Cross section view of branch show heavily calcified in outer cortical cells.

Galaxaura schimperi Decaisne 1842: 104. Halysium cylindricum (J. Ellis et Solander) Kützing 1843: 323.

Thalli (gametophyte) erect, caespitose, epilithic, pale red to grey-red, 3-7 cm high, dichotomously 86 Algal Flora of Korea·Nemalian Red Algae or rarely trichotomously branched, attached by a discoidal holdfast. Branches terete, 1-1.5 mm broad, glabrous, organized with medulla and cortex. Medul- la of longitudinal filaments (3-15 μm in diameter) giving rise to radiating, dichotomously branched filaments that form the cortex. Cortex 3-4 layered with inflated cells, innermost 18-36 μm in diameter, somewhat longer than broad, grading to outer cortical cells 9-21 μm diameter, 5-6 sided in surface view. Calcifi- cation present in the cortex. Spermatangial cavities spherical, immersed in medulla, 270-280 μm in diameter. Cystocarps spherical to subspherical, immersed in medulla, 265-280 μm in diameter. Tetrasporophyte not observed.

TYPE: West Indies. SEASONALITY: Probably observed all season in subtidal zone. DISTRIBUTION: Worldwide, subtropical to tropical seas. KOREA: Jeju Island (Lee 2008). SPECIMEN EXAMINED: KN7211 (Munseom islet, Jeju Fig. 62. Distribution of Tricleocarpa cylin- Island: 7.i.2010). drica. REMARKS: This species was reported by Lee and Lee (1989) as named Galaxaura oblongata from Jeju Island. Until recently, the taxa Tricleocarpa cylindrica and T. oblongata were considered to represent a single species of Galaxaura, G. oblongata. Huisman and Borowitzka (1990) found that the two species could be recognized on reproductive features and that they represented a genus distinct from Galaxaura. T. cylindrica is commonly found in tropical and subtropical seas.

Family Liagoraceae Kützing 1843: 321, 328. Bun-hong-guk-su-mal-gwa (분홍국수말과: 신칭)

Thalli erect, rarely decumbent, dichotomously or irregularly branched, soft, gelatinous, slimy, or lightly calcified. Axes terete or slightly compressed, multiaxial with medulla of longitudinal filaments and bearing anticlinal subdichotomously to trichotomously branched assimilatory filaments forming the cortex. Cortical filaments loosely arranged, not forming a coherent epidermis. Cells cylindrical, ellipsoidal, obovoid or clavate. Gametophytes monoecious or dioecious. Spermatangia clustered, usually terminal on spermatangial mother cells which arising from outer cortical filaments or in whorls on mid-cortical cells. Carpogonial branches 3- to many celled, arising on cortical filaments, either straight or curved. Hypogynous cells of carpogonial branches usually naked without formation of involucral or nutritive branches. Gonimoblast developed directly from fertilized carpo- Nemaliophycidae: Nemaliales: Liagoraceae 87 gonium. Cystocarps immersed in medulla. Carposporangia are usually terminal on gonimoblastic filaments. Life history triphasic with macroscopic gametophytes and minute tetrasporophytes. Type genus: Liagora Lamouroux 1812: 185.

GENERA AND SPECIES: 20 genera (148). DISTRIBUTION: Worldwide, subtropical to tropical seas. KEY REFERENCES: Huisman and Womersley (1994), Huisman (2006c). REMARKS: Liagoraceae Kützing (1843: 321, 328) pre-dates Helminthocladiaceae J. Agardh (1852: 410). The latter family name was proposed for conservation by Silva (1980: 84, 106) but the proposal was not agreed to by the General Committee (see Taxon Vol. 42, 438). The family is much diver- sified in morphology including about 20 genera at present with widespread in temperature and tropical seas; 5 genera 7 species in Korea. Taxa of the Liagoraceae are known in the field mainly from the conspicuous gametophytes, but the life history is now well known, to involve minute acrochaetioid tetrasporophytes (Womersley 1994). Most of Korean members (5 genera 7 species listed up) of this family are rarely observed in subtidal zone except Nemalion vermiculare, and also some of them are poorly understood of their taxonomic entities and morphological characters at present.

Key to the genera of family Liagoraceae

1. Carpogonial branches 4-8 celled and linear ·································································Nemalion - Carpogonial branches 3-4 celled, usually curved, laterally situated on the middle or basal cells of cortical filaments··················································································································2 2. Thallus not calcified; first division of zygote usually oblique; terminal cells of cortical filaments usually enlarged·································································································Helminthocladia - Thallus calcified; first division of zygote transverse; terminal cells of cortical filaments not enlarged··························································································································Liagora

Genus Helminthocladia J. Agardh 1852: 412. nom. cons. Ga-ji-guk-su-na-mul-sok (가지국수나물속)

Thalli (gametophyte) erect, usually epilithic, soft and mucilaginous, usually much and variously branched, attached by a small discoid holdfast. Axes terete, multiaxially organized with a medulla of fairly slender filaments and a cortex of subdichotomous filaments. Cortex loosely compacted, usually with distinctly larger and oblong terminal cells. Gametophyte monoecious or dioecious. Spermatangia clustered on spermatangial mother cells, arising from upper cortical cells. Carpo- gonial branches lateral on inner cortical cells, 3- or 4-celled, curved, with conical carpogonium and long trichogyne. Gonimoblast initials developed directely from fertilized carpogonium, divided subdichotously and repeatedly 3-5 times to form a globular compact carposporophyte. Carposporo- phyte producing ovoid carpospoangia on terminals of gonimoblast filaments, enveloped or sub- tended by sterile post-fertilization filaments arising from cells adjacent to the supporting cell. Tetrasporophyte minute, filamentous, with terminal cruciately divided tetrasporangia. Type species: Helminthocladia calvadosii (Lamouroux ex Duby) Setchell. 88 Algal Flora of Korea·Nemalian Red Algae

SPECIES 17 (1 in Korea). DISTRIBUTION: Worldwide, subtropical to tropical seas. KEY REFERENCES: Huisman and Womersley (1994), Huisman and Womersley (2006a). REMARKS: Helminthocladia is easily distinguished by the loosely compacted multiaxial axis, the soft and the mucilaginous larger terminal cells of cortical filaments, and the curved carpogonial branch. In Korea, two species of Helminthocladia, H. australis Harvey and H. yendoana Narita, were described with photographic specimens by Kang (1968). Comparing the two photographic specimens given by Kang (1968), however, it is very difficult to see a morphological difference between them except the abundance of branches in H. yendoana. In our observations, Helminthocladia australis has some morphological diffrences between male and female plants. The male plants have slender axes and many branches than female plants as like those of H. yendoana. Thus, we puta- tively identified the two reported species as a single species H. australis in this work.

29. Helminthocladia australis Harvey 1863: 39 (Figs. 63, 64). Keun-ga-ji-guk-su-na-mul (큰가지국수나물)

Harvey 1863. pl. 272. synop. xxxix. Womersley 1965: 470. f. 46-53. pl. 5. Kang 1966: 60; 1968: 191. pl. 37. f. 61. Huisman and Womersley 1994: 90. f. 20E, F, 23. Huisman and Womersley 2006a: 49. f. 20. Dawes and Mathieson 2008: 211. pl. 27. f. 19-21. Lee 2008: 213.

SYNONYM: Helminthora tumens J. Agardh 1890: 41. Lucas 1935: 215. Lucas and Perrin 1947: 133. Helminthocladia californica (J. Agardh) Kylin 1941: 6. Helminthocladia gracilis N.L. Gardner 1926: 206. pl. 15. Helminthocladia macrocephala Yamada 1941: 207. f. 10. pl. 47. Helminthocladia tumens (J. Agardh) J. Agardh 1899: 95. De Toni 1924: 84.

Thallii (gametophytes) epilithic, usually solitary, purple to yellowish red, 10-30 cm high, irregulary and generally profusely branched, attached by small discoidal holdfast. Axes terete, 5-10 mm in diameter below, 1-4 mm in diameter above, multiaxially organized with a medulla and a cortex. Medulla filamentous, loosely associated, bearing peripheral girdle filaments which producing radially arranged cortical filaments. Cells of medulla colourless, elongated cylindrical or irregular, 20-40 μm broad, 150-220 μm long. Cortical filaments 6-7 celled in female plants, 4-5 celled in male plants, loosly associated and subdichotomously branched with large oblong terminal cells. Cells of cortex conatin one plastid per cell, cylindrical Fig. 63. Distribution of Helminthocladia in below of 15-20 μm broad and 40-75 μm long, oblong australis. Nemaliophycidae: Nemaliales: Liagoraceae 89

ABC

DEFG

Fig. 64. Helminthocladia australis. A, B. Habit of female plants (A. Iho beach, Jeju; B. Chuja islet, Jeju); C. Detail of cortical cells of female plant in cross section view showing large oblong terminal cells; D. Cross section view of medulla and cortex of female plant; E. Carpogonial branch; F. Habit of male plant (Chuja islet, Jeju); G. Spermatangial clusters on terminals of cortical filaments. 90 Algal Flora of Korea·Nemalian Red Algae in terminal cells of 25-35 μm broad and 35-55 μm long. Spermatangia clustered on spermatangial mother cells, arising from upper cortical cells. Carpogonial branches lateral on inner cortical cells, 4-celled, curved, with conical carpogonium and long trichogyne. Gonimoblast and carposporophytes not observed in this work. Tetrasporophyte unknown.

TYPE: Fremantle, Western Australia (in Herb. Harvey, TCD). SEASONALITY: Probably observed spring to summer season in subtidal zone. DISTRIBUTION: Worldwide, subtropical to tropical seas. KOREA: Jukbyeon, Guryongpo (Kang 1966, 1968), Jeju (Lee and Lee 1989). SPECIMEN EXAMINED: KN7321 (Iho, Jeju: 6.vi.2009), KN6231 (Daeseri, Chujado: 23.vii.2007). REMARKS: Korean specimens are basically identical with the descriptions and illustrations of southern Austrailan specimens provided by Womersley (1994). It is necessary, however, for the further studies including DNA sequencing work to confirm the taxonomic entity of Japan and Korean materials. Fortunately, we collected male and female plants at Chuja islet in same habitat. The male and female plants look like very different species by the colour and gross morphology. The male plants are almost similar with the photograpies provided by Lee (2008). Therefore, we considered this species may be shown dimorphisms between male and female plants.

Genus Nemalion Duby 1830: 953. Guk-su-na-mul-sok (국수나물속)

Thalli (gametophytes) erect, usually epilithic, soft and mucilaginous, simple to subdichotomously branched, sometimes with laterals, attached by a small discoid holdfast. Axes terete to subterete, multiaxial with a medulla and cortex. Medulla colourless, composed of slender filaments, irregularly entwined and tightly compacted. Cortex consisted of loosely compacted and subdichotomously branched of anticlinal filaments, differentiated into an inner hyaline region and outer assimilatory region, usually tapered to apex without enlarging of terminal cells. Plastids large, single per cell of assimilatory filament, stellate with a conspicuous central pyrenoid. Gametophyte usually monoecious. Spermatangia lateral or in digitate clusters on outer cortical cells. Carpogonial branches usually 4-8-celled, straight, similar to immature cortical filaments, with only the distal 3 to 4 cells modified. Gonimoblast initials developed directely from fertilized carpogonium; first zygote division transverse, the upper cell segmenting obliquely or longitudinally to form a dense, globular carposporophyte. Sterile filaments usually present, developing from lower cells of carpogonial branch after fertilization, forming a slight involucre around the carposporophyte. Tetraspo- rophyte minute, filamentous, with cruciate tetrasporangia or monosporangia. Type species: Nemalion lubricum Duby [=N. helminthoides (Velley) Batters].

SPECIES ca. 7 (1 in Korea). DISTRIBUTION: Worldwide, subtropical to cool temperate seas. KEY REFERENCES: Womersley (1994), Huisman and Womersley (2006b). REMARKS: The generic type species, Nemalion helminthoides, was well studied in detail of its ultra- structure of post-fertilizational process by Ramm-Anderson and Wetherbee (1982). The life history Nemaliophycidae: Nemaliales: Liagoraceae 91 of Nemalion was confirmed by laboratory culture studies (Masuda and Horiguchi 1988; Cunning- ham and Guiry 1989). Only one species, N. vermiculare, was reported as a popular species in east coast of Korea, but the anatomical structures and life history of Korean materials have not been studied at present.

30. Nemalion vermiculare Suringar 1874: 91 (Figs. 65, 66). Cham-guk-su-na-mul (참국수나물)

Suringar 1874: 91. pl. 13. Okamura 1916: 28. pl. 158. f. 1-16; 1936: 413. f. 191. Kang 1966: 60; 1968: 193. pl. 38. f. 63. Yoshida 1998: 522. pl. 3-9. f. F-H.

SYNONYM: Nemalion helminthoides var. vermiculare (Suringar) C.K. Tseng 1983: 50, nom. illeg.

Thalli (gemetophytes) erect, epilithic, simple or sparingly branched, vermicular, very soft and slippery but durable, 10-20 cm long, 1.5-2 mm thick, deep vinose-red fading to yellowish in age, attached by dicoidal holdfast. Axes terete to subterete, multiaxial with a medulla and cortex. Medulla colourless, tightly compacted with entangled slender filaments, 400-500 μm in diameter. Cells of medulla elongated cylindrical, 4-5 μm broad, 50-150 μm long. Cortex consisted of two layers, loosely compated inner layer of 400-450 μm length and densely clustered assimilatory outer cortical layer of 200-250 μm length. Cells of assimilatory layer cylindrical to ellipsoidal, 9-12 μm broad, 18-30 μm long, constricted at septem, slightly tepered to apex, ending short haylin hair, contained a stellate plastid with a conspicuous central pyrenoid. Gameto- phyte monoecious. Spermatangia digitate clusters on terminal of outer cortical cells. Carpogonial branches arising terminal of cortical filament, straight, similar to immature cortical filaments. Carposporophyte spherical, enveloped loosely by involucral branches, 70-80 μm diameter. Tetrasporophyte minute, filamentous, with cruciate tetrasporangia or monosporangia (Masuda and Horiguchi 1988).

TYPE: Japan (?). SEASONALITY: Spring to summer season. DISTRIBUTION: Japan, Russia (Yoshida et al. 1990, Perestenko 1980). KOREA: Around all coasts. SPECIMEN EXAMINED: KU6743 (Imwon, Samcheok: 18.vii.2009). REMARKS: This species is a more plentiful in easten Fig. 65. Distribution of Nemalion vermi- coast of Korea on waved upper intertial zone. Korean culare. 92 Algal Flora of Korea·Nemalian Red Algae

AB CD

EF GHI

Fig. 66. Nemalion vermiculare. A. Habit of plants (Imwon, Samcheok); B. Cross section view; C. Detail of cross section view; D. Detail of assimilatory cotical cells showing a stellate plastid per cell with conspicuous cenral pyrenoid; E. Detail of cortical filaments with spermatangia; F. Mature carposporophyte; G. Carpogonial branch on terrninal of cortical cells (arrow); H. Divisions of fertilized carpogonium (arrow); I. Monoecious occurring together young carposporopyte (arrow head) and spermatangia (arrow). Nemaliophycidae: Nemaliales: Liagoraceae 93 materials are almost identical with descriptions and illustrations of Japanese specimens provided by Okamura (1916; pl. 158, f. 1-16). Okamura (1916) mentioned that the Japanese plants are dioecious; however, Korean materials are usually monoecious.

Acknowledgments: We thank Dr. Chen-Jung Kwon and Miss Sun-Mi Kim for help with collection and preparing of manuscripts. 94

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Index to Korean Names

ㄱ 숨긴붉은털 69 숨긴붉은털속 68 가는홍다발솔 51 가락말속 84 가시홍다발솔 56 ㅇ 가위손말과 71 가위손말속 82 애기가위손말 83 가지국수나물속 87 애기붉은솜 15 고리방사털 74 애기손붉은솜 13 고리방사털속 73 얼레기붉은솜 25 고비홍다발솔 54 여린두가닥바닷말 79 국수나물목 71 왕홍다발솔 64 국수나물속 90 융단자리풀 33 국수나물아강 9 굵은자루붉은솜 19 ㅈ

ㄷ 자리풀속 33 제주붉은솜 17 다발붉은솜 12 대양홍다발솔 61 두가닥바닷말속 74 ㅊ 두포자홍다발솔 37 참국수나물 91 천사붉은솜 22 ㅁ 청각홍다발솔 39

매듭붉은솜 30 민가락말 84 ㅋ

큰가지국수나물 88 ㅂ 큰홍다발솔 42

분홍국수말과 86 붉은솜과 10 ㅍ 붉은솜목 9 붉은솜속 11 팔손이풀목 67 버짐더부살이과 67 편생홍다발솔 44 비녀붉은솜 24 뾰족두가닥바닷말 76 ㅎ

ㅅ 해면손홍다발솔 58 홍다발솔과 36 상투붉은솜 28 홍다발솔목 36 송이홍다발솔 47 홍다발솔속 36 106

Index to Korean Names as Pronounced

A Ga-si-hong-da-bal-sol 56 Ga-wi-son-mal-gwa 71 Ae-gi-bul-geun-som 15 Ga-wi-son-mal-sok 82 Ae-gi-ga-wi-son-mal 83 Go-bi-hong-da-bal-sol 54 Ae-gi-son-bul-geun-som 13 Go-ri-bang-sa-teol 74 Go-ri-bang-sa-teol-sok 73 Guk-su-na-mul-a-gang 9 B Guk-su-na-mul-mok 71 Guk-su-na-mul-sok 90 Beo-jim-deo-bu-sal-i-gwa 67 Gul-geun-ja-ru-bul-geun-som 19 Bi-nyeo-bul-geun-som 24 Bul-geun-som-gwa 10 Bul-geun-som-mok 9 H Bul-geun-som-sok 11 Bun-hong-guk-su-mal-gwa 86 Hae-myeon-son-hong-da-bal-sol 58 Hong-da-bal-sol-gwa 36 Hong-da-bal-sol-mok 36 C Hong-da-bal-sol-sok 36

Cham-guk-su-na-mul 91 Cheon-sa-bul-geun-som 22 J Cheong-gak-hong-da-bal-sol 39 Ja-ri-pul-sok 33 Je-ju-bul-geun-som 17 D

Da-bal-bul-geun-som 12 K Dae-yang-hong-da-bal-sol 61 Du-ga-dak-ba-dat-mal-sok 74 Keun-ga-ji-guk-su-na-mul 88 Du-po-ja-hong-da-bal-sol 37 Keun-hong-da-bal-sol 42

E M

Eol-rae-gi-bul-geun-som 25 Mae-deub-bul-geun-som 30 Min-ga-rak-mal 84

G P Ga-neun-hong-da-bal-sol 51 Ga-ji-guk-su-na-mul-sok 87 Pal-son-i-pul-mok 67 Ga-rak-mal-sok 84 Ppyo-jok-du-ga-dak-ba-dat-mal 76 Index to Korean Names as Pronounced 107

Pyeon-saeng-hong-da-bal-sol 44 W

Wang-hong-da-bal-sol 64 S

Sang-tu-bul-geun-som 28 Y Song-i-hong-da-bal-sol 47 Sum-gin-bul-geun-teol 69 Yeo-rin-du-ga-dak-ba-dat-mal 79 Sum-gin-bul-geun-teol-sok 68 Yung-dan-ja-ri-pul 33 108

Index to Scientific Names

A Ahnfeltiophycidae 4 Audouinella 9, 10, 11, 33, 36 Acrochaetiaceae 10, 11, 12, 36, 71 bispora 37 Acrochaetiales 4, 6, 9, 10, 36, 67, 68, 71 botryocarpa 44 Acrochaetium 9, 10, 11, 12, 28, 33 canariensis 12 bisporum 37 catenulata 13 canariense 11, 12, 13, 15, 17 codicola 39 catenulatum 19, 21, 22 codii 42, 44 codicola 39 compta 44 codii 42 crassipes 19 collopodum 21 daviesii 47 compactum 21 densa 15 comptum 44 dictyotae 51 crassipes 19, 21 elegans 54, 56 daviesii 12, 47 endozoica 60 densum 11, 13, 15, 17, 23 hermannii 12 dictyotae 51 hyalosiphoniae 56 effusum 60 infestans 58 elegans 54 microscopica 19 humile 30 moniliformis 19 hyalosiphoniae 56 pacifica 61 infestans 58 phacelorhiza 44 inkyui 11, 17, 19, 30 purpurea 33 iyengarii 61 rhizophora 44 microscopicum 11, 13, 15, 19, 21, 22 sancti-thomae 22 moniliforme 13, 15, 21, 22 saviana 67 pacificum 61 scapae 24 parvulum 32 secundata 25 plumosum 32 subimmersa 69 rhipidandrum 32 terminalis 28 sancti-thomae 11, 22, 24 thuretii 64 scapae 11, 24, 25 virgatula 32 secundatum 11, 12, 25, 28, 32 seriatum 46 spongicolum 60 B subimmersum 69 terminale 11, 19, 28, 30 Balbiania 4, 10 thuretii 64 Balbianiales 4, 6 virgatulum 11, 28, 30, 32 Ballia 4 Actinotrichia 72, 73 Balliales 4, 6 fragilis 73, 74 Batrachosperamaceae 71 rigida 73, 74 Batrachospermales 4, 6, 71 Index to Scientific Names 109

Bonnemaisoniaceae 71 densum 15 Bonnemaisoniales 71 microscopicum 19 Byssus 10 moniliforme 19 purpurea 33 secundatum 25 virgatulum 32 Colaconema 6, 10, 12, 36, 37 C bispora 37 bonnemaisoniae 36 Callithamnion codii 42 codicola 19, 37, 39, 42, 44, 46 Callithamnion daviesii 47 codii 37, 42, 44 Callithamnion daviesii var. secundatum 25 compta 37, 44, 46 Callithamnion microscopicum 19 daviesii 37, 47, 54, 56, 58 Callithamnion purpureum 33 dictyotae 37, 51, 54 Callithamnion rothii 33 elegans 37, 54, 56 Callithamnion virgatulum 30 hyalosiphoniae 37, 56, 58 Ceramiaceae 6 infestans 37, 58, 60 Ceramium 64 membranaceum 6 daviesii 47 pacificum 37, 61 rothii 33 secundatum 25 secundatum 25 subimmersum 6 Chaetangiaceae 71 thuretii 37, 64, 67 Chantransia 10 Colaconemataceae 12, 36 bispora 37 Colaconematales 4, 6, 9, 36 catenulata 13 Conferva daviesii 47 codii 42 Conferva purpurea 33 compta 44 Conferva violacea 33 corymbifera 64 Corallina cylindrica 84 crassipes 19 Corallina rugosa 83 dictyotae 51 Corallinales 4, 6 efflorescens var. thuretii 64 Corallinophycidae 4, 6 endozoica 60 infestans 58 microscopica 19 D moniliformis 19, 21 pacifica 61 Dermonenmataceae 71 sancti-thomae 22 Dichotomaria 72, 73, 74, 75, 82 secundata 25 apiculata 75, 76, 79, 82 thuretii 64, 67 falcata 75, 76, 79 thuretii var. agama 67 marginata 75, 76 thuretii var. amphicarpa 67 virgatula 32 Chantransiella 10 F Chromastrum 10 canariense 12 Florideophyceae 4 catenulatum 13 Fucus fragilis 74 crassipes 19 110 Algal Flora of Korea·Nemalian Red Algae

G Helminthora tumens 88 Hindenbrandiophycidae 4 Galaxaura 72, 73, 75, 82, 84, 86 apiculata 72, 73, 75, 76 clavigera 72 K collabens 83 conglutinata 84 Kylinia 10 cylindrica 84 catenulata 13 elongata 83 crassipes 19 falcata 72, 73, 75, 79 infestans 58 fastigiata 84 microscopica 19 filamentosa 72 moniliformis 19 glabriusculosa 83 scapae 24 hystrix 72, 73, 75, 76, 79 secundata 25 lapidescens 83 terminalis 28 marginata 72, 73, 75 virgatula 32 oblongata 86 pacifica 83 papillata 75 L pilosula 72, 73, 76 rigida 74 Lemanaceae 71 rudis 72, 83 Leptosarca 67 rugosa 72, 73, 75, 82, 83 Liagora 87 schimperi 85 pinnata 46 squalida 83 Liagoraceae 71, 86, 87 subfruticulosa 83 Liagorophila 10 subverticillata 73 Galaxauraceae 71, 72 Gloiophloea 72 M Grania 10 Meiodiscaceae 9, 67

H N Halossaccion 67 Halysium cylindricum 85 Naccariaceae 71 Helminthocladia 64, 87, 88 Nemaliaceae 71 australis 88 Nemaliales 4, 6, 67, 71 californica 88 Nemalion 87, 90, 91 calvadosii 87 helminthoides 90 gracilis 88 helminthoides var. vermiculare 91 macrocephala 88 lubricum 90 tumens 88 vermiculare 87, 91 yendoana 88 Nemaliophycidae 4, 6, 9 Helminthocladiaceae 71, 87 Nothogenia 72 Index to Scientific Names 111

P virgatulum 32 Rhododraparnaldia 4 Palmaria 67 Rhodogorgonales 4, 6 palmata 67 Rhodonematella 6, 33, 68 Palmariaceae 6, 67 subimmersa 67, 68, 69, 70 Palmariales 4, 6, 9, 33, 36, 67, 68 Rhodophysema 68 Pseudoacrochaetium 10 georgii 67 Pseudochantransia 10 Rhodophysemataceae 33, 67, 68 Rhodothamniella 4, 10 codii 42 R Rhodothamniellaceae 9, 67 Rhodymeniales 67 Rhodachlya 6 Rhodymeniophycidae 4 Rhodachlyales 4, 6, 9 Rubrointrusa 6 Rhodochorton 9, 10, 11, 12, 33 catenulatum 13 codicola 39, 42 S codii 42 daviesii 47 Scinaia 72 densum 15, 17 Scinaiaceae 71, 72 dictyotae 51 elegans 54, 61 floridulum 69 T hyalosiphoniae 56, 58 infestans 58 Thamnidium 10 intermedium 35 intermedium 35 islandicum 35 rothii 35 magnificum 51, 54, 61 Thorea 6 microscopicum 19 Thoreaceae 71 moniliforme 19 Thoreales 4, 6 pacificum 61 Trentepohlia 10 parasiticum 35 daviesii 47 penetrale 60 purpurea 35 plumosum 61 rothii 35 purpureum 12, 33, 35, 69 virgatula 32 rhizoideum 39, 42 Tricleocarpa 72, 84 rothii 35 cylindrica 84, 85, 86 sancti-thomae 22 oblongata 86 subimmersum 33, 67, 68, 69 tenue 35 terminale 28 W thuretii 64 variabile 61 Whidbeyella 72

Russia

JB GN JN

JJ South Sea Algal Flora of Korea

Volume 4, Number 2 Rhodophyta: Florideophyceae: Nemaliophycidae: Acrochaetiales, Colaconematales, Palmariales, Nemaliales Nemalian Red Algae

Flora and Fauna of Korea

National Institute of Biological Resources Ministry of Environment National Institute of Biological Resources Ministry of Environment