Flora and Fauna of Korea

Transcription

1 Algal Flora of Korea Volume 3, Number 2 Chrysophyta: Bacillariophyceae: Pennales: Araphidineae: Diatomaceae Freshwater Diatoms II Flora and Fauna of Korea National Institute of Biological Resources Ministry of Environment

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

3 Algal Flora of Korea Volume 3, Number 2 Chrysophyta: Bacillariophyceae: Pennales: Araphidineae: Diatomaceae Freshwater Diatoms II

4 Acknowledgement This work was derived from the Flora and Fauna of Korea project supported by the National Institute of Biological Resources, the Ministry of Environment, Korea.

5 Algal Flora of Korea Volume 3, Number 2 Chrysophyta: Bacillariophyceae: Pennales: Araphidineae: Diatomaceae Freshwater Diatoms II Gyeongje Joh, Jung Ho Lee 1, Kyung Lee 2 and Sook-Kyung Yoon 2 School of Environmental Science and Engineering, Inje University Gimhae, Gyeongsangnam-do 1 Department of Biology Education, Daegu University Gyeongsan, Gyeongsangbuk-do 2 Department of Life Science, The Catholic University of Korea Bucheon, Gyeonggi-do Flora and Fauna of Korea National Institute of Biological Resources Ministry of Environment

6 Copyright 2010 by the National Institute of Biological Resources Published by the National Institute of Biological Resources Environmental Research Complex, Gyeongseo-dong, Seo-gu Incheon , Republic of Korea 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 : Government Publications Registration Number Printed by Junghaengsa, Inc. in Korea on acid-free paper Publisher : Chong-chun Kim Editor : Sook Shin Reviewer : Hans U. Dahms Project Staff : Youn-Bong Ku, Chang-Hwan Bae, Moon-Soo Lim Published on April 23, 2010 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.

7 Preface In the wake of the Convention on Biological Diversity (CBD), which recognized national sovereignty over indigenous biological and genetic resources when adopted in 1992, countries around the world have been putting their best foot forward in unearthing raw biological materials reckoned as one of the crucial resources upon which national competitiveness depends to a great extent in the 21 st century. Being well aware of the priority of securing and managing biological resources, the National Institute of Biological Resources (NIBR) under the Korean Ministry of Environment decided to issue the Flora and Fauna of Korea in an attempt to attain systematic management and comprehensive conservation of biological resources at the national level. Endowed with diverse landscapes involving a wide range of topographic conditions, Korea is acclaimed as one of the nations with high levels of biological diversity. Purporting to establish a thorough record on national indigenous species, the NIBR embarked on the publication of the Flora and Fauna of Korea in Korean and English detailing those species inhabiting the Korean peninsula in Our dedication to research during the past three years led by a group of professionals in the field of systematics finally came to fruition with issuance of the first of a kind monograph for Korean animals, fungi and algae encompassing approximately 1,037 species in 158 families belonging to 9 phyla. It is my firm belief that this very first national the Flora and Fauna of Korea is indeed the culmination of our persevering scientific research efforts aimed at deepening our understanding on native species and acutely identifying Korean biota. It will not only serve as an important initiative for sustainable biodiversity conservation but also a catalyst for rational and far-sighted use of biological resources. I would like to extend my utmost gratitude to the team of over 29 professors and associated experts headed by Prof. Sook Shin of Sahmyook University for their unsparing efforts in producing this groundbreaking work. I earnestly hope that on-going publication of the Flora and Fauna of Korea initiated by the Ministry of Environment will significantly contribute to unveiling all Korean native species estimated up to 100,000 and expanding wise utilization of beneficial indigenous resources. Chong-chun Kim, Ph. D. President NIBR

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9 Bryozoa: Gymnolaemata: Cheilostmata: Inovicellata, Malacostega, Flustrina, Ascophora 1 Contents Asterionella, Diatoma, Meridion, Opephora, Tabellaria List of Taxa 7 Introduction 8 Materials and Methods 11 Taxonomic Notes Asterionella formosa Hassall Asterionella formosa var. gracillima (Hantzsch) Grunow Diatoma hyemalis (Roth) Heiberg Diatoma mesodon (Ehrenberg) Kützing Diatoma moniliformis Kützing Diatoma tenuis Agardh Diatoma vulgaris Bory Meridion circulare (Greville) Agardh Meridion circulare var. constrictum (Ralfs) Van Heurck Opephora olsenii Möller Tabellaria fenestrata (Lyngbye) Kützing Tabellaria flocculosa (Roth) Kützing 42 Literature Cited 45 Fragilaria, Hannaea, Pseudostaurosira, Punctastriata, Staurosira, Staurosirella List of Taxa 53 Introduction 54 Materials and Methods 55 Taxonomic Notes Fragilaria bidens Heiberg Fragilaria capitellata (Grunow) Petersen Fragilaria capucina Desmazieres Fragilaria capucina var. radians (Kützing) Lange-Bertalot Fragilaria crotonensis Kitton Fragilaria famelica (Kützing) Lange-Bertalot Fragilaria gracilis Ǿstrup Fragilaria mesolepta Rabenhorst 69

10 2 Algal Flora of Korea Freshwater Diatoms II 9. Fragilaria parva Tuji and D. M. Williams Fragilaria rumpens Kützing Fragilaria vaucheriae (Kützing) Petersen Hannaea arcus var. recta (Cleve) M. Idei Hannaea arcus var. subarcus (Iwahashi) Lee Pseudostaurosira brevistriata (Grunow) Williams and Round Punctastriata linearis Williams and Round Staurosira construens Ehrenberg Staurosira construens var. binodis (Ehrenberg) Hamilton Staurosira elliptica (Schumann) Williams and Round Staurosira venter (Ehrenberg) H. Kobayasi Staurosirella pinnata (Ehrenberg) Williams and Round 90 Literature Cited 92 Staurosira Staurosirella Synedra List of Taxa 99 Introduction 100 Materials and Methods 101 Taxonomic Notes Synedra acus Kützing Synedra delicatissima W. Smith Synedra delicatissima var. angustissima Grunow in Van Heurck Synedra goulardi Brébisson ex Cleve and Grunow Synedra pulchella (Ralfs ex Kützing) Kützing Synedra pulchella var. lacerata Hustedt Synedra rumpens var. meneghiniana Grunow in Van Heurck Synedra socia Wallace Synedra tabulata var. obtusa (Pantocsek) Hustedt Synedra tenera W. Smith Synedra tenuissima (Kützing) Kützing Synedra ulna (Nitzsch) Ehrenberg Synedra ulna var. amphirhynchus (Ehrenberg) Grunow Synedra ulna var. contracta Østrup Synedra ulna var. danica (Kützing) Van Heurck Synedra ulna var. obtusa Van Heurck Synedra ulna var. oxyrhynchus (Kützing) Van Heurck Synedra ulna var. oxyrhynchus f. mediocontracta (Forti) Hustedt Synedra ulna var. ramesi (Héribaud) Hustedt Synedra ulna var. spathulifera (Grunow in Van Heurck) Grunow in Van Heurck 138

11 Bryozoa: Gymnolaemata: Cheilostmata: InContentsovicellata, Malacostega, Flustrina, Ascophora 3 Literature Cited 140 Index to Korean Names 149 Index to Korean Names as Pronounced 150 Index to Scientific Names 152

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13 Bryozoa: Gymnolaemata: Cheilostmata: Inovicellata, Malacostega, Flustrina, Ascophora 5 Asterionella, Diatoma, Meridion, Opephora, Tabellaria Gyeongje Joh

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15 Bryozoa: Gymnolaemata: Cheilostmata: Inovicellata, Malacostega, Flustrina, Ascophora 7 List of Taxa Class Bacillariophyceae Order Pennales Suborder Araphidineae Family Diatomaceae Dumortier 1822 Genus Asterionella Hassall 1850 Asterionella formosa Hassall 1850 Asterionella formosa var. gracillima (Hantzsch) Grunow 1881 Genus Diatoma Bory 1824 Diatoma hyemalis (Roth) Heiberg 1863 Diatoma mesodon (Ehrenberg) Kützing 1844 Diatoma moniliformis Kützing 1833 Diatoma tenuis Agardh 1812 Diatoma vulgaris Bory 1824 Genus Meridion Agardh 1824 Meridion circulare (Greville) Agardh 1831 Meridion circulare var. constrictum (Ralfs) Van Heurck 1881 Genus Opephora Petit 1888 Opephora olsenii Möller 1950 Genus Tabellaria Ehrenberg 1840 ex Kützing 1844 Tabellaria fenestrata (Lyngbye) Kützing 1844 Tabellaria flocculosa (Roth) Kützing 1844

16 8 Introduction 1. Flora of the family Diatomaceae Diatoms belonging to family Diatomaceae are characterized by the rapheless valve as pennate forms. The group diatoms are widely and abundantly distributed in streams, rivers and lakes of freshwaters, while their dominances are relatively low in marine environments Approximately 40 genera and 1,700 species in the family have been reported in the world as the database, and approximately 400 species among them are currently accepted as valid taxa (Algaebase 2009). Among them, genera containing only 1 or 2 species are 25 taxa. The genera ordered serially in the number of species are as follows: Synedra (139), Fragilaria (108), Diatoma (34), Asterionella (17), Staurosira (12), etc. In Korea, the 24 genera and 164 species of araphid pennate diatoms have been recorded according to the literature survey by Lee (1995), KNCCN (1996) and Lee (1997). Among them, seven genera and 107 species (Asterionella 2, Diatoma 13, Meridion 2, Fragilaria 32, Synedra 52, Tabellaria 4, Tetracyclus 2 species) from freshwaters, and other 17 genera and 57 species from coastal and marine waters. Floristic studies in Korea have followed up the classification of Hustedt (1930, 1959). However, the system of Krammer and Lange-Bertalot (1991) is applied to the diatom flora in freshwaters, they are summarized as about 67 species (Asterionella 1, Diatoma 11, Meridion 2, Fragilaria 48, Synedra 1, Tabellaria 3, Tetracyclus 1 species). In Korea, taxonomic research of araphid pennate diatoms was conducted on the 19 Synedra species by Lee and Yoon (2001). In their studies, the Synedra taxa are differentiated into 5 types with morphological features and this is the unique taxonomic study on the familian diatoms. Generic and specific discrimination on araphid diatoms mainly have relied on the feature of valve morphology such as the presence or magnitude of axial and central area, striae, spines, pore fields and rimoportulae and, the forms of colony and plastids. Colonial forms of the araphid diatoms are very important in traditional classification. Among them, diatoms united into ribbon-like or linear colonies are simply assigned to Fragilaria, solitary, fan-shaped or rosette-like colonies to Synedra, and cells joined at their angles to the other genera. The araphid pennates show a variety of forms in the colonial morphology and their ecological dominances in freshwaters are great as plankton and periphyton. 2. Classification of the Diatomaceae The diatomoid diatoms are ubiquitous algae with great morphological and ecological diversity. Especially, generic circumscriptions between Fragilaria and Synedra have long been doubtful by many authors, and to keep them as independent genera has long been questioned. The microillustrations by light and electron microscopy supported the similarity or combinations of the two genera to reveal morphological variations within one genus (Round 1984; Poulin et al. 1986; Williams and Round 1987; Lange-Bertalot 1989). Hustedt (1930) and Patrick and Reimer (1966) already pointed out the indistinctness of the two genera. F. Hustedt discussed the problems to distinguish the isolated Fragilaria valves from those of Synedra. A similar problem has arisen in identification of Fragilaria and Opephora (Morales 2002). Lange-Bertalot (1980) combined the freshwater species of subgenus Synedra (Eusynedra) with Fragilaria and consequently transferred about 30 Syndera taxa to Fragilaria. In additions, the

17 Bryozoa: Gymnolaemata: Cheilostmata: IIntroductionnovicellata, Malacostega, Flustrina, Ascophora 9 taxonomic positions of the genus Fragilaria and Synedra have changed in the last three decades with the observation of the ultrastructure by SEM. Many species belonging to the genus Synedra have been separated, and placed under many different genera in addition to a newly circumscribed Synedra by Williams and Round (1986) and Round et al. (1990). The ultrastructures of frustules were examined and studied as follows; the striation pattern of valve faces, the structure of apical pore field, the presence and absence of rimoportulae, and girdle structure. The range of character variations were made to erect the following 5 genera. Catacombas (=Catacombus) Williams and Round 1986 (type: Navicula gaillonii Bory 1827) (marine) Hyalosynedra Williams and Round 1986 (type: Synedra laevigata Grunow 1877) (marine) Tabularia (Kützing) Williams and Round 1986 (type: Synedra barbatula Kützing 1844) (marine) Ctenophora (Grunow) Williams and Round 1986 (type: Synedra pulchella Ralfs ex Kützing 1844) Neosynedra Williams and Round 1986 (type: Synedra provincialis Grunow 1877) (marine) The similar taxonomic situations were applied to Fragilaria by Williams and Round (1987). In additions of the Synedra characters, the type of linking spines was considered important in revision of the Fragilaria. The genus Fragilaria was differentiated into the following six genera. The original description was recorded in December, 1987, however, the journal was actually published in February in Fragilariforma (Ralfs) Williams and Round 1988 (type: Fragilaria virescens Ralf 1843) - Pseudostaurosira Williams and Round 1987 (type: Fragilaria brevistriata Grunow in Van Heurck ) - Punctastriata Williams and Round 1987 (type: Punctastriata linearis Williams and Round 1987) - Stauroforma Flower et al (type: Fragilaria exiguiformis Lange-Bertalot 1996) - Staurosira (Ehrenberg) Williams and Round 1987 (type: Fragilaria construens (Ehrenberg) Grunow 1862) - Staurosirella Williams and Round 1987 (type: Fragilaria lapponica Grunow in Van Heurck ) Krammer and Lange-Bertalot (1991) consequently divided Fragilaria into five subgenera [Fragilaria, Alterasynedra Lange-Bertalot, Ctenophora (Grunow) Lange-Bertalot, Tabularia (Kützing) Lange- Bertalot and Staurosira (Ehrenberg) Lange-Bertalot)] in their micrographs. Interrelationships within araphid pennate diatoms are not yet well known and araphid diatom taxonomy would be expected to progress further than in the past decades. Three taxonomic systems are currently applied to the classification of the family Diatomaceae. They are the traditional system deriving from Hustedt (1930, 1959) and Patrick and Reimer (1966), the recombinational system proposed by Lange-Bertalot (1989) and Krammer and Lange-Bertalot (1991), and the natural system propounded by Williams and Round (1987) and Round et al. (1990). Three classifications of araphid diatoms were summarized as Table 1 and system 1 by F. Hustedt or system 2 by K. Krammer and H. Lange-Bertalot is applied to the generic level of this edition.

18 10 Algal Flora of Korea Freshwater Diatoms II Table 1. Three classification systems for genera of araphid diatoms [adopted from Kingston (2002)] System 1 (Hustedt 1930, 1959; Patrick and Reimer 1966) System 2 System 3 (Krammer and Lange-Bertalot 1991) (Williams and Round 1987; Round et al. 1990) 1. Tetracyclus Ralfs 1. Tetracyclus Ralfs 1. Tetracyclus Ralfs, transferred to family Tabellariaceae 2. Diatoma Bory 2. Diatoma Bory 2. Diatoma Bory 3. Meridion Agardh 3. Meridion Agardh 3. Meridion Agardh 4. Asterionella Hassall 4. Asterionella Hassall 4. Asterionella Hassall (freshwater) 5. Asterionellopsis Round (marine) 5. Tabellaria Ehrenberg 5. Tabellaria Ehrenberg 6. Tabellaria Ehrenberg, transferred to family Tabellariaceae 7. Oxyneis Round, transferred to family Tabellariaceae 6. Synedra Ehrenberg 6. Synedra Ehrenberg 8. Synedra Ehrenberg 9. Catacombus Williams and Round (marine) 10. Hyalosynedra Williams and Round (marine) 11. Tabularia (Kützing) Williams and Round (mainly marine) 12. Ctenophora (Grunow) Williams and Round 13. Neosynedra Williams and Round (mainly marine) 7. Fragilaria Lyngbye 7. Fragilaria Lyngbye 14. Fragilaria Lyngbye Subgenus - Fragilaria Lange-Bertalot 15. Fragilariforma (Ralfs) Williams and Round Subgenus - Alterasynedra Lange- 16. Pseudostaurosira Williams and Round Bertalot Subgenus - Ctenophora (Grunow) 17. Punctastriata Williams and Round Lange-Bertalot Subgenus - Tabellaria (Kützing) 18. Stauroforma Flower et al. Lange-Bertalot Subgenus - Stauorsira (Ehrenberg) 19. Staurosira (Ehrenberg) Williams and Lange-Bertalot Round 20. Staurosirella Williams and Round 8. Opephora Petit 8. Opephora Petit 21. Opephora Petit (marine) 22. Martyana Round (freshwater) 9. Hannaea Patrick 9. Hannaea Patrick 22. Hannaea Patrick 10. Centronella Voigt 10. Centronella Voigt 23. Centronella Voigt 11. Frankophila Lange-Bertalot

19 Bryozoa: Gymnolaemata: Cheilostmata: Inovicellata, Malacostega, Flustrina, Ascophora 11 Materials and Methods 1. Sampling and cleaning Diatom materials were sampled from planktons, periphytic or attached collection (stones, rocks, aquatic macrophytes or even filamentous algae), as well as benthic one in Korean freshwaters. They should be always studied in their natural, living state under light microscopy (LM). Samples were cleaned in the boiling bath of nitric acid, sulphuric acid or hydrogen peroxide to remove organic matters (APHA 1995). The pretreated materials were repeatedly washed by centrifugation or settling overnight and subsequent dilution with distilled water. The cleaned frustules are stored in ethanol after the final rinse and ready for mounting. Permanent diatom specimens were made using mounting media such as Pleurax or Hyrax resin. Diatoms were analyzed under compound photomicroscope (Zeiss, Olympus or Nikon) with high resolution DIC (differential interference contrast) optics. Additionally, the cleaned and dried frustules were mounted on aluminum stubs, and coated with gold to examine the high resolution of the morphology under the scanning electron microscopy (SEM). Images were captured with a monochromatic camera equipped to the microscope and a computer. A counts of 500 diatom frustules in minimum are required for quantitative analysis of floristic study. 2. Description of diatom flora All araphid pennate diatoms encountered in this collections were described and illustrated for taxonomic and iconographic studies in Korea. We mainly followed the taxonomic system of Hustedt (1930, 1959), Patrick and Reimer (1966), and Krammer and Lange-Bertalot (1991) to identify the diatoms. We refer and consult many other literatures to expand high diversity of freshwater diatoms in Korean peninsula. The morphological variations in a group or assemblage were also reevaluated for the autoecological and taxonomic aspects. To provide synonyms and basionyms of diatom species, we refer to Catalogue of Diatom Names of academic website, (CAS 2009), which has been held by California Academy of Sciences (CAS) for world life s diversity. The informations about diatom flora over the world came from the global database ( Algaebase 2009). Catalogues or check-lists authorized by Lee (1995), KNCCN (1996) and Lee (1997) were source documents to sum up the diatom flora reported or described in Korea.

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21 13 Taxonomic Notes Family Diatomaceae Dumortier Mak-dol-mal-gwa ( ) Among pennate diatoms, this family is composed of the diatoms having no raphe on valve faces and being non-motile. This group contains all elongated diatoms which are pennate forms with no true raphe along the apical axis. Valves have transapical striae arranged in both sides of an apical axis. The valve has a pore field at each apex. Some genera have one or two rimoportulae at the pole or in the central area. The diatoms of this group do not have complex cingula. Some genera have also spines arranged in valve margins and united to make colonies. Some taxa or diatoms have septa in girdle band. The common features of these groups are to have linear valve, one rimoportula near one of both apices, one apical pore or thick silicification at each apex, uniseriate striae composed of poroid areolae and numerous narrow intercalary bands. GENERA AND SPECIES: There are 40 genera and approximately 1,700 species in the world and, among them, about 400 speces are currently accepted as valid taxa (Algaebase 2009). Genera containing only 1 or 2 species are 25 taxa. The flora of this group diatom in Korean freshwaters are the 7 genera (Asterionella, Diatoma, Meridion, Fragilaria, Synedra, Tabellaria, Tetracyclus) and 70 species according to the summarization of literature studies (Lee 1995; KNCCN 1996; Lee 1997). This monographic edition include eight generic flora of diatoms, the seven freshwater taxa and one brackish taxon, Opephora. DISTRIBUTION: Araphid diatoms are partly planktonic, and largely attached on stone (epilithic), sediment (benthic), sand (epipsammic), mud (epipelic), and aquatic macrophytes (epiphytic). The diatoms of this group are usually colony and, as plankton, periphyton or bottom diatoms, more rich and abundant than other family ones in freshwaters. The major parts of this group are mostly restricted to freshwaters and minor parts to the marine environments. KEY REFERENCE: Poulin et al. (1986), Williams (1986), Krammer and Lange-Bertalot (1991), Williams and Round (1986, 1987), Hasle and Syvertsen (1996). Key to the genera of family Diatomaceae 1. Septum present in the intercalary band 2 Septum absent in the intercalary band 3 2. Transverse costae distinct in valve view Tetracyclus Transverse costae absent Tabellaria 3. Transverse costae in valve view 4 Transverse costae absent 5 4. Cells clavate in valve view to be asymmetry to the transapical axis. Circular fan-shaped colonies Meridion Cells fusiform in valve view and isopolar. Often zigzag or ribbon shaped colony

22 14 Algal Flora of Korea Freshwater Diatoms II Diatoma 5. Cells elongated with inflated ends and somewhat heteropolar in valve view. Cells forming stellate colonies in girdle view Asterionella having not above-mentioned characteristics 6 6. Valve entirely asymmetry to transapical axis, brackish diatom Opephora Valves isopolar 7 7. Cells in ribbon-like colonies. Cells usually μm long Fragilaria Cells solitary or in clusters. Cells usually μm long Synedra (Krammer and Lange-Bertalot 1991) REMARKS: The classification of araphid and rod-shaped diatoms is the most problematic due to their simple forms and characteristics. They are comparatively less studied than the other group of diatoms. All aspects of araphid groups have led early authors to clump the genera (Round et al. 1990). Based on valve faces and the related characteristics, it is often very difficult to differentiate Fragilalria, Opephora and Synedra under light microscopy. Electron microscopic studies and other advanced researches probably show a revision of this diatom group. Genus Asterionella Hassall 1850: pl. 2. Byeol-dol-mal-sok ( ) Valves elongate, more or less linear and swollen the apices. Apical pore fields present at both ends. Spines arranged along margins of the valve. Several girdle bands, each with one or two rows of pores. Striae very fine to be parallel, perpendicular to apical axis, and in 10 μm. From 4 to 32 elongated cells joining at base to form stellate colonies. Plastids numerous, small and plate-like, in a cell. Transversely oriented rimoportula at either end of a valve opening externally via a pore, somewhat larger than areolae. Lectotype: Asterionella formosa Hassall 1850: 10. pl. 2. f. 5. SPECIES: Among 54 species reported up to date, 17 species are currently accepted as taxonomically valid taxa (Algaebase 2009). Two taxa, A. formosa and A. formosa var. gracillima, have been recorded in freshwaters of Korea. DISTRIBUTION: In many boreal and temperate lakes, Asterionella is a common and important component of the phytoplankton in lakes, rivers and the other freshwaters. ECOLOGY: These are typical and popular planktons in open water such as lowland lakes and ponds, and rivers and streams having low concentrations of dissolved salts. Some populations are particularly susceptible to infestations by fungus chytrids. KEY REFERENCE: Patrick and Reimer (1966), Körner (1970), Round et al. (1990), Krammer and Lange- Bertalot (1991). Key to the species of genus Asterionella 1. The pole inflated with triangle-form in the girdle view A. formosa The pole slightly or scarcely inflated in the girdle view A. ralfsii

23 Araphidineae: Diatomaceae: Asterionella 15 REMARKS: By proposal by Round et al. (1990), marine species formerly included in Asterionella were transferred to new genus Asterionellopsis (type: Asterionella glacialis Castracane) and Bleakeleya (type: Asterionella bleakeleyi var. notata Grunow). Newly defined Asterionella is exclusive in freshwater. 1. Asterionella formosa Hassall 1850: 10 (Figs. 1, 2). Hustedt 1930: 147. f Patrick and Reimer 1966: 159. pl. 9. f. 1. Krammer and Lange-Bertalot 1991: 103. pl f. 1. SYNONYM: Asterionella gracillima var. formosa (Hassall) Wislouch 1921: 107. Cells forming star or rosette shaped colonies. Valves and girdle views narrow-linear, apices of valve capitately widened. Capitate apex (foot pole) much larger than that of the other apex (head pole) in both valve and girdle view. Pseudoraphe narrow, central area absent. Transapical striae very delicate in 10 μm, often obscure on valves. Valve μm, in length and 1 3 μm in breadth. Colonies typically comprising of 4 or 8 cells, occasionally spiraling slightly. Some varieties have been discussed by authors. Huber-Pestalozzi (1942) recorded the curved forms as a variety (A. formosa var. acaroides Lemmermann), and described it as pathological forms of the type species. TYPE: Locality-Thames at Brentford, England, drinking water of Grand Junction Company in SEASONALITY: The species usually exhibits a maximum growth in spring and less in autumn. They are the most frequent and typical planktonic algae in lakes or streams of temperate regions. Sometimes, they grow well in the season when nutrient concentrations are relatively high and water temperature is low. The species was the most abundant component in surface water under icecover in the fjord regions of Canada (Chassé and Côté 1991) and dominated the diatom populations with winter maximum in an oligo-mesotrophic lake of Northwest Spain (Negro et al. 2000). DISTRIBUTION: A. formosa is a common and well studied planktonic diatoms which may be found in all kinds of freshwaters. In many temperate regions, A. formosa is the most important plankton in many boreal and temperate lakes and, on the other hands, is subdominant by the preoccupation of the centric diatoms among phytoplankton. As euryhaline and eutrophic algae, the species is a remarkable pennate diatom taxa in plankton distribution. The population variability was early studied in Windermere Lake of England and its ecology was examined by Lund (1949). Similar seasonal changes in abundance were known from lakes and reservoirs in other many regions - the Rhine River, the Meuse River, some Mediterranean reservoirs of France in Europe (Ibelings et al. 1998; Bertrand et al. 2003), Canadian fjord regions, and lakes of North America (Chassé and Côté 1991). This is one of the top ten diatoms which have been recorded frequently during the recent century in Finland lakes (Lepistö et al. 2006). The species dominates the diatom populations of the mesotrophic, eutrophic, and hypereutrophic waters (Negro et al. 2000), and can grow even under more extremely alpine oligotrophy (Bozniak and Kennedy 1968; Ilmavirta 1975; Spaulding et al. 1993). In the Southern Hemisphere, this dominated highly the lake plankton in New Zealand (Duthie and Stout 1986). Though this is a typical plankton, the species has found

24 16 Algal Flora of Korea Freshwater Diatoms II G F H I J A B C D 10 μm E L K Fig. 1. Asterionella formosa. A-F. the valve of cells; G-K. the girdle view of cells ( 2,000); L. a starshaped colony ( 400) (LM); A-D, H-K. the images of LM. as an important epiphyte on reed plants (Albay and Akcaalan 2003). The species has increased in some lakes since the 1990s to show paleolimnological evidence in USA (Eilers et al. 2004). On the other hands, it is relatively decreased since 1955 in the River Meuse of Germany (Ibelings et al. 1998). KOREA: Asterionella population, A. formosa and A. formosa var. gracillima - is the most important one in the frequency and abundance of the phytoplankton in Korean freshwaters. A. formosa was recorded in Han River (Chung and Kim 1970), abundant in the brackish area of the Nakdong River before the estuarine dam construction (Chung et al. 1987). In the Han River, the species was the most important plankton throughout the year from 1977 to 2004 (Jung et al. 2003; Lee and Jung 2004). Asterionella population had a peak in spring and October to April next year (Jung et al. 2003).

25 Araphidineae: Diatomaceae: Asterionella 17 In addition, the species showed significant dominance in lakes; Lake Angye and Unmun (Lee et al. 2002), Lake Andong, Yeongcheon, and Hapcheon (Kim and Lee 1996), Lake Paldang (Kim 1996, 1998), Lake Imha (Kim et al. 1997), and Lake Gachang (Park and Kim 2003). In Lake Paldang, the species ranked as a subdominant among planktons in May. Besides eutrophic rivers and streams, the species was observed in a mountainous stream of Mountain Baekun (Chung et al. 1986). In contrast to the river waterbody, the species exhibited a very low frequency in the Haman lowland wetland (Chung and Noh 1987). SPECIMEN EXAMINED: (Planktonic diatoms collected from agricultural reservoirs of Gimhae regions: iv. 1993). ECOLOGY: This is a representative in planktonic algae of mesotrophic and eutrophic waters, and has generally been described as an indicator of eutrophic conditions (Hutchinson 1967; Reynolds 1984). The colony size of A. formosa is affected by phosphorus and silica limitation (Tilman et al. 1976), and water temperature (Hayakawa et al. 2004). The number of cells per colony increased from 6 8 up to under Fig. 2. Distribution of Asterionella formosa. the condition of silica depletion, but most colonies have only 2 4 cells under phosphorus limitation (Tilman et al. 1976). Morphometric changes of A. formosa colony could be a useful indicator of phytoplankton nutrient status. In addition to cell numbers, rapid cell reduction was reported in cultures and natural populations of A. formosa, and there was an increase in cell numbers per colony (Kling 1993). In the fossil records of a Canadian Precambrian Shield lake, A. formosa increased concurrently with European settlement ca in North America (Clerk et al. 2000). On the other hand, this can grow well in the oligotrophic natural freshwaters. The species can additionally tolerate the significant saline conditions of estuarine waters. REMARKS: Hustedt (1930) and Patrick and Reimer (1966) differentiated A. formosa var. gracillima (Hantsch) Grunow [=A. gracillima (Hantzsch) Heiberg] from A. formosa, according to the identity of two apex morphologies. Although some authors have traditionally classified the two taxa into a single species, A. formosa, A. gracillima was considered as a variation in the type species (Hutchinson 1967; Körner 1970; Krammer and Lange-Berlalot 1991). 2. Asterionella formosa var. gracillima (Hantzsch) Grunow 1881: pl. 51 (Figs. 3 6). Hustedt 1930: 147. f Patrick and Reimer 1966: 160. pl. 9. f. 4.

26 18 Algal Flora of Korea Freshwater Diatoms II BASIONYM: Diatoma gracillimum Hantzsch in Rabenhorst 1861: 1104b. f. a. SYNONYM: Asterionella gracillima (Hantzsch in Rabenhorst) Heiberg 1863: 68. pl. 6. f. 19. Valve and girdle view similar with the type species except for equal capitate ends. End portions of the valve strongly widened on both basal (attaching) and apical poles of valve. Transapical striae very fine in 10 μm. Valve μm in length and 2 3 μm in breadth. The variety can be distinguished from type species by equal sizes of capitate ends of valve. TYPE: Locality-Elbufer im grossen Gehege bei Dresden, November, SEASONALITY: This variety occurs more abundantly in spring and winter than in the warmer season. D C 10 μm E F G H I J K A B M N O Fig. 3. Asterionella formosa var. gracillima. A-C. the girdle view of cells ( 2,000); D-K. the valve of cells ( 2,000); M-O. the colonial forms of 8, 16 and 32 cells ( 400) (LM); B-H. DIC images of LM.

27 Araphidineae: Diatomaceae: Asterionella 19 A B D C E F Fig. 4. Asterionella formosa var. gracillima. A D. girdle views of cells, foot and head poles; E, F. a rimoportular pore in the inside of valves (SEM).

28 20 Algal Flora of Korea Freshwater Diatoms II 10 6 Asterionella formosa var. gracillima Nakdong River (Gupo Bridge) Asterionella (cells/ml) Seonakdong River Fig. 5. The seasonal variations of Asterionella formosa var. gracillima (grey area) and total phytoplankton (solid lines) in the Nakdong and the Seonakdong River. The taxon showed clear seasonality from late autumn (October) to spring (May) of next year in the downstreams of the Nakdogn River, South Korea (Fig. 5). Water temperature during the Asterionella blooms ranged from 10 C to 20 C in the Nakdong River. DISTRIBUTION: This is widely distributed as plankton in boreal freshwaters. A. formosa var. gracillima was a dominant plankton in April and May in Lake Oshima-Ohnuma of Japan (Takano et al. 2001). KOREA: Among two neighboring populations, A. formosa var. gracillima was more abundant and persistent than A. formosa in Korean rivers. The gracillima taxon was very abundant in the brackish regions of the Nakdong River before the dam construction in the river mouth (Chung et al. 1987). Its abundance decreased after the estuarine dam construction to record around 1,500 cells/ml in maximum. A. formosa was subdominant in November in the midstream (Waegwan) of the Nakdong River (Lee et al. 2002). Variable forms of Asterionella were recorded as A. formosa var. gracillima in the Han River (Chung and Kim 1970). SPECIMEN EXAMINED: (Planktonic diatoms collected from the Seonakdong River and the lower reaches of the Nakdong River: iv.1993).

29 Araphidineae: Diatomaceae: Diatoma 21 ECOLOGY: The variety is also eutrophic like the species and would have similar ecological status. It seems to prefer cool to cold water. REMARKS: A. formosa and its variety did not exhibit a different geographical distribution, and are found together in same locality. A. formosa var. gracillima was accepted as A. formosa by many authors (Krammer and Lange-Bertalot 1991). The detailed reexaminations are required to discern the distributional patterns or their taxonomic determinations. Two sibling taxa, formosa and gracillima of Asterionella, may not be differentiated in the course of many phytoplankton studies in Korean freshwaters and gracillima variety has been likely reported as A. formosa. Fig. 6. Distribution of Asterionella formosa var. gracillima. Genus Diatoma Bory 1824: 461. Mak-dol-mal-sok ( ) Valve faces of cell elliptical to elongate. Apices of the valve variable, from rounded to swollen. Striae fine, transverse at centre, becoming radiate towards apices. Narrow axial area with no central area. Transapical costae clearly present, with a single rimoportula as a thickened area at one end of the valve. The outline of girdle view rectangular and girdle bands with two rows of pores. Cells joining either along valve face to form ribbon-shaped colonies, or at one corner to form zigzag or stellate colonies. Many small discoid or slightly elongate plastids in a cell. Lectotype: Diatoma rigidum A.P. de Candolle 1805: 49. SPECIES: Among 223 species reported up to date, the 34 ones are currently accepted as taxonomically valid taxa (Algaebase 2009). Eleven Diatoma taxa were recorded in South Korea until They are Diatoma anceps (Ehrenberg) Kirchner, D. elongatum Agardh, D. hyemalis (Roth) Heiberg, D. mesodon (Ehrenberg) Kützing, D. moniliformis Kützing, D. tenuis Agardh, and D. vulgaris Bory and its four varieties. DISTRIBUTION: As periphyton or plankton, they are commonly distributed in streams and lakes of Korean freshwaters, however, rare in brackish waters. ECOLOGY: As periphytic diatom assemblages, Diatoma species was dominant in habitats with higher N:P ratios in the regions of the Great Lakes (Marks and Power 2001). Diatoma population is one of the epiphyte assemblages attached on filamentous green algae, Cladophora species (Marks

30 22 Algal Flora of Korea Freshwater Diatoms II and Power 2001). KEY REFERENCE: Hustedt (1930), Patrick and Reimer (1966), Round et al. (1990), Williams (1990), Krammer and Lange-Bertalot (1991). Key to the species of genus Diatoma 1. Transverse costae more than 6 in 10 μm and cells united to be zigzag colony 2 Transverse costae less than 5 in 10 μm and cells united with valve face to make colony 5 2. Rimoportula present at the both valve poles, transapical striae very fine and invisible. Valve spindle-shaped or linear without prodruding end D. moniliformis A rimoportula in a pole of the valve 3 3. Valve 5 μm or less wide, narrow and long D. tenuis Cells strong and more than 6 μm wide 4 4. Valve margin linear with capitate end D. ehrenbergii Valve broad, linear or elliptical-lanceolate, generally without capitate or protruding end D. vulgaris 5. Valve linear with distinctly capitate end D. anceps Valve end not capitate and protruding end 6 6. Valve very strong and longer more than 40 μm, and striae distinct in 10 μm D. hyemalis Valve more delicate and less than 40 μm, and striae very fine in 10 μm D. mesodon (Krammer and Lange-Bertalot 1991) REMARKS: The genera Diatoma and Meridion are traditionally distinguished from the other araphid diatoms by the possession of highly silicified costae or transapical ribs, and by lack of septa on girdle bands (Williams 1990). They differ each other in the form of valves, isopolarity of Diatoma and heteropolarity of Meridion. 3. Diatoma hyemalis (Roth) Heiberg 1863: 58 (Figs. 7 9). Hustedt 1930: 129. f Patrick and Reimer 1966: 107. pl. 2. f. 7. Krammer and Lange-Bertalot 1991: 99. pl. 97. f. 6. SYNONYM: Conferva hyemalis Roth 1800: 506. pl. 1. f. 22. Odontidium hyemalis Kützing 1844: 44. pl. 17. f. 1. Diatoma hyemale (Roth) Heiberg 1863: 58. Cells in rectangular girdle view and intercalary bands present. Valves connected to form more and less long and ribbon-shaped colony. Valve linear to lanceolate and isopolar with straight or convex margins and rounded ends of the valve. Sometimes, the ends of the valve wedge-shaped. Transapical costae very wide and perpendicular to transapical axis, sometimes irregularly or obliquely distributed. Costae strong, 2 4 in 10 μm, μm. Axial area very wide and striae in 10 μm. Valve (12) μm in length and 7 13 μm in breadth. TYPE: Locality - In piscinis prope Schoenebeck non procul vegesack primo observavi, postea

31 Araphidineae: Diatomaceae: Diatoma 23 passim in fossis. SEASONALITY: The species is frequently found in cold season or water body. DISTRIBUTION: The species seems to prefer cool waters and common in alpine and mountain streams of the northern regions. The species was common and abundant in a small high mountain pond of Poland (Kownacki et al. 2006) and the most frequent and abundant diatom in alpine waters A 10 μm B C D E K F H G M L J I Fig. 7. Diatoma hyemalis. A I. valve faces of cells ( 2,000); J, K. girdle views of cells ( 2,000); L, M. the ribbon-shaped forms of colonies ( 400, LM).

32 24 Algal Flora of Korea Freshwater Diatoms II C D B 10 μm A E Fig. 8. Diatoma hyemalis. A D. valve faces of cells ( 2,000); E. a girdle view of cell ( 2,000, LM); A E. DIC images of LM. of Swiss mountains (Robinson et al. 2010). KOREA: In Korea, this was frequently reported in the rivers and lakes (Lake Paldang, Uiam and Paro) of the Han River, the Nam River and Lake Juam in the other river system, and some mountain streams (Chung and Kim 1970; Chung and Koh 1974; Chung et al. 1987). D. hyemalis was exclusively dominant as attached algae around the Jeolmul Spring of Mountain Halla, Jeju Island, in November SPECIMEN EXAMINED: (Periphytic diatoms attached on wood substrates around the Jeolmul Spring in Jeju Island were collected in: xi.2009). ECOLOGY: The species is distributed in northern or alpine area and prefers relatively low conductivity and reduced alkalinity in freshwaters. REMARKS: The species is easily distinguished from D. mesodon by coarseness or breadth of the costae and the width of axial area on the valve. The length of the species is longer than 30 μm in the monographe of Krammer and Lange-Bertalot (1991). However, more smaller sized diatoms were abundantly observed in Fig. 9. Distribution of Diatoma hyemalis.

33 Araphidineae: Diatomaceae: Diatoma 25 the specimens collected from the Jeolmul Spring. The lengths of specimens sampled from the Jeolmul Spring ranged from 12 μm to 50 μm. 4. Diatoma mesodon (Ehrenberg) Kützing 1844: 47 (Figs. 10, 11). Hustedt 1930: 129. f Patrick and Reimer 1966: 108. pl. 2. f. 8. Krammer and Lange-Bertalot 1991: 100. pl. 98. f. 7. SYNONYM: Fragilaria mesodon Ehrenberg 1839: pl. 2. f. 9. Odontidium mesodon (Kützing) Kützing 1849: 12. Diatoma hiemale var. mesodon (Ehrenberg) Grunow in Van Heurck 1881: pl. 51. f. 3. Valves linear-elliptical to lanceolate with blunt apices. The central portion of the valve crossed by below 5 strong costae, 3 6 in 10 μm. Transapical striae in 10 μm. Striae indistinct, in 10 μm. Girdle view square to rectangular, often deeper than valve length and numerous intercalary bands distinct. Cells united with valve faces to form compact chains usually in ribbon-shaped colony. Valves μm in length and 6 14 μm in breadth. D. mesodon is the most smaller species among genus Diatoma. TYPE: Bacillarien: 47. pl. 17. f. XIII (1844) (L). In Bächen des thüringer Waldes, des Harzes, Auch bei Berlin: Ehrenberg. SEASONALITY: The genus is frequently found in cold season or water body. DISTRIBUTION: The species was distributed widely in the alpine areas and mountains of Europe as benthic or attached diatoms. This is a cosmopolitan benthos and often dominant in the mountain or alpine streams, in central Colorado streams (Vavilova and Lewis 1999), in forest basin of Willamette River in Oregon (Carpenter and Waite 2000), in streams in a high elevation catchment of the Swiss Alps (Robinson and Kawecka 2005) and in alpine-zone streams of the Tatra National Park in Poland (Kawecka and Robinson 2008). They occurred abundantly in ponds, bogs, and running waters in the plains. D. mesodon dominated the sampling sites of New Zealand rivers in winter (Biggs and Price 1987). KOREA: The frequency of this species is relatively low in Korean freshwaters. The species was recorded in the Yongneup Alpine Swamp (Chung and Kim 1987), in streams originated from Mountain Jiri and among epilithic diatom assemblages in some shallow streams. Fig. 10. Distribution of Diatoma mesodon.

34 26 Algal Flora of Korea Freshwater Diatoms II SPECIMEN EXAMINED: (Epilithic diatoms collected from the Imcheon Stream around Jiri Mountain, the tributary of the Gyeongho River: 22.xi.2007). ECOLOGY: The species is distributed in northern or alpine areas, usually found in the plain and A C D E F G B 10 μm H I J K L M N O Fig. 11. Diatoma mesodon. A G. valve faces of cells ( 2,000); H M. girdle views of cells ( 2,000, LM); N, O. the valve and girdle view of cells (SEM); F I. DIC images of LM.

35 Araphidineae: Diatomaceae: Diatoma 27 occasionally predominant in some flowing waters. This is cold-water species and an oligotrophic diatom typically in unpolluted streams. 5. Diatoma moniliformis Kützing 1833: 580 (Figs. 12, 13). Krammer and Lange-Bertalot 1991: 98. pl. 96. f. 11. SYNONYM: Diatoma tenuis var. moniliformis Kützing 1833: 52. f. 60. Cells in narrow rectangular girdle view and intercalry bands present, but indistinct. Cells connected to form zig-zag colony. Valve elliptical, elliptical-lanceolate to lanceolate with convex margins. The ends of valve broadly rounded to wedge-shaped, or slightly protracted. Transapical costae thin and dense, 7 12 in 10 μm. Apical axial area very narrow. Transapical striae invisible and unresolved under light microscopy, in 10 μm under electron microscopy. Valve 8 40 μm in length and μm in breadth. TYPE: Linnaea 8(5): 580 (1833). SEASONALITY: The species was abundant in spring in the running waters (Ziemann et al. 2001). A B C D E F G 10 μm H I J K L M N O P Q R S T Fig. 12. Diatoma moniliformis. A J. valve faces of cells ( 2,000); K O. girdle views of cell ( 2,000); P T. valve faces of cell ( 2,000, LM); A K. DIC images of LM.

36 28 Algal Flora of Korea Freshwater Diatoms II DISTRIBUTION: The species was one of the dominating diatoms in benthic diatom communities in three Estonian rivers (Vilbaste, 2001) and in salt-loaded reaches of the River Wipper in Germany (Ziemann et al. 2001). Epiphytes of the Tisza River, the Hungarian Reaches of the Danube River was dominated by D. moniliformis and other important species (Szabó et al. 2005). This had high abundance in waters contaminated with mining discharge across the continental United States (Potapova and Charles 2003). Especially, this was found in the digestive apparatus of barbell (Barbus meridionalis) from different parts of the Shkumbini River, Albania (Miho et al. 2005). KOREA: In Korea, this was only reported in the lower reaches of the Nakdong River in the past (Lee et al. 1995). The species was observed abundantly in the Eogok Stream of Danyang, a tributary of the Namhan River in January The freshwater of the Eogok Stream was alkaline to show 216 mg CaCO 3 /l of alkalinity and 8.9 of ph value, respectively. SPECIMEN EXAMINED: (Epilithic diatoms collected from the Eogok Stream, Danyang were examined for the D. moniliformis which was important in the stream: i.2010). Fig. 13. Distribution of Diatoma moniliformis. ECOLOGY: D. moniliformis favors the comparatively high ion contents as halphilous diatom or saltindicating taxon (Ziemann et al. 2001; Potapova and Charles 2003). In additions, this is known to be abundant in brackish waters, below 1,000 μs/cm (Potapova and Charles 2003). 6. Diatoma tenuis Agardh 1812: 15 (Figs. 14, 15). Hustedt 1930: 128. f Patrick and Reimer 1966: 108. pl. 2. f. 5. Krammer and Lange-Bertalot 1991: 97. pl. 96. f. 1. Genkal 2004: 24. f. 1. SYNONYM: Diatoma tenue Agardh 1824: 4. Diatoma elongatum (Lyngbye) Agardh 1824: 4. Odontidium elongatum var. tenuis (Agardh) Patrick 1939: 5. Diatoma elongatum subsp. tenuis (Agardh) Skabichevskii 1960: 235. f. 78. Odontidium tenue (Agardh) Kuntze 1898: 418. Diatoma elongatum var. tenuis (tenue) (Agardh) Van Heurck : 160. Candollella tenuis (Agardh) Gaillon 1833: 34. Colony typically zig-zag in form. Valves linear to linear-lanceolate and the capitate valve end broader than the rest of the valve. Slightly convex margins in valve views, being above 10 in the

37 Araphidineae: Diatomaceae: Diatoma 29 ratio of length to width. A row of spines along edge of valve face, leading to join sibling cells. Cells secrete mucus at one corner via apical pore fields joining neighbor cells in zigzag or rarely in stellate colonies. Cells elongated in girdle view, and intercalrary bands indistinct. Transapical costae conspicuous, 6 10 in 10 μm and striae delicate, over 40 in 10 μm. Number of striae in 10 μm and valve width most stable features, while valve length and the number of costae in 10 μm most labile (Genkal 2004). Valves μm in length, 3 5 μm in breadth. TYPE: Locality - In aquis dulcibus Scandinaviae. D A B C 10 μm E F G H I J K L M N O P Q R Fig. 14. Diatoma tenuis. A K. valve views of cells ( 2,000); L R. girdle views of cells ( 2,000, LM); A C, L N. DIC images of LM.

38 30 Algal Flora of Korea Freshwater Diatoms II SEASONALITY: The species mainly occurs during spring in lakes and ponds. DISTRIBUTION: The species is widely distributed and common, often bloomed massively in lentic waters and also found in the slightly brackish waters. This is a common diatom over the world. The species was found as both the periphyton and plankton in shallow brackish lakes of England (Moss 1981), and as the first dominant algae in some lakes of New Zealand (Duthie and Stout 1986). The araphid diatoms such as Diatoma elongatum, Meridion circulare, and Tabellaria flocculosa were dominant in plexiglass attachment in a small stream Near Abisko, Swedish Lapland (Muller-Haeckel and Håkansson 1978). In South Finnland, Lake Vesijärvi was loaded by sewage from the City of Lahti for 60 years until 1976 when the discharge was diverted. Paleolimnological analyses revealed Asterionella formosa and D. elongatum reached their maximum occurrence between when the hypolimnion of the lake was aerated artificially (Liukkonen et Fig. 15. Distribution of Diatoma tenuis. al. 1993) D. elongatum has largely developed in Lake Tjeukemeer (a Dutch fen lake) 5 6 weeks later than those of the centric diatoms such as Aulacoseira and unicellular centric diatoms, owing to intensification of agriculture surrounding the lakes (Moed and Hoogveld 1975). During survey of phytoplankton among 153 lakes, rivers, and temporary pools in Canada s Northwest Territories lying above a latitude of about 55 degrees, Asterionella formosa and D. tenue var. elongatum were common in many southern lakes. KOREA: The species is rare in frequency and lower in abundance in Korean freshwaters. It was recorded in lower reaches of the Namhan and Bukhan River under the name of D. elongatum Agardh (Chung and Kim 1970; Chung and Lee 1978; Lee and Cho 1994) and also in the lower parts of the Han River. SPECIMEN EXAMINED: (Epipiphytic diatoms collected from Lake Hyeongeum and Daein near the estuary of the Bukcheong-Namdaecheon Stream in North Korea: 27.vii.1998). ECOLOGY: The species is often observed in lakes or standing waters with relatively high conductivity. The species is classified as halophilic taxon (Ziemann et al. 2001). REMARKS: This species is frequently recorded as D. elongatum Agardh. However, the valid name of this taxon is D. tenuis Agardh, due to the nomenclatural priority. 7. Diatoma vulgaris Bory 1824: 461 (Figs. 16, 17). Hustedt 1930: 127. f Patrick and Reimer 1966: 109. pl. 2. f. 9. Krammer and Lange-Bertalot 1991: 95. pl. 93. f. 1.

39 Araphidineae: Diatomaceae: Diatoma 31 SYNONYM: Diatoma vulgare Bory 1831: pl. 20. f. 1. Bacillaria vulgaris (Bory) Ehrenberg 1836: 53. Diatoma vulgare var. productum Grunow 1862: 363. Odontidium vulgare (Bory) Pfitzer 1871: 121. pl. 6. f. 20. Neodiatoma vulgare (vulgaris) (Bory) Kuntze 1891: 906. A B C 10 μm D E F G H I Fig. 16. Diatoma vulgaris. A G. valve views of cells ( 2,000); H. a girdle view of cell ( 2,000); I. the zig-zag form of a colony ( 400, LM); F, G. DIC images of LM.

40 32 Algal Flora of Korea Freshwater Diatoms II Cells rectangular in girdle view, and sometimes its side slightly convex. Cells united to form zigzag colonies by mucilage pads at corners. Zigzag colony often in long length. Valves broadly elliptical to lanceolate with bluntly rounded to rostrate ends. Rimoportula distinct near one of apices of valve. No central area in the middle parts of valve. Tranverse costae crossing valve face. Striae fine, hard to resolve under light microscopy, although narrow axial area being usually distinct. Valves 8 75 μm in length, 7 18 μm in breadth. TYPE: Uncertain, probably France. SEASONALITY: The diatoms seem to prefer cool and mainly occur during low-flow period in winter. However, their occurring seasons are variable in spring and autumn, even in summer (Ziemann et al. 2001). DISTRIBUTION: The species is found in brackish water up to 4 NaCl salinity and often in water with fairly high nutrient concentrations. It is commonly Fig. 17. Distribution of Diatoma vulgaris. abundant on stones as periphytics or benthics. As benthic diatoms attached on the gravel in streams, epilithic colonization during winter of low flow was characterized by D. vulgaris and its relatives in the River Garonne in France (Eulin and Cohu 1998), in a Pampean river of Argentina (Martínez de Fabricius et al. 2003), and in the St. Lawrence River, Quebec in Canada (Vis et al. 1998). D. vulgaris is a representative epiphyte on Cladophora filaments (Marks and Power 2001). In addition, it comprised the major proportion of total abundance and biomass of the phytoplankton in littoral zone of an urban Lake Jeziorak Ma ly, Poland (Zębek 2007). KOREA: The species was observed in high frequency as epipilthic algal assemblages in many streams (Chung and Kim 1991). This has a higher frequency than any other species of Diatoma in Korean freshwaters. It was observed as plankton in the lower, middle reaches of the Han River (Chung et al. 1965; Chung et al. 1968; Chung and Kim 1970; Chung 1972), the Imjin River above 10% relative abundance (Lee and Yoon 2002), however, rare in lakes. They were observed in the Yongneup Alpine Swamp (Chung and Kim 1987). SPECIMEN EXAMINED: (Epilithic diatoms sampled from streams of the Suambo Spa in Chungju: 19.i.2010), (plankton collected from the estuary of the Namdaecheon Stream in Uljin: 2.i.2008). ECOLOGY: The species is frequent in slightly flowing waters and often observed in fairly eutrophic waters. The species was classified as differential taxon indicating β-mesosaprobic and oligohalobic (indifferent) taxon (Lange-Bertalot 1979). It is often found within epilithic and epiphytic assemblages, or as tychoplankton in both lakes and rivers. In colonization experiments using the artificial substrata in the River Danube near Budapest, the first colonizer was D. vulgaris which adhered on apical pad after a few hours (Ács et al. 2000). And further, after 24 hours the algal community was already diverse with a total of 35 species. REMARKS: D. vulgaris has a wide range of variations and has been divided into some varieties and forma (Fukushima et al. 1986). Krammer and Lange-Bertalot (1991) classified the species into seven

41 Araphidineae: Diatomaceae: Meridion 33 morphotypic groups in their monographs according to the shapes of valves, valve apices or other morphological features. This species has relatively a great deal of synonyms. Genus Meridion Agardh 1824: 2. Bu-chae-dol-mal-sok ( ) In valve view, cell outline either wedge-shaped or linear with rounded, sub-capitate to capitate apices. Thickened costae cross valve at irregular intervals. Striae fine and barely resolvable under light microscopy. Axial area very narrow and no central area. Small spines along margins of valve face. A rimoportula towards head pole and a poorly-defined pore field towards foot pole. Cells rectangular to wedge-shaped in girdle view, joined by valve faces to form straight to fan-shaped colonies. The large septa easily seen in the girdle view. Many discoid plastids in cell. Lectotype: Meridion vernale Agardh 1824: 2. SPECIES: Among 17 species reported up to date, 7 taxa are currently accepted as taxonomically valid taxa (Algaebase 2009). Two taxa, M. circulare and M. circulare var. constrictum, were recorded in Korea. DISTRIBUTION: The species are often abundant in slow flowing waters and ditches. ECOLOGY: This is alkaliphilous taxa in freshwater. KEY REFERENCE: Hustedt (1930), Williams (1990), Round et al. (1990), Krammer and Lange-Bertalot (1991), Rhode et al. (2001). REMARKS: The diatoms are typical freshwater periphyton or plankton. Traditionally distinction between Diatoma and Meridion was by the shape of valves, the former is isopolar, whilst the latter is heteropolar. However, Williams (1985) has transferred Diatoma anceps to Meridion, so as to admit the unreasonableness of new combination. 8. Meridion circulare (Greville) Agardh 1831: 40 (Figs. 18, 19). Hustedt 1930: 130. f Patrick and Reimer 1966: 113. pl. 2. f. 15. Krammer and Lange-Bertalot 1991: 101. pl f. 1. SYNONYM: Echinella circularis Greville 1823: 35. Exilaria circularia (Greville) Greville 1827: 37. Exilaria circularis (Greville) Agardh 1831: 40. Cells wedge-shaped with heteropoles having a narrow foot apex and wide head apex. Cells united each other with valves to form fan-shaped colonies. Transverse costae strongly developed, entirely or partly crossing valve face in 3 5 in 10 μm. Axial area of valve face narrow, transpical striae in 10 μm. Many discoid plastids in cytoplasm. Cells μm in length, 4 8 μm in breadth.

42 34 Algal Flora of Korea Freshwater Diatoms II TYPE: Consp. Crit. Diat.: 40 (1831). Locality-Rivulet near Dumbryden Quarries, England. SEASONALITY: The species generally achieves maximum abundances in spring. M. circulare was dominated early in spring in a shaded, woodland stream of eastern Tennessee (Steinman and Parker 1990), in winter in the Llobregat River of Spain (Tomas and Sabater 1985), and in three alpine streams B C F G A 10 μm H I J K D E L M N O P Fig. 18. Meridion circulare. A K. valve views of cells; L N. girdle views of cells ( 2,000, LM); O, P. the outer and inner face of the valve (SEM); A E. DIC images of LM.

43 Araphidineae: Diatomaceae: Meridion 35 in Kosciusko National Park of Australia (Chapman and Simmons 1990). DISTRIBUTION: As more benthic or periphytic diatom, the species shows a wide distribution in freshwaters of the world. M. circulare was the most common taxon on epilithic substrates as a group to make 61 95% of the diatom assemblages in streams surrounding Lake Sophia of Cornwalls Island, Canada (Antoniades and Douglas 2002). This was the widespread species from soft bottom sediments of 46 streams in the lowlands of West Estonia (Vilbaste and Truu 2003). Araphid diatoms including M. circulare were dominant in plexiglass attachment in the Njakojokk stream near Abisko, Sweden (Muller-Haeckel and Håkansson 1978). A bloom of benthic algae occurred following the destruction of zoobenthic fauna in a mountain stream of Japan (Yasuno et al. 1982). This is seldom in the tropical regions. KOREA: The species occurs in high frequency among the streams of Korean freshwaters and common as epilithic diatom assemblages in inland streams. It Fig. 19. Distribution of Meridion circulare. had rare frequency and low abundance in lakes and, for example, was observed once in course of the phytoplankton survey in Lake Imha (Kim et al. 1997). The diatoms exhibited very low frequency in the Haman lowland wetland (Chung and Noh 1987). SPECIMEN EXAMINED: (Benthic diatoms collected from the Sandeulneup Bog of Mountain Jaeyak : 25.x.2006). ECOLOGY: This species prefers the cool and flowing waters, where it attaches to stones and plants. The species is common in springs and brooks, and usually considered to be rheobiont. M. circulare is often observed as macroscopic colonies in streams. The brown mucilaginous colonies were found to attach on plants and other substrates (Krejci and Lowe 1987). As alkaliphilous diatoms, they are not tolerant of low ph or organic pollution. 9. Meridion circulare var. constrictum (Ralfs) Van Heurck 1881: 51 (Figs. 20, 21). Hustedt 1930: 131. f Patrick and Reimer 1966: 114. pl. 2. f. 16. Krammer and Lange-Bertalot 1991: 102. pl f. 8. BASIONYM: Meridion constrictum (constricta) Ralfs 1843: 458. pl. 18. f. 2. SYNONYM: Meridion circulare f. constricta (Ralfs) Cleve-Euler 1932: 15. Meridion constrictum (constricta) Ralfs 1843: 458. pl. 18. f. 2. Meridion circulare var. constrictum (Ralfs) Brun 1880: 128.

44 36 Algal Flora of Korea Freshwater Diatoms II Cells wedge-shaped with narrower foot-pole and wider head-pole. Valve with capitate foot-pole and sharp constriction on neck. Cell walls very thick and strong. Transapical costae 3 5 in 10 μm, striae in 10 μm. Cells μm in length, 4 8 μm in breadth. TYPE: Diat. Alpes: 128. (1880). SEASONALITY: The variety has the same seasonality in its distribution as the species. DISTRIBUTION: This variety is commonly observed in freshwater, especially in flowing waters and occasionally blooms. The taxon shows the same distributional patterns as the species. It is not so frequent as the species. E A B C D F G H I J K L M N O 10 μm Fig. 20. Meridion circulare var. constrictum. A I. valve views of cells; J O. girdle views of cells ( 2,000, LM); A D, J L. DIC images of LM.

45 Araphidineae: Diatomaceae: Opephora 37 KOREA: The variety was observed in the Han River and its tributaries, the Namhan and Bukhan River, Lake Paldang, and in the lower reaches of the Nakdong River (Chung et al. 1968; Chung and Lee 1984; Lee et al. 1995). SPECIMEN EXAMINED: (Periphytic diatoms collected from the Sandeulneup Bog of Mountain Jaeyak: 25.x. 2006). ECOLOGY: This is an alkaliphilous taxon, and the ecology and habitat characteristics are same as the species. Fig. 21. Distribution of Meridion circulare var. constrictum. Genus Opephora Petit 1888: 130. Mong-chi-dol-mal-sok ( ) Cells narrowly rectangular, not trapezoid in girdle view. Valve heteropolar like wedge, club and clavate forms to be asymmetrical to transverse axes. The apices of valves rounded. Striae strongly broad and present along margins of valve. The genus can be differentiated with neighboring taxa by valve shapes and the structure of striae. Lectotype: Opephora pacifica (Grunow) Petit SPECIES: Among 22 species reported up to date, 10 taxa are currently accepted as taxonomically valid taxa (Algaebase 2009). Opephora has been usually recorded in coastal areas in Korea, however, its dominance and frequency were not high. The species was less frequently reported in Korean fresh and brakish water, 2 species (O. olsenii and O. martyi Héribaud) in total. DISTRIBUTION: These diatom groups are important components in marine and brackish waters as periphyton attached to sand grains or other substrates, aquatic plants. ECOLOGY: Opephora diatoms are commonly observed from sediments of diverse coastal habitats. KEY REFERENCE: Sundbäck (1987), Round et al. (1990), Krammer and Lange-Bertalot (1991), Sabbe and Vyverman (1995). REMARKS: These diatoms are too small to clarify the diagnostic characteristics by light microscopy. Although Opephora was diagnosed as an expansion of Petit s original description, it has been

46 38 Algal Flora of Korea Freshwater Diatoms II considered, as a complex group sharing a combination of morphological features (Morales 2002). The existing Opephora is suggested to be transferred to other genera or new taxa by some authors through cummulative resolutions of electron microscopy. There is considerable confusion as to the boundary of the genus or species identity (Round et al. 1990). 10. Opephora olsenii Möller 1950: 197 (Figs. 22, 23). Krammer and Lange-Bertalot 1991: 166. pl f. 9. Sabbe and Vyverman 1995: 241. f. 13. SYNONYM: Fragilaria pacifica Grunow 1862: 59. pl. 5/8. f. 19. Opephora pacifica (Grunow) Petit 1888: 131. Cells rectangular to weakly wedge-shaped or trapezoidal with rounded corners in girdle view. Cells solitary or connected to form simple colony with a few cells. Valve ovate to clavate, slightly heteropolar with rounded apices, nearly showing elliptical outlines although smaller diatoms often mostly isopolar. Axial area linearly narrow. Striae on valve face coarse, and their densities variable, 6 14 in 10 μm, arranged to be parallel. Striae arranged alternately throughout the valve. Valve 7 60 μm in length, μm in breadth. Two plastids plate-like, parietal in cytoplasm. TYPE: Folia Geogr. Dan. 3: 197. f. 4 (1950) (E). Locality-Zealand: Praestrø Fjord, Denmark. SEASONALITY: The maximum number of Opephora diatoms are usually observed in spring on sediment. DISTRIBUTION: The species showed a wide distribution in littoral zone of the coastal and brackish waters. This was frequently observed in brakish or marine areas, as the epipsammic or epiphytic diatoms in the brackish water (Sunbäck 1987), in a sandy tidal flat of eastern North Sea, as the abundant in Pack Bay of Poland, Southern Baltic Sea (Witkowski 2007), as the common in the Chesapeake Bay and its tidal estuaries (Marshall et al. 2005). On the other hand, the species was made assemblages in the epilithon of a spring-fed stream in Texas (Sherwood and Sheath 1999) and at high elevation regions (1,600 2,800 m) in the streams throughout the Lahontan Basin, California (Blinn and Herbst 2003). KOREA: The species was observed with very low frequency in the estuary of the Gwangcheon Stream in Uljin (Lee et al. 1994) and the Nakdong River. SPECIMEN EXAMINED: (Epipsammic diatoms collected from the estuary of the Bukcheong- 10 μm A B C D Fig. 22. A C. Opephora olsenii ( 2,000, LM).

47 Araphidineae: Diatomaceae: Tabellaria 39 Namdaecheon Stream, Hamgyeongnam-do of North Korea: 24.x.1997). ECOLOGY: The cells attach to sand grains by mucilage stalks as typically episammic diatoms and are also found as epiphytes. REMARKS: Like its genus, diatoms of species O. olsenii also contain comprehensive characteristics and share combination of morphological features: the presence of spines along valve margin, the feature of areolar type in striae, the morphology of striae and their alternation. Especially, small Opephora diatoms have never been studied in detail and a considerable confusion exists over the true nature of the taxa (Sabbe and Vyverman 1995). Some taxa belonging to genus Opephora and related taxa were transferred to new genera Pseudostaurosira (Sabbe and Vyverman 1995; Morlaes 2002). The classification of genus Opephora is still problematic today, even though some revisions have been made for the taxonomic delineation. After examination of isotype materials of the old Fig. 23. Distribution of Opephora olsenii. synonym of O. olsenii, its name was correctly designated to be O. mutabilis by Sabbe and Vyverman (1995) as the oldest legitimate name has priority. Additionally, they suggested that O. pacifica nominated by P. Petit differed with O. mutabilis (Grunow) Sabbe and Vyerman in size, striae density and the alternation of the striae to be probably a different taxon. Genus Tabellaria Ehrenberg 1840 ex Kützing 1844: 127. Bol-rok-ppyeo-dol-mal-sok ( ) Valves elongate, approximately linear in outline but with swollen poles and central regions. Valves flat with irregularly spaced striae that are either parallel or slightly radiate. Small spines along valve margins. An apical pore field at both poles of the valve, and a single rimoportula, found near the centre of the valve. Cells square or rectangular in girdle view, usually seen in girdle view. Cells joining to form zigzag or partly linear colonies. Cells typically lying in girdle view, with septa clearly visible on copulae. Copulae either complete, in which case a septum present at one pole, or open, in which case no septum present. Short strip-like plastids between septa. Lectotype: Tabellaria fenestrata (Lyngbye) Kützing 1844: 127. pl. 17. f. 22. pl. 18. f. 2. (=Diatoma fenestratum Lyngbye pl. 61. f. E 3). SPECIES: The 12 species have been reported up to date and 5 ones are currently accepted as taxonomically valid taxa (Algaebase 2009). Three Tabellaria - T. binalis, T. fenestrata, T. flocculosa - and

48 40 Algal Flora of Korea Freshwater Diatoms II one variety were described in South Korea. DISTRIBUTION: As freshwater diatoms, they are abundant in the littoral area of freshwaters. However, they ae rarely in brackish waters. ECOLOGY: They have wide tolerance from oligotrophic to eutrophic waters. KEY REFERENCE: Hustedt (1930), Patrick and Reimer (1966), Round et al. (1990), Krammer and Lange-Bertalot (1991). Key to the species of genus Tabellaria 1. Valve concave in the middle area, however, valve outline almost elliptisch T. binalis Valve convex both in the middle and on the end 2 2. Spine absent in the valve margin if focus on, the apical pore field present and deep septa in girdle view T. fenestrata Fine to stong spines in valve margin and others not above 3 3. Valve margin parallel and inherently 4 septa in intercalary bands. Under the EM field, spine relatively strong and longer than 0.5 μm and a rimoportula in the middle of valve T. quadriseptata One or more puncta in valve margin and especially numerous septa in the intercalary band 4 4. Valve breadth μm in the center and punctuate striae distinct. A rimoportula present at the pole under electron microscopy field T. ventricosa Valve breadth less than 10 μm and a rimoportula near the middle parts in valve face T. flocculosa (Krammer and Lange-Bertalot 1991) REMARKS: In the micro-structure, genus Tabellaria has two forms of copulae - with septa occluding almost half the length and no septa with ligulae. These have been considered an important systematic character (Round et al. 1990). 11. Tabellaria fenestrata (Lyngbye) Kützing 1844: 127 (Figs. 24, 25). Hustedt 1930: 122. f. 99. Patrick and Reimer 1966: 103. pl. 1. f. 1. Krammer and Lange-Bertalot 1991: 106. pl f. 2. SYNONYM: Diatoma fenestratum Lyngbye pl. 61. f. E3. Tabellaria flocculosa var. fenestrata (Lyngbye) Rabenhorst 1847: 40. Striatella fenestrata (Lyngbye) Kuntze 1898: 432. Cells rectangular in girdle view and many septa penetrating near the middle. Cells forming zigzag clonies. Valves long and narrow in width and swollen in middle with capitate apices. The valve width in the middle and terminal area almost the same or in middle slightly wider than on ends. Transapical striae delicate in 10 μm and pseudoraphe narrow. A rimoportula present in the middle portions of valves. Cells μm long and 5 10 μm wide in middle inflation. More slender form and less common than T. flocculosa.

49 Araphidineae: Diatomaceae: Tabellaria 41 TYPE: Bacillarien 127 (1844). Locality-Habitat in rivulo leniter fluent ad Naes Norvegiae. SEASONALITY: The species occurs maximumly in spring and fall and often or frequently more in winter or cold season. DISTRIBUTION: It is common species in standing or slowly flowing waters in the world. The species was reported to be significantly abundant in Fjord waters of Canada in winter (Chassé and Côté 1991) and dominated the diatom population in oligo-mesotrophic waters in Spain (Negro et al. 2000). The diatoms were found as dominant epiphytes on reed plants (Albay and Akcaalan 2003). 10 μm A B C D E F Fig. 24. Tabellaria fenestrata. A E. valve views of cells; F. girdle views ( 2,000, LM).

50 42 Algal Flora of Korea Freshwater Diatoms II On the other hand, the species made assemblages in the streams at high elevation regions (1,600 2,800 m) throughout the Lahontan Basin, California (Blinn and Herbst 2003). This species is the overall dominant plankton in Lake Michigan. T. flocculosa is cosmopolitan, however, seldom in the tropical regions. KOREA: This is the most frequent in diatom distribution in Korean freshwaters. T. fenestrata was already recorded in lakes in 1920s by a pioneer diatomist in Korea, B. W. Skvortzow (Skvortzow 1929). The species was recorded as a common flora in the middle reaches and the lakes in the Han River basin (Chung 1972, 1974). In addition of rivers and large streams, the species was observed in mountainous streams of Mountain Baekun and Wolchul in Korea (Chung et al. 1986; Lee 1989). The species exhibit relatively high frequency and low abundance in the Haman lowland wetland (Chung and Noh 1987). T. fenestrata was observed in the Yongneup Swamp and as subaerial diatoms attached on the moss in the swamp, which is a important alpine wetland in South Korea (Chung 1974). SPECIMEN EXAMINED: (Epilithic diatoms collected Fig. 25. Distribution of Tabellaria fenestrata. from the Bukcheong-Namdaecheon Stream and adjacent reservoirs in North Korea: 3.v.1998). ECOLOGY: The species prefers standing waters as planktonic or epiphytic diatoms, and has wide ecological distribution from oligotrophic to eutrophic freshwaters. This has been reported to be widely distributed in the world and rarely observed in planktons. REMARKS: A analysis shows that the two diatoms T. fenestrata and T. flocculosa never occur together in high numbers, which can be related to different trophic needs. This has been obscured by the fact that in most cases both species are present but not in large numbers (Beyens 1982). 12. Tabellaria flocculosa (Roth) Kützing 1844: 127 (Figs. 26, 27). Hustedt 1930: 123. f Patrick and Reimer 1966: 104. pl. 1. f. 4. Krammer and Lange-Bertalot 1991: 108. pl f. 1. SYNONYM: Conferva flocculosa Roth 1797: 192. pl. 4. f. 4. pl. 5. f. 6. Bacillaria flocculosa (Roth) Leiblein 1827: 258. Bacillaria flocculosa (Roth) Ehrenberg 1832: 84. Candollella flocculosa (Roth) Gaillon 1833: 34. Bacillaria tabellaris Ehrenberg 1835: 232. Striatella flocculosa (Roth) Kuntze 1898: 432. Tabellaria flocculosa (Roth) Knudson 1952: 436.

51 Araphidineae: Diatomaceae: Tabellaria 43 Cells rectangular in girdle view and united to the corners to form zigzag colonies. Numerous intercalary bands to make longer pervalvar axes than valvar axes. However, the ratios of length to width are less than 4:1. Valves swollen in middle portions and wider than the apices. Valve somewhat asymmertrical to either transverse or apical axis, sometimes slightly asymmetrical to both axes. The positions of rimoportulae variable. Transapical striae in 10 μm. Valves μm long and 5 16 μm wide. TYPE: Bacillaria 127 (1844). SEASONALITY: T. flocculosa dominates the other algae in June in some freshwaters and makes often the spr-ing pulses to accompany other diatoms. Its maximum occurs in spring and autumn. DISTRIBUTION: The species is common in freshwaters in the world and occasionally blooms in standing waters. The species was dominant throughout Holocene sediment of slight acidic lakes in the diatom records of Canada (Prather and Hickman 2000). This is rather periphytic diatom, A B C 10 μm D E F G I J K H Fig. 26. Tabellaria flocculosa. A H. valve views of cells; I, J. girdle views of cells ( 2,000, LM); K. the inner face of a valve and a rimoportular pore in central area (SEM); A, B. DIC images of LM.

52 44 Algal Flora of Korea Freshwater Diatoms II however, occurs often as tychoplankton. KOREA: This is one of diatoms having the most high frequency in streams, rivers, lakes or reservoirs and mountains in Korea. T. flocculosa was already recorded from Korean lakes in 1920s by Skvortzow (1929). The species was observed in middle areas of the Han River (Chung 1972). Besides the rivers, the species was observed in mountainous streams of Mountain Baekun and Wolchul in Korea (Chung et al. 1986; Lee 1989). The species showed very low frequency in the Haman lowland wetland (Chung and Noh 1987). The diatoms were observed in the Yongneup Swamp and in subaerial moss of the swamp, the unique alpine wetland (Chung 1974). T. flocculosa was observed with low abundance in epipilthic algal assemblages of some streams (Geumho River) (Lee and Chung 1992). SPECIMEN EXAMINED: (Epilithic diatoms collected from the Bukcheong-Namdaecheon Stream: 3.v.1998). ECOLOGY: The species was dominant in many acidotrophic-oligotrophic lakes in North America and a member of the important in the oligotrophic lakes of Finland (Lepistö and Rosenström 1998). This was rare in a few eutrophic lakes and ponds. It was classified as an acidic diatom in the peatland or rather Fig. 27. Distribution of Tabellaria flocculosa. acidic waters. The variety was relatively abundant in two mountain bogs, the Danjoneup in Mountain Yeongchuk in Yangsan, September 2006, and Daeseongneup Bog and its adjacent wetlands near Mountain Jeongjok in Ulsan, December REMARKS: The specimens with shorter frustules are more often found in acid water of bogs and ponds, whereas those with longer frustules seem to be more often found in oligotrophic to mesotrophic water (Lepistö and Rosenström 1998).

53 45 Literature Cited Ács, É., K.T. Kiss, K. Szabó and J. Makk, Short-term colonization sequence of periphyton on glass slides in a large river (River Danube, near Budapest). Algological Studies 100: Albay, M. and R. Akcaalan, Comparative study of periphyton colonisation on common reed (Phragmites australis) and artificial substrate in a shallow lake, Manyas, Turkey. Hydrobiologia : Algaebase, Antoniades, D. and M.S.V. Douglas, Characterization of high arctic stream diatom assemblages from Cornwallis Island, Nunavut, Canada. Canadian Journal of Botany 80: APHA, Standard methods for the examination of water and wastewater. 19th edition. American Public Health Association (APHA) Bertrand, C., S. Fayolle, E. Franquet and A. Cazaubon, Responses of the planktonic diatom Asterionella formosa Hassall to abiotic environmental factors in a reservoir complex (south-eastern France). Hydrobiologia 501: Beyens, L., On the difference in ecology between Tabellaria fenestrata and T. flocculosa (Diatomophyceae). Bulletin de la Société Royale de Botanique 115: Biggs, B.M. and G.M. Price, A survey of filamentous algal proliferations in New Zealand rivers. New Zealand Journal of Marine and Freshwater Research 21: Blinn, D.W. and D.B. Herbst, Use of diatoms and soft algae as indicators of environmental determinants in the Lahontan Basin, USA. Annual report for California State Water Resources Board Contract Agreement CT pp. Carpenter, K.D. and I.R. Waite, Relations of habitat-specific algal assemblages to land use and water chemistry in the Willamette basin, Oregon. Environmental Monitoring and Assessment 64: CAS, by California Academy of Sciences (CAS). Chapman, J.C. and B.L. Simmons, The effects of sewage on alpine streams in Kosciusko National Park, NSW. Environmental Monitoring and Assessment 14: Chassé, R. and R. Côté, Aspects of winter primary production in the upstream section of Saguenay Fjord. Hydrobiologia 215: Chung, J. and Y.O. Kim, An assessment of water quality by epilithic diatoms of Paek-chon water system. Journal of Environmental Science Institute, Kyungpook National University 5: Chung, Y.H., A study on the microflora of the Han River (VI). Taxonomy of phytoplankton for the South Han River and estimation of water pollution levels on the central area of the Han River. Korean Journal of Botany 15: Chung, Y.H., A study on the microflora of the Han River (VIII). On the subaerial diatoms from Yongneup (swamp) in Mt. Daeam near DMZ in Korea. Korean Journal of Botany 17: Chung, Y.H. and K.T. Kim, A study on the microflora of the Han River (V). The taxonomy of phytoplankton and its seasonal variation in the area of middle course of the Han River. Report for the IBP, Korea 4: Chung, Y.H. and K.T. Kim, Phytoplankton from the swamp Yong-neup located in Mt. daeam as a water source of the North Han River. Korean Journal of Environmental Biology 5: Chung, Y.H. and C.H. Koh, A study on the microflora of the Han River. VII. Taxonomy of diatoms of the Paroho. Korean Journal of Botany 17: Chung, Y.H. and K. Lee, A study on the microflora of the Han River (IX). Taxonomy of phytoplankton and environmental conditions in Paldang Dam reservoir. Proceedings of College of Natural Science, Seoul

54 46 Algal Flora of Korea Freshwater Diatoms II National University 3: Chung, Y.H. and K. Lee, A study on the microflora of the Han River. X. Distribution of diatoms in the Mid-Han River. Proceedings of College of Natural Science, Seoul National University (2): Chung, Y.H. and K.H. Noh, The diatom flora of lowland swamp in Haman county, Korea. Proceedings of College of Natural Science, Seoul National University, Korea 12: Chung, Y.H., E.S. Kay and D.W. Park, A study on the microflora of the Han River. II. The phytoplanktons and its seasonal variation in the area of the Chun-chon and the Chung-pyong reservoir of the Han River. Korean Journal of Botany 11: Chung, Y.H., K.H. Noh and O.M. Lee, A study on the flora and standing crop of phytoplankton at watershed in Mt. Paegun. Report of the Korean Association for Conservation of Nature (KACN) 24: Chung, Y.H., J.H. Shim and M.J. Lee, A study on the microflora of the Han River. I. The phytoplanktons and the effect of the marine water in the lower course of the Han River. Korean Journal of Botany 8: Clerk, S., P. Hall, R. Quinlan and P. Smol, Quantitative inferences of past hypolimnetic anoxia and nutrient levels from a Canadian Precamrian Shield lake. Journal of Paleolimnology 23: Duthie, H.C. and V.M. Stout, Phytoplankton periodicity of the Waitaki Lakes, New Zealand. Hydrobiologia 138 (Dev. Hydrobiol. 33): Eilers, J.M., J. Kann, J. Cornett, K. Moser and A. St. Amand, Paleolimnological evidence of change in a shallow, hypereutrophic lake: Upper Klamath Lake, Oregon, USA. Hydrobiologia 520: Eulin, A. and R.L. Cohu, Epilithic diatom communities during the colonization of artificial substrates in the River Garonne (France): Comparison with the natural communities. Archiv für Hydrobiologie 143: Flower, R.J., V.J. Jones and F.E. Round, The distribution and classification of problematic Fragilaria (virescens v.) exigua Grun./Fragilaria exiguiformis (Grun.) Lange-Bertalot: A new species or a new genus? Diatom Research 11: Fukushima, H., T. Kobayashi, K. Terao and S. Yoshitake, Morphological variability of Diatoma vulgare Bory var. grande. 9th Diatom Symposium: Genkal, S.I., Morphological variability and taxonomy of Diatoma tenue Ag. (Bacillariophyta). International Journal of Algae 6: Hasle, G.R. and E.E. Syvertsen, Marine diatoms. In: Tomas C.R. (ed.), Identifying Marine Diatoms and Dinoflagellates. Academic Press, Inc., San Diego, California. pp Hayakawa, T., S. Kudoh, Y. Suzuki and M. Takahashi, Temperature-dependent changes in colony size of the freshwater pennated diatom Asterionella formosa (Bacillariophyceae) and their possible ecological implications. Journal of Phycology 30: Huber-Pestalozzi, G., Das Phytoplankton des Süβwassers. Systematik und Biologie. In: Thienemann A. (ed.), Die Binnengewässer. E. Schweizerbart sche Verlag. Stuttgart. 16(2) Häfte 2, Supplement: Hustedt, F., Bacillariophyta (Diatomeae). In: Pascher A. (ed.). Die Süβwasserflora Mitteleuropas Vol nd ed. Verlag von Gustav Fisher, Jena. 466 pp. Hustedt, F., Die Kieselalgen. Deutschlands, Österreichs und der Schweiz: unter Berücksichtigung der übrigen Länder Europas sowie der angrenzended Meeresgebiete. In: Dr. L. Rabenhorst s Kryptogamen-Flora von Deutschland, Österreich und der Schweiz. Teil 2. Otto Koeltz Science Publishers, Koenigstein, Germany. pp Hutchinson, G.E., A Treatise on Limnology. Vol. 2. Introduction to lake biology and the limnoplankton. Wiley pp. Ibelings, B., W. Admiraal, R. Bijkerk, T. Ietswaart and H. Prins, Monitoring of algae in Dutch rivers: Does it meet its goals? Journal of Applied Phycology 10:

55 Literature Cited 47 Ilmavirta, V., Dynamics of phytoplanktonic production in the oligotrophic lake Pääjärvi, southern Finland. Annales Botanici Fennici 12: Jeong, S.W., J.H. Lee and J.S. Yu, Environmental Studies of the Lower Part of the Han River - 5. Blooming Characteristics of Phytoplankton Communities. Korean Journal of Phycology 18: Kaweckal, B. and C.T. Robinson, Diatom communities of lake/stream networks in the Tatra Mountains, Poland, and the Swiss Alps. Oceanological and Hydrobiological Studies 37: Kim, Y.J., J.S. Choi and H.S. Kim, Structure of phytoplankon communtiy in lake Imha. Korean Journal of Limnology 30: Kim, Y.J., Ecological study of phytoplankton community and trophic states using indications in Lake Paldang. Korean Journal of Limnology 29: Kim, Y.J., Ecological characteristics of phytoplankton community in Lake Paldang Dam. Korean Journal of Limnology 31: Kim, Y.J. and J.H. Lee, Comparison of phytoplankton communities of six dam lakes in the Nakdong River system. Korean Journal of Limnology 29: Kingston, J.C., Araphid and monoraphid diatoms. In: Wehr J.D. and Sheath R.G. (eds) Freshwater Algae of North America: Ecology and Classification. Academic Press. pp Kling, H.J., Asterionella formosa Ralfs: The process of rapid size reduction and its possible ecological significance. Diatom Research 8: KNCCN., Literature Survey for Biological Species in Korea. V. Lower plant (algae). The current state of biological species in Korea. Vol. 11. The Korean National Council for Conservation of Nature (KNCCN). Körner, H., Morphologie und Taxonomie der Diatomeengattung Asterionella. Nova Hedwigia 20: Kownacki, A., E. Dumnicka, J. Kwandrans, J. Galas and M. Ollik, Benthic communities in relation to environmental factors in small high mountain ponds threateded by air pollutants. Boreal Environmental Research 11: Krammer, K. and H. Lange-Bertalot, Bacillariophyceae. Teil 3. Centrales. Fragilariaceae, Eunotiaceae. Bd 2/3 von Süsswasserflora von Mitteleuropa (Ettl H., Gerloff J., Heynig H. and Mollenhauer D., eds.). Gustav Fischer Verlag. Berlin. 598 pp. Krejci, M.E. and R.L. Lowe, The seasonal occurrence of macroscopic colonies of Meridion circulare (Bacillariophyceae) in a spring-fed brook. Transactions of the American Microscopical Society 106: Lange-Bertalot, H., Toleranzgrenzen und Populationsdynamik benthischer Diatomeen bei unterschiedlich starker Abwasserbelastung. Archiv fur Hydrobiologia, Supplement 56, Algological Studies 23: Lange-Bertalot, H., Zur systematischen Bewertung der bandförmigen Kolonien bei Navicula und Fragilaria - Kriterien für die Vereinigung von Synedra (subgen. Synedra) Ehrenberg mit Fragilaria Lyngbye. Nova Hedwigia 33: Lange-Bertalot, H., Könne Staurosirella, Punctastriata und weitere Taxa sensu Williams and Round als Gattungen der Fragilariaceae kritischer Prufung standhalten? Nova Hedwigia 49: Lee, E.J. and K.S. Cho, Yearly variation of phytoplankton in Lake Soyang. Korean Journal of Limnology 27: Lee, J.H. and J. Chung, Station variation of epilithic diatoms according to pollution degree from the Kumho River. Korean Journal of Limnology 25: Lee, J.H. and S.W. Jung, Environmental studies in the lower part of the Han River. VII. Long term variations and prospect of the phytoplankton community. Algae 19: Lee, J.H., J.S. Choi, Y.J. Kim and J. Chung, 1995a. Flora of planktonic diatoms in the Naktong River Estuary,

56 48 Algal Flora of Korea Freshwater Diatoms II Korea. Korean Journal of Limnology 28: Lee, J.H., J. Chung and T. Gotoh, Diatoms of the Kwang River (Kwangchun) South Korea 1. Centrales, Pennales except Naviculaceae. Diatom 9: Lee, K., On the flora of phytoplankton in the watersheds of Mt. Wolch ul. Report of the Korean Association for Conservation of Nature (KACN) 27: Lee, K. and S.K. Yoon, Seasonal changes of the phytoplankton community in the Imjin River. Korean Journal of Limnology 35: Lepistö, L. and U. Rosenström, The most typical phytoplankton taxa in four types of boreal lakes. Hydrobiologia : Lepistö, L., P. Kauppila, J. Rapala, M. Pekkarinen, I. Sammalkorpi and L. Villa, Estimation of reference conditions for phytoplankton in a naturally eutrophic shallow lake. Hydrobiologia 568: Liukkonen, M., T. Kairesalo and J. Keto, Eutrophication and recovery of Lake Vesijärvi (south Finland): Diatom frustules in varved sediments over a 30-year period. Hydrobiologia : Lund, J.W.G., Studies on Asterionella. I. The origin and nature of the cells producing seasonal maxima. Journal of Ecology 37: Marks, J.C. and M.E. Power, Nutrient induced changes in the species composition of epiphytes on Cladophora glomerata Kütz. (Chlorophyta). Hydrobiologia 450: Marshall, H.G., L. Burchardt and R. Lacouture, A review of phytoplankton composition within Chesapeake Bay and its tidal estuaries. Journal of Plankon Research 27: Martínez De Fabricius, A.L., N. Maidana, N. Gómez and S. Sabater, Distribution patterns of benthic diatoms in a Pampean river exposed to seasonal floods: The Cuarto River (Argentina). Biodiversity and Conservation 12: Miho, A., A. Cake and E. Çarçani, Diatoms in the stomach content of barbel (Barbus meridionalis) from Shkumbini river (Central Albania). Journal of Environmental Protection and Ecology (JEPE) 6: Moed, J.R. and H.L. Hoogveld, Dominant diatoms in Tjeukemeer: The appearance of Diatoma elongatum (Lyngb.) Agardh. Freshwater Biology 5: Morales, E.A., Studies in selected fragilarioid diatoms of potential indicator value from Florida (USA) with notes on the genus Opephora Petit (Bacillariophyceae). Limnologica 32: Moss, B., The composition and ecology of periphyton communities in fresh-waters. 11. Interrelationships between water chemistry, phytoplankton populations and periphyton populations in shallow lakes and associated experimental reservoirs (Lund Tubes). British Phycological Journal 16: Müller-Haeckel, A. and H. Håkansson, The diatom flora of a small stream near Abisko (Swedish Lapland) and its annual periodicity, judged by drift and colonization. Archiv für Hydrobiologie 84: Negro, A.I., C. de Hoyos and J.C. Vega, Phytoplankton structure and dynamics in Lake Sanabria and Valparaíso reservoir (NW Spain). Hydrobiologia 424: Park, J.K. and S.K. Kim, Annual variation of fresh water quality in the Gachang Dam Reservoir. Korean Journal of Chemical Engineering 20: Patrick, R. and C.W. Reimer, The Diatoms of the United States Exclusive of Alaska and Hawaii. Vol. 2, Part 1. Monographs of the Academic of Natural Science of Philadephia, Number pp. Potapova, M. and D.F. Charles, Distribution of benthic diatoms in U.S. rivers in relation to conductivity and ionic composition. Freshwater Biology 48: Poulin, M., L. Berard-Therriault and A. Cardinal, Fragilaria and Synedra (Bacillariophyceae): A morphological and ultrastructural approach. Diatom Research 1: Prather, C. and M. Hickman, History of a presently slightly acidic lake in northeastern Alberta, Canada,

57 Literature Cited 49 as determined through analysis of the diatom record. Journal of Paleolimnology 24: Reynolds, C.S., The Ecology of Freshwater Phytoplankton. Cambridge University Press, Cambridge. 396 pp. Rhode, K.M., J.L. Pappas and E.F. Stoermer, Qunatitative analysis of shape variation in type and modern populations of Meridion (Bacillariophceae). Journal of Phycology 37: Robinson, C.T. and B. Kawecka, Benthic diatoms of an alpine stream/lake network in Switzerland. Aquatic Sciences: Research Across Boundaries 67: Robinson, C.T., B. Kawecka, L. Füreder and A. Peter, Biodiversity of flora and fauna in Alpine Waters. Alpine Waters 6: Round, F.E., The classification of the genus Synedra. Nova Hedwigia, Beiheft 64: Round, F.E., The circumscription of Synedra and Fragilaria and their subgroupings. 7th Diatom- Symposium: Round, F.E., R.M. Crawford and D.G. Mann, The Diatoms. Biology and Morphology of the Genera. Cambridge University Press, Cambridge. 747 pp. Sabbe, K. and W. Vyverman, Taxonomy, morphology and ecology of some widespread representatives of the diatom genus Opephora. European Journal of Phycology 30: Sherwood, A.R. and R.G. Sheath, Seasonality of macroalgae and epilithic diatoms in spring-fed streams in Texas, USA. Hydrobiologia 390: Skvortzow, B.W., Freshwater diatoms from Korea, Japan. The Philippine Journal of science 38: Spauling, S.A., J.V. Ward and J. Baron, Winter phytoplankton dynamics in pelahial of small artificial reservoirs in Spring. International Review of Hydrobiology 83: Steinman, A.D. and A.F. Parker, Influence of substrate conditioning on periphytic growth in a heterotrophic woodland stream. Journal of the North American Benthological Society 9: Sundbäck, K., The epipsammic marine diatom Opephora olsenii Müller Diatom Research 2: Szabó K., K. Kiss, G. Taba and E. Ács, Epiphytic diatoms of the Tisza River, Kisköre Reservoir and some oxbows of the Tisza River after the cyanide and heavy metal pollution in Acta Botanica Croatia 64: Takano, K., Y. Ishikawa, H. Mikami, S. Ban, T. Yoshida, T. Aono, K. Imada, R. Yasutomi, K. Takeuchi and S. Hino, Analysis of the change in dominant phytoplankton species in unstratified Lake Oshima- Ohnuma estimated by a bottle incubation experiment. Limnology 2: Tilman, D., S.S. Kilham and P. Kilham, Morphometric changes in Asterionella formosa colonies under phosphate and silicate limitation. Limnology and Oceanography 42: Tomas, X. and S. Sabater, Diatom flora of the Llobregat River and its relation to water quality. International Vereinig theor angew Limnologie 22: Vavilova, V.V. and W.M.J. Lewis, Temporal and altitudinal variations in the attached algae of mountain streams in Colorado. Hydrobiologia 390: Vilbaste, S., Benthic diatom communities in Estonian rivers. Boreal Environmental Research 6: Vilbaste, S. and J. Truu, Distribution of benthic diatoms in relation to environmental variables in lowland streams. Hydrobiologia 493: Vis, C., C. Hudon, A. Cattaneo and B. Pinel-Alloul, Periphyton as an indicator of water quality in the StLawrence River (Québec, Canada). Environment and Pollution 101: Williams, D.M., Morphology, taxonomy and interrelationships of the ribbed araphid diatoms from the genera Diatoma and Meridion (Diatomaceae: Bacillariophyta). Bibliotheca Diatomologica 8. Vaduz: Cramer. Williams, D.M., Comparative morphology of some species of Synedra Ehrenb. with a new definition of

58 50 Algal Flora of Korea Freshwater Diatoms II the genus. Diatom Research 1: Williams, D.M., Cladistic analysis of some freshwater araphid diatoms (Bacillariophyta) with particular reference to Diatoma and Meridion. Plant Systematics and Evolution 171: Williams, D.M. and F.E. Round, Revision of the genus Synedra Ehrenb. Diatom Research 1: Williams, D.M. and F.E. Round, Revision of the genus Fragilaria. Diatom Research 2: Witkowski, A., Diatoms of the Puck Bay coastal shallows (Poland, Southern Baltic). Nordic Journal of Botany 11: Yasuno, M., S. Fukushima, J. Hasegawa, F. Shioyama and S. Hatakeyama, Changes in the benthic fauna and flora after application of temephos to a stream on Mt. Tsukuba. Hydrobiologia 89: Zębek, E., Qualitative and quantitative changes of diatoms with relation to physiochemical water parameters in the littoral zone of the urban Lake Jeziorak Ma ly. Oceanological and Hydrobiological Studies 36: Ziemann, H., L. Kies and S. Clause-Jürgen, Desalinization of running waters, III. Changes in the structure of diatom assemblages caused by a decreasing salt load and changing ion spectra in the river Wipper (Thuringia, Germany). Limnologica 31:

59 Bryozoa: Gymnolaemata: Cheilostmata: Inovicellata, Malacostega, Flustrina, Ascophora 51 Fragilaria, Hannaea, Pseudostaurosira, Punctastriata, Staurosira, Staurosirella Jung Ho Lee

60

61 Bryozoa: Gymnolaemata: Cheilostmata: Inovicellata, Malacostega, Flustrina, Ascophora 53 List of Taxa Family Diatomaceae Dumortier 1822 Genus Fragilaria Lyngbye 1819 Fragilaria bidens Heiberg 1863 Fragilaria capitellata (Grunow) Petersen 1946 Fragilaria capucina Desmazieres 1825 Fragilaria capucina var. radians (Kützing) Lange-Bertalot 1991 Fragilaria crotonensis Kitton 1869 Fragilaria famelica (Kützing) Lange-Bertalot 1980 Fragilaria gracilis Ǿstrup 1910 Fragilaria mesolepta Rabenhorst 1861 Fragilaria parva Tuji and D. M. Williams 2008 Fragilaria rumpens Kützing 1844 Fragilaria vaucheriae (Kützing) Petersen 1938 Genus Hannaea Patrick 1966 Hannaea arcus var. recta (Cleve) M. Idei 2006 Hannaea arcus var. subarcus (Iwahashi) Lee 1992 Genus Pseudostaurosira Williams and Round 1987 Pseudostaurosira brevistriata (Grunow) Williams and Round 1987 Genus Punctastriata Williams and Round 1987 Punctastriata linearis Williams and Round 1987 Genus Staurosira Ehrenberg 1843 Staurosira construens Ehrenberg 1843 Staurosira construens var. binodis (Ehrenberg) Hamilton 1992 Staurosira elliptica (Schumann) Williams and Round 1987 Staurosira venter (Ehrenberg) H. Kobayasi 2002 Genus Staurosirella Williams and Round 1987 Staurosirella pinnata (Ehrenberg) Williams & Round 1987

62 54 Introduction Diatoms are the most common and diverse primary producers in freshwaters and in most aquatic systems provide the basis of their food webs (Round et al. 1990). Because diatoms are effective indicators of environmental changes, their use for biomonitoring of pollution in rivers becomes increasingly important (Watanabe et al. 2005). Fragilarioid diatoms, genus Fragilaria (sensu lato), are common in planktonic and benthic samples from both, rivers and lakes, where they typical account for a considerable portion of the total biomass produced by algae (Round et al. 1990). Fragilarioid diatoms have also been used as good indicators of changes in nutrient levels, salinity and ph. However, the taxonomy of some frailarioid diatoms is still problematic (Paull et al. 2008). For instance, Krammer and Lange-Bertalot (1991) combined almost of the common freshwater Fragilarioid species in the genus Fragilaria. Contrary to this lumping move, separated Williams and Round (1987, 1990) five genera (Staurosirella, Staurosira, Pseudostaurosira, Punctastriata, Fragilariforma) as independent genera from Fragilaria. Especially in Korea, the taxonomy of fragilarioid species is rarely pursued. There is also a lack of detailed analyses using light microscopy and scanning electron microscopy. In this work, taxonomical characterizations of twenty fragilarioid taxa collected from rivers and lakes in Korea are provided. Ecological information on each taxon is also presented from various references.

63 Bryozoa: Gymnolaemata: Cheilostmata: Inovicellata, Malacostega, Flustrina, Ascophora 55 Materials and Methods The diatom samples were cleaned by the permanganate method (Hendey 1974) and mounted in Pleurax. Microscopic slides were examined with a Zeiss Axioscope 2 microscope. For scanning electron microscopy, the cleaned material was dehydrated in ethanol, then a drop of the suspension was air-dried on stubs, sputter-coated with graphite and gold, and examined with a JEOL-JSM-840 SEM. The terminology used here follows Anonymous (1975) and Ross et al. (1979).

64

65 57 Taxonomic Notes Family Diatomaceae Dumortier Mak-dol-mal-gwa ( ) Araphid diatoms are an as yet phylogenetically unresolved group consisting of diatoms that can not be accomodated in the centric or the raphid diatoms (Williams 1985). The taxa occur as unicells or in colonies: ribbon-like, stellate, zigzag, or attached to a mucilage pad at one end of the cells. Certain planktonic populations in large lakes can selectively develop either offshore (Stoermer and Yang 1970) or nearshore (Stoermer 1968). Key to the genera of family Diatomaceae 1. Frustules lacking a raphe system Araphid diatoms 2 Frustules with a raphe system on one valve only Monoraphid diatoms 2. Transapical ribs present 3 Transapical ribs absent 6 3. Septa present 4 Septa absent 5 4. Septa large Tetracyclus Septa small, not noticeable by light microscopy Meridion 5. Raised sternum, pore fields on mantle and face, girdle structure more complex Diatoma Without sternum, pore fields on mantle only, girdle structure more simple Distrionella 6. Septa present 7 Septa absent 8 7. Cells elongate, capitate, wider at the center, large polar pore field on face and mantle Tabellaria Cells elliptical or panduriform, apical pore field of fine striae rows Oxyneis 8. Long, narrow cells; two labiate processes per valve; solitary, stellate, pincushion, or bandshaped colonial habit 9 Shorter cells; labiate process either one per valve or lacking; solitary, zig-zag or band-shaped colonies Heteropolar; slender, capitate cells; broader at cell ends than at the cell center; planktonic; stellate colonies Asterionella Isopolar, broader at the center than at the poles, benthic or planktonic Cells arcuate, with central thickening on the ventral side Hannaea Cells linear to linear-lanceolate Two terminal spines at each pole in many species, areolae with simple vela Synedra No spines, areolae with complex cribra, thickened fascia, brackish waters Ctenophora 12. Broad striae composed of linear areolae or composed of multiseriate rows of areolae 13 Narrow striae composed of uniseriate rows of round to transapically elongate areolae Striae composed of multiseriate rows of areolae Punctastriata

66 58 Algal Flora of Korea Freshwater Diatoms II Striae composed of linear areolae Cells ovate-elliptical, valve depressed at head-pole, lacking spines Martyana Cells elliptical, linear, or cruciform; possessing spines Staurosirella 15. Cells longer, linear to linear-lanceolate, valve margins not undulate Fragilaria Cells shorter; linear, elliptical, or cruciform Cells elliptical, lanceolate, or linear; striae very finely punctate; sternum very narrow 17 Striae composed of larger areolae, sternum wider Cell margins often undulating; one labiate process per valve Fagilariforma Cells sometimes with subrostrate ends; no labiate processes Stauroforma 18. Striae made of regularly spaced, round areolae Staurosira Striae made of few transapically elongate areolae Pseudostaurosira (Wehr and Sheath 2003) Genus Fragilaria Lyngbye Gim-bal-dol-mal-sok ( ) Frustules rectangular in girdle view, usually forming linear colonies, or some as growing singly. Cells in girdle view are either oblong or swollen at the centre, and then linked only by themselves. Valves are linear to elliptical and apices rostrate to capitate. Striae are more or less evenly spaced. At the valve centre the striae are often occluded, appearing as ghost structures. Sternum is narrow. Apical pore field is present, situated at the apices of the mantle. The pore fields are of the ocellulimbus type. A single rimoportula is present, usually lying at a pole. Williams and Round (1987) restricted Fragilaria to taxa that form colonies and have simple rows of areolae and a single rimoportula. Representatives of the genus occurs in freshwater. Type species : F. capucina Desmaz. Key to the species of genus Fragilaria 1. Apical axis weakly sickle- or banana-shaped or valves and sharply widened in the centre and irregularly curved to bent. Ventral side of the valve more or less distended in the central region F. arcus complex (Hannaea arcus complex) in part Apical axis usually straight 2 2. Valves with short striae along edge, axial area therefore expanded to a large central area 9 Not with the above combination of characteristics 3 3. Central area symmetrical and abruptly set off from the axial area, encircled with a distended ring 7 Central area not as above or absent 4 4. Valves needle-shaped, gradually narrowing from the centre to the ends 5 If valves upon population average shorter in relation to width, see also the following taxa: F. delicatissima in part, F. tenera in part, F. famelica in part, F. capucina species complex in part, F. ulna species complex in part 5. Valves more or less concave in the middle 6

67 Araphidineae: Diatomaceae: Fragilaria 59 Valves more or less distended in the centre, ends less elongate and more widely rounded F. construens (Staurosira construens) 6. Valve edge more undulating, very rarely also F. pinnata (Staurosirella pinnata), F. construens (Staurosira construens) in part Not as above 8 7. Central area more or less distinctly unilateral, valve length usually less than 50 μm; in addition to other tribes or individual examples of these complexes F. vaucheriae and F. capitellata. Not with the above combination of characteristics 9 8. Valves without central area set off from the axial area F. construens (Staurosira construens) and its related taxa complex in part Central area more or less distinctily clearly set off, compare also F. famelica in part. F. capucina spcies complex in part 9. Valves rhomboid to broadly lanceolate or with distended enlarged area elongated close to the poles. Striae very coarse, maximum of 12/10 μm, nevertheless never punctate, at most appearing lanceolate F. pinnata (Staurosirella pinnata) in part Valves linear, elliptical, narrowly lanceolate. Striae distinctly punctuate, central area moderately wide F. elliptica (Staurosira elliptica) in part. (Krammer and Lange-Bertalot 2000) 1. Fragilaria bidens Heiberg 1863 (Figs. 1, 2). Krammer and Lange-Bertalot 1991: 127. f. 111: Metzeltin and Lange-Bertalot 1998: taf. 1. f. 29, 30. BASIONYM: Fragilaria bidens Heiberg SYNONYM: Synedra pulchella var. minuta Hustedt in Schmidt et al Synedra rumpens var. fragilarioides f. constricta Hustedt Frusules rectangular in girdle view, forming bandlike colonies. Valve linear-lanceolate, swollen on each side of the central area. Apices somewhat rounded. Axial area linear. Central area longer than broad, somewhat swollen. Striae almost parallel, in 10 μm. Length μm. Width of the valve in swollen area 3 4 μm. TYPE: Not clear. SEASONALITY: Present throughout the year. DISTRIBUTION: Cosmopolitan, frequently occurring in northern Europe. KOREA: This species occurs possibly throughout Fig. 1. Distribution of Fragilaria bidens.

68 60 Algal Flora of Korea Freshwater Diatoms II Korea; Nakdong River (Lee 2002) and Namchun stream in Kyungsan city (Jung 2008). SPECIMEN EXAMINED: LJ217 (Kwang river, Ulchin county, Kyungbuk). ECOLOGY: The species is an oligo-meso eutrophic taxon (Krammer and Lange-Bertalot 1991). REMARKS: This species is very similar to Fragilaria parva Tuji and D. M. Williams and Synedra E A 10 μm B C D F G H Fig. 2. Fragilaria bidens. A D. (LM. 2,000); E H. (SEM); E, F. external views of valves; G, H. external views of valve apicies. Note that a rimoportula is present on only one apex.

69 Araphidineae: Diatomaceae: Fragilaria 61 familiaris f. major Grunow in Van Heruck. It differs from F. parva (18 20 striae in 10 μm) and S. familiaris f. major (18 19 striae in 10 μm) by its coarser density of striae. 2. Fragilaria capitellata (Grunow) Petersen 1946: 54 (Figs. 3, 4). Krammer and Lange-Bertalot 1991: 124. f. 109: Lee et al. 1994: 19. f. 15, 16. Kawashima and Kobayasi 1994: 9. f. 1B G. Kobayasi et al. 2006: 56. pl. 71, 72. BASIONYM: Synedra capitellata Grunow in Van Heruck SYNONYM: Fragilaria vaucheriae var. capitellata Ross Fragilaria capucina var. capitellata (Grun.) Lange-Bertalot Valve linear-lanceolate, narrowed towards capitate apices. Pseudoraphe narrow. Central area unilateral, somewhat swollen. Rimoportula on the pole of only one valve. Striae in 10 μm. Length μm, width μm. Outer opening in each areola ornamented with rota occlusion under SEM. TYPE: Not clear. SEASONALITY: All around year. DISTRIBUTION: Worldwide in temperate freshwater. KOREA: This species occurs in various running waters in Korea-Lake Imha (Kim 1993), Lake Kwang River (Kwangchun) in Ulchin county (Lee et al. 1994), Kumho River (Park 1995), Banbyeon Stream (Banbyeonchun) in Cheongsong county (Kim 1993), Tongwha stream (Tongwhachun) in Daegu (Hong 1990), Nammaeji reservoir (Kim et al. 1993) and Nam stream (Namchun) in Kyungsan city (Jung 2008). SPECIMEN EXAMINED: LJ207 (Kwang river, Ulchin county, Kyungbuk). ECOLOGY: The taxon is β-saproxenous, ph circumneutral, and eutraphenic. REMARKS: The taxon is distinguished by the broader valve width and finer striae density different from Flagilaria vaucheriae (Kütz.) Petersen (Kobayasi et al. 2006). F. capitella has external opening of areolae with rota occlusion against F. vaucheriae, a very similar species, having simple pore external areolar opening (see, Kobayasi et al. 2006: pl. 72, 78). Kawashima and Kobayasi (1994) treated F. capitellata as a solitary taxon not forming colonies because they have not been able to observe its connecting spines. This species has been commonly found in the samples from lakes and reser- Fig. 3. Distribution of Fragilaria capitellata.

70 62 Algal Flora of Korea Freshwater Diatoms II A 10 μm B C D E G F H Fig. 4. Fragilaria capitellata. A E. (LM. 2,000); F H. (SEM); F. external view of valve; G, H. external views of valve apices. Note that a rimoportula is present in only one apex.

71 Araphidineae: Diatomaceae: Fragilaria 63 voirs (Kawashima and Kobayasi 1994). 3. Fragilaria capucina Desmazieres 1825 (Figs. 5, 6). Krammer and Lange-Bertalot 1991: 121. f. 108: 1 8. Lee et al. 1994: 19. pl. 1. f. 13. Tuji Tuji and Williams Watanabe 2006: 102. pl. IIA 8: BASIONYM: Fragilaria capucina var. capucina Desmazieres SYNONYM: Fragilaria capucina var. lanceolata Grunow in Van Heruck Synedra rumpens var. familiaris f. major Grunow in Van Heruck Synedra fallax Grunow in Van Heruck Synedra rumpens var. acuta (Ehrenberg) Rabenhorst 1864 sensu Grunow. Fragilaria intermedia sensu auct nonnull. Fragilaria product sensu auct nonnull. Frustules forming long filaments, in girdle view linear, narrower towards the ends, sometimes forming short chains of two to three. Valve linear, attenuated toward the ends, thicker at the central area; not distinctly swollen. Apices swollen, somewhat capitate. Pseudoraphe narrow. Central area usually with distinct longer than broad transverse fascia. Striae parallel, not distinctly punctuate, in 10 μm. Width μm. TYPE: Lectotype: BM in BM. SEASONALITY: Present throughout the year. DISTRIBUTION: Worldwide in temperate freshwater. KOREA: This species occurs in various streams, lakes and ponds-all sites in Nakdong River (Lee 2002; Kim and Lee 1991), Sinchun river in Daegu (Choi et al. 1993), Lake Imha (Kim et al. 1995), Milyang River (Choi and Chung 1995), Imha Lake (Kim et al. 1997), Somjin River (Lee and Yoon 1999), Imjin River (Lee and Yoon 2002), Kwang River in Ulchin county (Lee et al. 1994), Kumho River in Daegu (Lee and Chung 1992), Guem River (Kim et al. 2000), Banbyeon Stream in Cheongsong county (Kim 1993) and Tongwha stream in Daegu (Hong 1990). SPECIMEN EXAMINED: LJ1008 (Seomjin river, Sunchang county, Jeonbuk). ECOLOGY: The species is widely distributed on freshwater lakes, ponds, or slow-flowing streams. They are saproxenous and alkaliphilous taxa. REMARKS: According to the observation of the type material of Fragilaria capucina Desmazieres (label no. 453 of Desmazieres) by Tuji and Williams (2006a), this Fig. 5. Distribution of Fragilaria capucina.

72 64 Algal Flora of Korea Freshwater Diatoms II E A B C 1 D F 1 H I J G 10 μm K Fig. 6. Fragilaria capucina. A D. Fragilaria capucina. (LM. 2,000); E G. (SEM); E. external view of valve; F, G. external views of valve apices; H K. Fragilaria capucina var. radians (LM. 2,000).

73 Araphidineae: Diatomaceae: Fragilaria 65 species has two rimorportulae per valve, which are located in each pole. It is inconsistent with general description of the genus Fragilaria, which has one rimorpotula per valve (Krammer and Lange-Bertalot 1991). The species closely resembles Synedra rumpens Kützing, however, it differs in looser striae distribution. 4. Fragilaria capucina var. radians (Kützing) Lange-Bertalot 1991 (Fig. 7). Krammer and Lange-Bertalot 1991: 122. f. 109: 17, 18. BASIONYM: Synedra radians Kützing SYNONYM: Fragilaria capucina var. rumpens (Kütz.) Lange-Bertalot in Krammer and Lange-Bertalot Fragilaria capucina ssp. rumpens (Kütz.) Lange-Bertalot Cells linear, attenuated toward ends. Apices swollen, somewhat capitate. Pseudoraphe narrow. Central area usually with distinct transverse fascia, longer than broad; often valve appearing thicker at central area; not distinctly swollen. Striae parallel, in 10 μm. Length about 45 μm, breadth about 4 μm. TYPE: Lectotype BM SEASONALITY: All around year. DISTRIBUTION: Not clear. KOREA: This taxon is mainly collected from various dam lakes; Nakdong River around Changwon city (Lee 2002), Lake Imha, Lake Yungchun, Lake Unmun, Lake Andong, Lake Hapchun and Lake Chinyang (Lee and Kim 1996). SPECIMEN EXAMINED: LJ915 (Yungchun dam reservoir and its tributaries). ECOLOGY: Not clear. REMARKS: The specimens from the Namchun stream closely matched with the photographs of lectotype BM (18192), presented by Krammer and Lange-Bertalot (1991; f. 109: 17, 18). This taxon differs from F. capucina var. capucina by its coarser striae density and slightly more swollen valve margin in center. However, taxonomic distinctions between F. capucina var. radians and F. capucina var. diatans (Grunow) Lange-Bertalot, an earlier synonym of Synedra rumpens var. fragilarioides Grunow, are problematic (see, Van Heruck 1881: pl. 40. f. 12; Krammer and Lange-Bertalot 1991: f. 109: 16; f. 113: 16 21; Fukushima et al. 1991: pl. 1 15). Fig. 7. Distribution of Fragilaria capucina var. radians.

74 66 Algal Flora of Korea Freshwater Diatoms II 5. Fragilaria crotonensis Kitton 1869 (Figs. 8, 9). Patrick and Reimer 1966: 121. pl. 3. f. 11, 12. Krammer and Lange-Bertalot 1991: 130. f. 116: 1 4. BASIONYM: Fragilaria crotonensis Kitton C D F G E 10 μm A B Fig. 8. Fragilaria crotonensis. A, B. Fragilaria crotonensis; C. Fragilaria famelica; D, E. Fragilaria gracilis; F, G. Fragilaria mesolepta (LM. 2,000).

75 Araphidineae: Diatomaceae: Fragilaria 67 SYNONYM: Fragilaria smithiana Grunow in Van Heruck Synedra crotonensis (Kitton) Cl. and Möll Synedra crotonensis var. prolongata f. belgica Grunow in Van Heruck Fragilaria crotonensis var. prolongata Grunow in Van Heruck Frustules in girdle view linear, swollen at center, slender toward ends, attaching at middle portion to form ribbon-like filaments. Valve linear, distinctly swollen to lanceolate shape in middle portion, lesser extent, at somewhat capitate apices. Sternum indistinctive. Central area usually rectangular in shape, extending to margins of valve or to marginal striae. Rimoportula in valve pole. Striae parallel, in 10 μm. Valve length μm. Width in middle portion 2 4 μm. TYPE: BM type locality: U.S.A., Croton water, New York. SEASONALITY: Present throughout the year. DISTRIBUTION: Worldwide in temperate freshwater lakes. Fig. 9. Distribution of Fragilaria crotonensis. KOREA: This species frequently occurs in almost all reservoirs and lakes in Korea-Nakdong River (Kim and Lee 1991; Lee 2002; Cho et al. 1993; Shin 2004), Seomjin River in Imsil county (Lee and Yoon 1999), Guem River in Yeonggi county, Gongju city and Nonsan city (Kim et al. 2000), Imjin River (Lee and Yoon 2002a), Yungchun dam reservoir (Lee et al. 1992), Lake Chungju (Kim et al. 1998), Lake Janjwa (Lee and Yoon 2002b), Lake Unmun in Chungdo county (Lee and Kim 1996) and Nammaeji reservoir (Kim et al. 1993). SPECIMEN EXAMINED: LJ936 (Yungchun dam reservoir). ECOLOGY: The species is a cosmopolitan, alkaliphilous, mesotrophic or eutrophic planktonic taxon (Kobayasi et al. 2006). REMARKS: The species is in valve view, swollen, lanceolate at middle portion, which is the most distinctive characteristic (Patrick and Reimer 1966). 6. Fragilaria famelica (Kützing) Lange-Bertalot 1980 (Fig. 10). Krammer and Lange-Bertalot 1991: 129. f. 115: 1 5. Patrick and Reimer 1966: 139. pl. 5. f. 9. BASIONYM: Synedra famelica Kützing SYNONYM: Synedra minuscule Grunow in Van Heruck 1881.

76 68 Algal Flora of Korea Freshwater Diatoms II Frustules forming band, in girdle view linear, narrower toward ends. Valve linear to lanceolate, attenuated toward rostrate ends. Pseudoraphe narrow. Central area variable, not distinctly swollen. Striae parallel, not distinctly punctuate, in 10 μm. Valve length μm, width μm. TYPE: Herbar Kützing 179 (B. M, 18189), aus Halle. SEASONALITY: All around year. DISTRIBUTION: Not clear. KOREA: Low reaches of the Seomjin River. SPECIMEN EXAMINED: LJ1023 (the Seomjin River). ECOLOGY: The species prefers eutrophic water of high mineral content (Patrick and Reimer 1966). REMARKS: This taxon is characterized by its small size and fine striae (Patrick and Reimer 1966). 7. Fragilaria gracilis Ǿstrup 1910 (Fig. 11). Fig. 10. Distribution of Fragilaria famelica. Krammer and Lange-Bertalot 1991: 123. f. 110: f. 111: 1 3. f. 113: Lee et al. 1994: 19. pl. 1. f. 14. Watanabe 2006: 110. pl. IIA 10. Tuji BASIONYM: Fragilaria gracilis Ǿstrup SYNONYM: Synedra rumpens var. familiaris (Kützing) Grunow Synedra familiaris sensu auct nonnull. Synedra famelica sensu auct nonnull. Cells linear-lanceolate, narrowed toward subcapitates to acute apices. Central area very weakly present. Axial are present in LM or not distinct ( μm). Spines absent along valve margins, Rimoportula present in only on valve pole. Pore fields in both pole well developed. Striae ca. 20 in 10 μm. Valve length 63 μm. Width μm. TYPE: Lectotypus Coll. Østrup SEASONALITY: Present throughout the year. DISTRIBUTION: Worldwide occurrence in temperate freshwater. KOREA: This species occurs in various rivers and reservoirs-nakdong River (Lee 2002), Lake Imha (Kim et al. 1995), Somjin River around Imsil county (Lee and Yoon 1999), Imjin River around Yeoncheon county (Lee and Yoon 2002), Kwang River (Kwangchun), Ulchin county (Lee et al. 1994), Kumho River (Lee and Chung 1992), Tongwha stream in Daegu (Hong 1990), and Nam Stream (Namchun), in Kyungsan city (Jung 2008). SPECIMEN EXAMINED: LJ207 (Kwang river, Ulchin county, Kyungbuk). ECOLOGY: The species occurs on β-saproxenous, ph circumneutral and eutrophic areas.

77 Araphidineae: Diatomaceae: Fragilaria 69 REMARKS: The Fragialria capucina/vaucheriae species complex sensu Lange-Bertalot in Krammer and Lange- Bertalot (1991) may have representatives with cosmopolitan distribution (Patrick and Reimer 1966; Kobayasi et al. 2006; Watanabe 2006). In spite of it frequency of occurrence, the identification of the taxa is still problematic (Krammer and Lange-Bertalot 1991; Lee et al. 1991; Tuji 2004, 2007; Tuji and Williams 2006). Fragilaria gracilis was first described in 1910, and Hustedt (1950) transfered it to a variety of F. capucina. Krammer and Lange-Bertalot (1991) designated a lectotype slide and exhibited LM photograhps of the plants. However, F. gracilis has one rimoportula per valve, much denser striae, and a more slender valve shape against F. capucina having two rimoportulae on a valve and a lower density of striae. Therefore, F. gracilis may not be a variety of F. capucina but a distinct species (Tuji 2007). The taxon is distinguishable by the shape of the valve and fine striae from Flagilaria vaucheriae (Kütz.) Petersen. Also, it should be compared with Fragilaria famelica (Kütz.) Lange-Bertalot which has more coarse striae (Tuji 2007). Fig. 11. Distribution of Fragilaria gracilis. 8. Fragilaria mesolepta Rabenhorst 1861 (Fig. 12). BASIONYM: Fragilaria mesolepta Rabenhorst Patrick and Reimer 1996: 119. pl. 3. f. 6. Krammer and Lange-Bertalot 1991: 123. f. 110: 14 21, 23, 24. Chung et al. 1993: 52. Lee et al. 1994: 19. Lee and Kim 1996: 20. Tuji and Williams SYNONYM: Fragilaria subconstricta Oestrup Fragilaria tenuistriata Oestrup Cells linear to linear-lanceolate, constricted at central area. Valve somewhat attenuated, rostrate in apes. Pseudoraphe very narrow. Central area variable. Striae parallel, in 10 μm. Valve length μm. Width in narrowest portion in the middle of valve 2 4 μm. Rimoportula on stria in mantle closed to valve junction near valve ends. TYPE: Lectotype: slide 3616 in FH (England finder O37-0). SEASONALITY: Throughout the year. DISTRIBUTION: Worldwide occurrence in temperate freshwater. KOREA: This species occurs in various localities; Nakdong River (Lee 2002), Lake Imha (Kim et al. 1995), Lake Chinyang (Lee and Kim 1996), Kwang River (Kwangchun) in Ulchin county (Lee et al. 1994), Kumho River (Lee and Chung 1992), Banbyeon Stream (Banbyeonchun) in Cheongsong county

78 70 Algal Flora of Korea Freshwater Diatoms II (Kim 1993), Tongwha stream (Tongwhachun) in Daegu (Hong 1990), Nam stream (Namchun) in Kyungsan city (Jung 2008). SPECIMEN EXAMINED: LJ055 (Yungchun dam lake, Yungchun city, Kyungbuk). ECOLOGY: This is a saproxenous and alkaliphilous taxon. REMARKS: Fragilaria mesolepta was first described by Rabenhorst from the material in his Die Algen Eurpas exsciccata (Rabenhorst 1861: Decas , packed 1041; see Kociolek 2004). Later, Rabenhorst (1864) proposed a combination, Fraglaria capucina var. mesolepta (Rabenhorst) Rabenhorst for the species (Tuji and Williams 2008). However Krammer and Lange-Bertalot (1991) treated F. tenuistriata and F. subconstricta, to be very similar to F. mesolepta, as synonyms of the latter. Although the outer line of the valves of F. subconstricta and F. tenuistriata, are very similar, both species should be considered as different taxa, because of the different positions of the rimoportulae (Tuji and Williams 2008). The taxon is also distinguished from the other varieties among the species by a constriction in the middle portion of the valve. The correct name for this taxon Fig. 12. Distribution of Fragilaria mesolepta. at the species level is Fragilaria bipunctata Hempr. and Her., but F. mesolepta is the correct name as a variation (Patrick and Reimer 1966). 9. Fragilaria parva Tuji and D. M. Williams 2008 (Figs. 13, 14). Tuji and Williams 2008: 29. f Krammer and Lange-Bertalot 1991: 123. Patrick and Reimer 1966: 143. pl. 5. f. 20. BASIONYM: Synedra familiaris f. parva Grunow SYNONYM: Synedra familiaris Kütz Synedra familiaris f. parva Grun. in Van Heruck Synedra familiaris f. major Grun. in Van Heruck Synedra rumpens var. familiaris (Kütz.) Hustedt Fragilaria capucina var. gracilis (Ǿstrup) Hustedt 1950 sensu Krammer and Lange-Bertalot Cells linear-lanceolate, swollen on each side in central area. Apices somewhat capitates. Axial area linear. Central area longer than wide, somewhat swollen. Striae parallel, in 10 μm. Length μm. Width of valve in swollen area 3 5 μm. TYPE: Holotype: slide 2654 in Grunow collection in W.

79 Araphidineae: Diatomaceae: Fragilaria 71 SEASONALITY: Present throughout the year. DISTRIBUTION: Worldwide occurrence in temperate freshwater. KOREA: This species collected only from the Namchun stream around Kyungsan city in Kyungbuk province, Korea (Jung 2008). 10 μm D A B C E F Fig. 13. Fragilaria parva. A C. (LM. 2,000); D F. (SEM); D. external view of valve; E, F. external views of valve apices. Note that apical pore fields are present in both apices.

80 72 Algal Flora of Korea Freshwater Diatoms II SPECIMEN EXAMINED: LJ1112 (Namchun stream around Kyungsan city, Kyungbuk). ECOLOGY: This species is widely distributed in freshwater lakes, or ponds, or slow-flowing streams. They are saproxenous and alkaliphilous taxa. REMARKS: This taxon is distinguished by central swellings of valve or the finer striae (Patrick and Reimer, 1966). Synedra familiaris Kütz., a synonym of Fragilaria parva Tuji and D.M. Williams, is an important taxon in ecological studies for freshwater attaching algal communities, because it appears to be cosmopolitan, is reported from various localities (Hustedt 1930; Patrick and Reimer 1966; Krammer and Lange- Bertalot 1991; Watanabe et al. 2005; Tuji and Williams 2008). S. amiliaris was originally described by Kützing (1844). Later, Grunow (in Van Heruck 1881) added two new varieties to this taxon: S. familiaris f. parva and S. familiaris f. major. Hustedt (1930) classified S. familiaris as a variety of S. rumpens (Kütz.) Carlson, which was subsequently followed by several diatomists. In fact, it may be impossible to detect morphological Fig. 14. Distribution of Fragilaria parva. characteristics of Synedra familiaris originally described by Kützing, because his original figures are too simple. According to examinations of the lectotype (slide BM 18307), designated after Kützing s material (Kützing s collection no. 1370) under the name of Synedra familiaris by Tuji and Williams (1988), there is no specimen fitted to the current definition and description of Synedra familiaris. They observed only three possible taxa that might be compared with S. familiaris. The first taxon might be identified as Fragilaria vaucheriae (Kütz.) Petersen, the second as Tabularia fasciculate (Agardh) Williams and Round, and the third as Ctenophora pulchella (Kütz.) Williams (Tuji and Williams 2008). There, basionyms of the three species were described before S. familaris. Therefore, the latter should be a later synonym of any one of the three taxa. The name S. familiaris is invalid. Accordiong to the observation on the holotype of Synedra familiaris f. parva Grun. by Tuji and Williams (2008), it was well congruent with S. familiaris sensu auct nonnull. Thus, Tuji and Williams (2008) made a new combination for the taxon as Fragilaria parva with the basionym Synedra familiaris f. parva Grun. in Van Heruck Fragilaria rumpens Kützing 1844 (Figs. 15, 16). Patrick and Reimer 1966: 143. pl. 5. f. 19. Krammer and Lange-Bertalot 1991: 122. f. 108: 16 21; f. 110: 1 6A. Tuji and Williams BASIONYM: Synedra rumpens Kütz., Bacill. 69. pl. 16. f. IV. 4, 5. SYNONYM: Fragilaria capucina var. rumpens (Kütz.) Lange-Bertalot in Krammer and Lange-Bertalot

81 Araphidineae: Diatomaceae: Fragilaria Fragilaria capucina ssp. rumpens (Kütz.) Lange-Bertalot E A B C 10 μm D F G H Fig. 15. Fragilaria rumpens. A D. (LM. 2,000); E H. (SEM). E, F. internal views of valves; G, H. internal views of valve apices showing apical pore fields. Note that a rimoportula is present on only one apex.

82 74 Algal Flora of Korea Freshwater Diatoms II Cells lanceolate, fusiform, Length μm long, width 3 4 μm. Central area with wide transverse fascia, sometimes unilaterally to bilaterally gibbous. Striae parallel throughout, in 10 μm. Colonies ribbon-like, adhering by their valve faces. Pervalver axis thick at central area and thin at ends, cells sometimes separated from each other at the poles. Spines irregular, located on costae, at mantle-valve face junction. Rimorpotulae present eccentric, adjacent to sternum, one per valve, apical pore fields rectangular. TYPE: Holotype Kützing packet 194 in AWH. SEASONALITY: Present throughout the year. DISTRIBUTION: Worldwide occurrence in temperate freshwater. KOREA: Here, this species occurs in streams, dam lakes and ponds; Nakdong River around Andong (Lee 2002), estuary of Nakdong River (Kim and Lee 1991), Lake Chuam (Choi et al. 1994), Lake Imha Lake (Kim et al. 1997), Somjin River (Lee and Yoon 1999), Imjin River (Lee and Yoon 2002), Kumho River (Lee and Chung 1992), Tongwha Stream in Daegu (Hong 1990). SPECIMEN EXAMINED: LJ1008 (Seomjin river, Sunchang county, Jeonbuk 2008). Fig. 16. Distribution of Fragilaria rumpens. ECOLOGY: The species is widely distributed in freshwater lakes, or ponds, or slow-flowing streams. They are saproxenous and alkaliphilous. REMARKS: The taxon is presumed to be a cosmopolitan and common tychoplanktonic-benthic species, reported from many freshwater lakes, or ponds, or slow flowing streams (Patrick and Reimer 1986). Yet the species is very similar to both, Fragilaria capucina Desm. and Fragilaria vaucheriae (Kütz.) Petersen, and its taxonomic characteristics have been confusing (Tuji and Williams 2006b). Krammer and Lange-Bertalot (1991) examined the type materials of these taxa, and considered F. rumpens to be a part of the F. capicina complex. Although the new combination Fragilaria capucina var. rumpens (Kütz.) Lange-Bertalot was proposed, it is not validly published as no basionym was designated (Tuji and Williams 2006b). Later, Lange-Bertalot (1993) proposed a different combination, Fragilaria capucina ssp. rumpens (Kütz.). Lange-Bertalot, in the revised edition of Krammer and Lange-Bertalot (1991, 2000), were using the specific name Fragilaria rumpens (Kütz.) Carlson for this taxon (Tuji and Williams 2006b). Fragilaria vaucheriae is easily distinguished from F. rumpens and F. capucina by its form and lower striae density. There are also several differences between F. rumpens and F. capucina. F. rumpens has striae in 10 μm, and that is denser than both, F. capucina (15 16 striae in 10 μm) and F. vaucheriae (9 14 striae in 10 μm). Valve poles of F. capucina are not as rostrate as those of F. rumpens. F. capucina has two small rimoportulae (Tuji and Williams 2006a). These are located at the poles, whereas F. rumpens and F. vaucheri-

83 Araphidineae: Diatomaceae: Fragilaria 75 ae have only one per valve (Tuji and Williams 2006b). The simplest way to distinguish F. rumpens from both species F. capucina and F. vaucheriae is in girdle view, because the frustules of F. rumpens are much thicker at the center than at the poles, and the colonies of F. rumpens are sometimes separated from each other at their ends, similar to the related species, Fragilaria crotonensis Kitton (Tuji and Williams 2006b). Colonies of F. capucina and F. vaucheriae differ from F. rumpens; the depth of frustules in the former two taxa is equal at both central parts and poles of valve, and adjacent valves are completely attached by linking spines from pole to pole (Tuji and Williams 2006b). Many variaties have been described for F. rumpens (VanLandingham 1978; Patrick and Reimer 1966). Most are not found as plankton but as periphyton. These varieties do not form ribbon-like colonies, but occur as either single cells or in rosette-like colonies. This is the most probable reason that these taxa should not be accepted as varieties of F. rumpens, and further studies are required to clarify their taxonomic relation (Tuji and Williams 2006b). 11. Fragilaria vaucheriae (Kützing) Petersen 1938 (Figs 17, 18). Krammer and Lange-Bertalot 1991: 124. f. 108: Lee et al. 1992: 49. f. 23. Lee, Chung and Gotoh 1994: 19. Kawashima and Kobayasi 1994: 13. f. 4A K. Kobayasi et al. 2006: 61. pl. 78. BASIONYM: Exilaria vaucheriae Kützing Linnaea 8: 560. pl. 15. f. 38. SYNONYM: Staurosira intermedia Grunow Fragilaria intermedia Grunow in Van Heruck Synedra rumpens var. meneghiniana Grunow in Van Heruck Fragilaria capucina var. vaucheriae (Kütz.) Lange-Bertalot Frustules usually in short to fairly long chains, occasionally occurring separate. Valve linear to linearlanceolate, narrowed toward rostrate rounded apices. Pseudoraphe narrow. Central area usually on only one side of valve. Rimoportula in only one valve pole. Striae parallel to slightly radiate, occasionally slightly shortened opposite to central area. Striae 9 14 in 10 μm. Valve length μm. Width 4 5 μm. TYPE: Type slide: BM SEASONALITY: Present throughout the year. DISTRIBUTION: Worldwide occurrence in temperate freshwater. KOREA: This species is mostly collected from streams and standing waters in Korea; Nakdong River (Lee Fig. 17. Distribution of Fragilaria vaucheriae.

84 76 Algal Flora of Korea Freshwater Diatoms II A 10 μm B C D E F G H Fig. 18. Fragilaria vaucheriae. A C. (LM. 2,000); D H. (SEM); D. internal views of valves; E, F. internal views of valve apices showing a rimoportula; G, H. extenal views of valve apices.

85 Araphidineae: Diatomaceae: Hannaea ; Cho et al. 1993; Shin 2004), Lake Imha (Kim et al. 1995; Kim et al. 1997; Kim 1993), Lake Chinyang (Lee and Kim 1996), Kwang River (Kwangchun) in Ulchin county (Lee et al. 1994), Kumho River (Lee and Chung 1992; Park 1995), Yungchun dam reservoir (Lee et al. 1992), Banbyeon Stream (Banbyeonchun) in Cheongsong county (Kim 1993), Tongwha stream (Tongwhachun) in Daegu (Hong 1990), Milyang river (Choi and Chung 1995), Nammaeji reservoir (Kim et al. 1993) and Nam stream (Namchun) in Kyungsan city (Jung 2008). SPECIMEN EXAMINED: LJ218 (Kwang river, Ulchin county, Kyungbuk 1992). ECOLOGY: The species is saproxenous and alkaliphilous. REMARKS: The taxon is distinguished by its asymmetrical, usually slightly swollen central area. Genus Hannaea Patrick Cells are arcuate. Sometimes they are free living but often in cluster or short colonies, attaching to stones in freshwater streams and springs especially in mountainous areas (Round et al. 1990). In valve view the dorsal margin is convex and the ventral margin is concave, but swollen on each side of the central area, which is also swollen or smoothly concave except for the swelling of the central area (Patrick and Reimer 1966). The frustules are asymmetrical to the apical axis, and symmetrical to the transverse axis (Patrick and Reimer 1966). Areolae are small, round, and poroidal. Rimoprotulae are very distinct internally, one at each pole but occasionally lacking at one end, and copulae are split with a row of areolae (Round et al. 1990). Patrick and Reimer (1966) first proposed a new name, Hannaea, genus instead Ceratoneis is the taxon accomodating only two species. These species were originally included in Ceratoneis, but are now allocated to Nitzschia and Pleurosigma, making another name necessary (Round et al. 1990). The shape of the cell is almost the only feature distinguishing Hannaea from Fragilaria, and the former genus can be maintained only on its distinctive morphological characteristic and possibly its very restricted ecological amplitude (Round et al. 1990). Type species: H. arcus (Ehrenberg) Patrick. 12. Hannaea arcus var. recta (Cleve) M. Idei 2006 (Figs. 19, 20). Kobayasi et al. 2006: 64. pl. 81. BASIONYM: Fragilaria arcus var. recta Cleve SYNONYM: Fragilaria awqualis var. inaequidentata Lagerstedt Ceratoneis recta (Skvortzow and Meyer) Iwahashi Ceratoneis arcus var. linearis f. recta (Skvortzow and Meyer) Proschkina-Lavrenko in Sabelina et al Frustules in girdle view slightly bent, forming short bands. Valve very slightly curved to almost stright, with slightly convex dorsal margin. Ventral margin almost straight except the swelling of

86 78 Algal Flora of Korea Freshwater Diatoms II the unilateral central area. Apices of valve attenuaterostrate to somewhat capitate. Central area distinctly swollen, only on ventral side of pseudoraphe. Pseudoraphe distinct, narrow. Striae parallel to slightly radiate toward valve apices. Striae in 10 μm. Valve length μm. Width 6 7 μm. TYPE: Not clear. SEASONALITY: Present throughout the year. DISTRIBUTION: Distributed streams in Korea and Japan. KOREA: This taxon is rare and collected a few streams; Nam stream (Namchun) in Kyungsan city (Jung 2008). SPECIMEN EXAMINED: LJ1124 (Nam stream Namchun in Kyungsan city). ECOLOGY: The species is saproxenous (Kobayasi et al. 2006). REMARKS: This taxon differs from H. arcus by its almost straight shape of the valve margin (Kobayasi et al. 2006). Fig. 19. Distribution of Hannaea arcus var. recta. 13. Hannaea arcus var. subarcus (Iwahashi) Lee 1992 (Figs. 20, 21). Lee et al. 1992: 50. pl. 2. f. 34. Lee et al. 1994: 19. BASIONYM: Ceratoneis recta (Skvortzow and Meyer) Iwahashi f. subarcus Iwahashi SYNONYM: None. Frustules in girdle view slightly bent, forming short bands. Valve slightly curved, with slightly convex dorsal margin. Ventral margin a little concave except for swelling of unilateral central area. Apices of valve attenuate-rostrate to somewhat capitate. Central area distinctly swollen, only on ventral side of the pseudoraphe. Pseudoraphe distinct, narrow. Striae parallel to slightly radiate toward apices of valve. Striae in 10 μm. Valve length μm. Width μm. TYPE: Not clear. SEASONALITY: Presentl throughout the year. DISTRIBUTION: Known from a few unpolluted streams in Korea and Japan. KOREA: This taxon mainly occurs in the Nakdong River system in Korea; Lake Imha (Kim et al. 1995; Kim et al. 1997; Kim 1993), Kwang River (Kwangchun) in Ulchin county (Lee et al. 1994), Shinchun river in Daegu (Choi et al. 1993), Yungchun dam reservoir (Lee et al. 1992), Banbyeon Stream (Banbyeonchun) in Cheongsong county (Kim 1993), Tongwha stream (Tongwhachun) in

87 Araphidineae: Diatomaceae: Hannaea 79 E C A F G H B D 10 μm I Fig. 20. Hannaea arcus var. subarcus. A, B. Hannaea arcus var. recta; C, D. Hannaea arcus var. subarcus; E. Pseudostaurosira brevistriata; F H. Punctastriata linearis. (LM. 2,000); I. external valve view of Punctastriata linearis showing connecting spines (SEM).

88 80 Algal Flora of Korea Freshwater Diatoms II Daegu (Hong 1990), Milyang river (Choi and Chung 1995), Nammaeji reservoir (Kim et al. 1993) and Nam stream (Namchun) in Kyungsan city (Jung 2008). SPECIMEN EXAMINED: LJ121 (Jaho river, Yungchun city, Kyungbuk). ECOLOGY: The species is saproxenous. REMARKS: This taxon fits well with the description and the illustration of Ceratoneis recta f. subarcus. According to Skvortzow and Meyer (1928), Ceratoneis recta f. recta has almost a straight shape of the valve margin. It differs from C. recta f. recta only by the slightly curved valve margin. Therefore, should these taxa belong to Ceratoneis arcus (Ehrenberg) Kützing, as varieties. However, as proposed by Patrick (in Patrick and Reimer 1966), the genus Hannaea should be used instead of the name Ceratoneis. Fig. 21. Distribution of Hannaea arcus var. subarcus. Genus Pseudostaurosira Williams and Round 1987: 275. Cells joined tightly to form filaments. Plastids probably parietal and plate-like. Frustules rectangular, forming chains. Valves elliptical to linear, often undulate, occasionally cruciform. Striae uniseriate consisting of a few large elliptical areolae and often smaller round areolae. Sternum wide. Apical pore fields not always present. Spines situated along the valve edge. This genus differs from Staurosira by the nature of the striae, the shape of the central area and the small size of the pore fields (Williams and Round, 1987). The most characteristic feature of this genus are sparse marginal areolae (Round et al. 1990). Type species: P. brevistriata Williams and Round. 14. Pseudostaurosira brevistriata (Grunow) Williams and Round 1987 (Fig. 22). Williams and Round 1987: 276. Krammer and Lange-Bertalot 1991: 162. f. 130: Kobayasi et al. 2006: 67. pl. 85. BASIONYM: Fragilaria brevistriata Grunow in Van Heruck

89 Araphidineae: Diatomaceae: Punctastriata 81 SYNONYM: Fragilaria brevistriata var. subacuta Grunow in V. H Fragilaria brevistriata var. pusilla Grunow in V. H Fragilaria brevistriata var. subcapitata Grunow in V. H Frustules rectangular, forming chains. Valves slightly cruciform, 2 5 μm in breadth, μm in length. Closing plate delicate, highly branched. Rimoportulae absent. Mantle plaques present along mantle edge. Spines situated along valve edge usually in somewhat plain region of valve, enlarged at tips. Striae uniseriate consisting of a few ovoid areolae, no more than four per striae at centre, less at poles, in 10 μm. TYPE: BM SEASONALITY: All around year. DISTRIBUTION: Worldwide occurrence in standing waters. KOREA: This taxon is rare in Korea. SPECIMEN EXAMINED: LJ1009 (Seomjin river, Jeonnam). ECOLOGY: The species occurs in oligo-saprobic, oligo- to mesoeutrophic water with low electric conductivity (Krammer and Lange-Bertalot 1991). Fig. 22. Distribution of Pseudostaurosira brevistriata. REMARKS: This taxon is distinguished from the other species belonging to Pseudostaurosira by its fine marginal striae (Patrick and Reimer 1966). Genus Punctastriata Williams and Round 1987: 278. Cells are forming short filaments to branching clustersin fresh waters. Valves are linear-elliptical and sometimes with central inflation. Valves are heteropolar. Striae are multiseriate, lying between raised ribs and curving onto valve mantle. Sternum is narrow. One apical pore field is present, usually difficult to detect. Rimoportulae are absent. This genus is quite difficult to identify. It has been identified as Fragilaria, or as Opephora in the literature (Williams and Round 1987). Type species: P. linearis Williams and Round. 15. Punctastriata linearis Williams and Round 1987 (Fig. 23). Williams and Round 1987: 278. Kawashima and Kobayasi, 1994: 17. f. 7A G. Idei and Nagumo

90 82 Algal Flora of Korea Freshwater Diatoms II 1995: 238. f. 1m, n, o, f. 6. Kobayasi et al. 2006: 68. pl. 88. BASIONYM: Punctastriata linearis Williams and Round SYNONYM: Fragialria pinnata sensu Krammer and Lange-Bertalot 1991 part. Valves linear to elliptical, 3 6 μm in width, 5 32 μm in length. Striae composed of multiseriate areolae, 8 10 in 10 μm. Spines situated along valve face/mantle junction on interstriae that are crossing the striae, spines short, pointed, usually paired. Single apical pore field present. TYPE: BM SEASONALITY: Not clear. DISTRIBUTION: May be cosmopolitan. KOREA: This taxon is very rare and new to Korea. SPECIMEN EXAMINED: LJ1010 (Seomjin river, Jeonnam). ECOLOGY: Not clear. REMARKS: This genus is quite difficult to identify by light microscopy, and even at SEM level (Williams and Round 1987). It has been identified as Fragilaria pinnata Fig. 23. Distribution of Punctastriata linearis. (Krammer and Lange-Bertalot 1991: f. 133: 1 18), Fragilaria pinnata var. lanccetula, or as a species of the genus Opephora in the literature (Williams and Round 1987). With this difficulty, it is impossible to trace original names that have been used to describe it, without an SEM study of all relevant material (Williams and Round 1987). This taxon is the type species of the genus Punctastriata Williams and Round. This species is very similar to Staurosirella pinnata (Ehrenberg) Williams and Round. It is very difficult to distinguish both taxa by light microscopic observation. In LM the striae of Punctastriata linearis are widened towards the valve margin, while those of S. pinnata are almost same in width from the valve center to the margin. Significant feature of P. linearis are the striae, which are composed of multi-seriate areolae, they are an important characteristic of the genus Punctastriata and are only observed here (Idei and Nagumo 1995). Genus Staurosira Ehrenberg 1843: 45. Cells are solitary to form straight zig-zag filaments with a few to many cells attached to one another. They are attached to the substratum but often are found in a free-living state. Frustules are rectangular in girdle view. Valves are elliptical and expanded centrally. Sternum is narrow except in the centre. Striae are uniseriate. Areolae are elliptical to linear, and vela are present. Apical pore fields are present, but usually greatly reduced. No rimoportulae are present, and

91 Araphidineae: Diatomaceae: Staurosira 83 plastids become arranged in two large plates. This genus, formerly in Fragilaria, differs from Fragilaria by the absence of Rimoportulae, nonareolate copulae, wide valvocopulae, and relatively narrow copulae (Round et al. 1990). Staurosira differs from Martyana since the latter has no marginally linking spines, but has an apical pore field at one end only, a step in the valve at the other end, and a silt areole (Round et al. 1990; Idei and Nagumo 1995). Type species: S. construens Ehrenberg. 16. Staurosira construens Ehrenberg 1843 (Figs. 24, 25). Patrick and Reimer 1966: 125. pl. 4, f. 4. Williams and Round 1987: f Kawashima and Kobayasi 1994: 8A. Idei and Nagumo 1995: f. f, g. Kobayasi et al. 2006: 71. pl. 89. BASIONYM: Staurosira construens Ehrenberg SYNONYM: Odontidium tabellaria W. Smith Fragilaria construens (Ehrenberg) Grunow Frustules rectangular in girdle view. Valves strongly swollen in middle portion, often somewhat asymmetrical, rounded at the ends, sometimes slightly capitate. Sternum distinct, linear to linearlanceolate. Striae radiate throughout most of the valve, crossed by fine lines. Striae in 10 μm. Areolae elliptical to linear with simple cribra. Apical pore fields present, consisting of 4 8 pores. Spines interstriae, broadening outward, bigger, especially in the central part, Rimoportulae absent. TYPE: Type locality: Newhaven in Nord-Amerika. SEASONALITY: Present throughout the year. DISTRIBUTION: Occurring worldwide. KOREA: This species occurs in various localities; Nakdong River (Kim and Lee 1991; Lee 2002; Cho et al. 1993), Kumho River (Park 1995), Tongwha stream (Tongwhachun) in Daegu (Hong 1990), Milyang river (Choi and Chung 1995), Guem River (Kim et al. 2000) and Nam stream (Namchun) in Kyungsan city (Jung 2008). SPECIMEN EXAMINED: LJ275 (Nakdong River). ECOLOGY: The species is meso-saprobic (Kobayasi et al. 2006). REMARKS: This taxon is the type species of the genus Staurosira. Characteristics of this species are many cingulum bands, smaller apical pore fields, and simple pore-shaped areolae (Kobayasi et al. 2006). It Fig. 24. Distribution of Staurosira construens.

92 84 Algal Flora of Korea Freshwater Diatoms II is very similar to Fragilaria leptostauron in shape, but differs in having much finer striae. This species is highly variable in shape (Patrick and Reimer 1966). 10 μm A B C D E F G H Fig. 25. Staurosira construens. A D. (LM. 2,000); A. Staurosira construens; B H. Staurosira construens var. binodis. E H. (SEM); E. external valve view; F. internal valve view; G, H. copulae showing cingulum bands and connecting spines.

93 Araphidineae: Diatomaceae: Staurosira Staurosira construens var. binodis (Ehrenberg) Hamilton 1992 (Fig. 26). Patrick and Reimer 1966: 125. pl. 4. f. 7. Lee et al. 1992: 49. pl. 2. f. 28. Hamilton et al. 1992: 29. Lee et al. 1994: 19. Morales 2005: 118. f , Kobayasi et al. 2006: 72. pl. 90. BASIONYM: Fragilaria binodis Ehrenberg SYNONYM: Fragilaria construens var. binodis Grunow Fragilaria subconstricta Oestrup Fragilaria tenuistriata Oestrup Fragilaria construens f. binodis (Ehrenberg) Hustedt Frustules rectangular with undulate relief in girdle view. In valve view, valves linear to constricted in the middle portion with rostrate to subrostrate ends. Pseudoraphe lanceolate, variable in width. Central area not clearly differentiated from axial area. Striae composed by narrow slit-like areolae, continual from valve face to valve mantle. Costae thickened, somewhat raised with respect to striae. Apical pore fields composed of round poroids located on valve mantle. Rimoportula absent. Striae dense, in 10 μm. Valve length μm. Width at widest portion 4 6 μm. Fig. 26. Distribution of Staurosira construens var. binodis. TYPE: Not clear. SEASONALITY: Present throughout the year. DISTRIBUTION: Occurring worldwide in temperate freshwaters. KOREA: This taxon is collected from most localities in Korea; Nakdong River (Lee 2002; Shin 2004), Lake Imha (Kim et al. 1995), Kwang River (Kwangchun) in Ulchin county (Lee et al. 1994), Kumho River (Lee and Chung 1992; Park 1995), Yungchun dam reservoir (Lee et al. 1992), Banbyeon Stream (Banbyeonchun) in Cheongsong county (Kim 1993), Tongwha stream (Tongwhachun) in Daegu (Hong 1990), and Nam stream (Namchun) in Kyungsan city (Jung 2008). SPECIMEN EXAMINED: LJ991 (Nakdong river, Milyang city, Kyungnam). ECOLOGY: The species is oligosaprobous, meso-eutraphentic, and alkaliphilous (Van Dam et al. 1994). REMARKS: Hamilton (1992) validly made a new combination with the name Staurosira construens var. binodis (Hamilton et al. 1992), and there are some other name combinations (Morales 2005). Bukhtiyarova (1995) published the same combination as Hamilton, but is invalid under the rule of priority by international Code of Botanical Nomenclature. Edlund (1994) made another combination, Pseudostaurosira construens var. binodis (Ehrenberg) Edlund. However, the author himself understood the errata, and stated that it was an invalid combination, because the name Pseudostau-

94 86 Algal Flora of Korea Freshwater Diatoms II rosira construens was alreadly existing (Morales 2005). Pseudostaurosira robusta (Fusey) Williams and Round is very similar in its outer appearance to S. construens var. binodis, however, the areolar structure and some micro-characteristics under SEM show that P. robusta should be included in the genus Pseudostaurosira (Williams and Round 1987). This taxon is characterized by a constriction in the middle portion of the valve. Van Dam et al. (1994) and Watanabe (2006) reported the taxon to be alkaliphilous and mesoeutraphentic. 18. Staurosira elliptica (Schumann) Williams and Round 1987 sensu auct. nonnull. (Figs. 27, 28). Williams and Round 1987: 272. f Krammer and Lange-Bertalot 1991: 155. f. 130: Morales 2005: 114. f Edlund et al Kobayasi et al. 2006: 75. pl. 93. BASIONYM: Fragilaria elliptica Schumann SYNONYM: Fragilaria construens var. pumila Grunow in Van Heruck Fragilaria construens f. subsalina (Hustedt) Hustedt, Staurosira construens f. subsalina (Hustedt) Bukhtiyarova, Sturosira construens var. subsalina (Hustedt) Andersen, Stoermer and Kreis. Fragilaria elliptica Schumann sensu Lange-Bertalot 1991 and sensu auct nonnull. Frustules rectangular in girdle view, forming chains with connecting spines. Valves lanceolate with acute to broadly lanceolate or even round apices in valve view. Sternum linear, very narrow. Rimoportulae absent. Striae parallel, clearly punctuate under LM, slightly radiate near valve apices. The areolae broadly round. Apical pore fields more or less developed, positioned at valve face/ mantle junction. Striae in 10 μm, composed of areola in 10 μm. Valve length 6 22 μm. Width 3 5 μm. TYPE: Not clear. SEASONALITY: All around the year. DISTRIBUTION: Worldwide in temperate freshwaters. KOREA: This taxon occurs in various rivers and standing waters-nakdong River (Lee 2002; Shin 2004), Milyang River (Choi and Chung 1995), Guem River (Kim et al. 2000) and Kwang River (Kwangchun) in Ulchin county (Lee et al. 1994). SPECIMEN EXAMINED: LJ1007 (Seomjin river, Sunchang county, Jeonbuk 2008). ECOLOGY: The species is meso-saprobous (Kobayasi et al. 2006). REMARKS: Our specimens accord well with the characters of F. elliptica sensu auct nonnull. (excl. F. elliptica Schumann in type slide (ANSP G.C ), shown by the material from Schumann s collection in ANSP). Staurosira elliptica was described as Fragilaria elliptica in 1867 from Prussian freshwater material collected near Königsberg. Schumann (1867) provided a very short description and a single illustration of his new taxon (Edlund et al. 2006). The illustration showed an elliptic valve with a broad axial area and clearly punctuate and radiate striae. Van Heruck ( ) described Fragilaria elliptica and its form minors with a narrower axial area and parallel striae compared to Schumann s illustration. Haworth (1975) reported some specimens

95 Araphidineae: Diatomaceae: Staurosira 87 A B C 10 μm D E F G H I Fig. 27. Staurosira elliptica. A C. (LM. 2,000); D I. (SEM); D G. external views of various valves showing simple openings of areolae and connecting spines; H. internal valve view with cribra; I. frustules.

96 88 Algal Flora of Korea Freshwater Diatoms II from Scottish lake sediments identified as F. elliptica with narrow axial area, round to dash-like shape of the areolae, no apical pore field, and flat spines on each interstria (Edlund et al. 2006). Then, Williams and Round (1987) proposed to transfer F. elliptica to the genus Staurosira Ehrenberg, as S. elliptica (Schumann) Williams and Round. This general description for F. elliptica or S. elliptica has been accepted by other authors (Edlund et al. 2006). Especially Krammer and Lange-Bertalot (1991) illustrated a more variable morphotype of F. elliptica. They showed various morphotypes of parallel to radiate, coarsely punctuate striae, oval, elliptical, and lanceolate valve shapes. However, in all of the previous reports, no analysis on Schumann s (1867) original slides or material was attempted (Edlund et al. 2006). F. elliptica and S. elliptica, having a narrow axial area and a simple round outer opening of areolae, generally reported in various literatures (Williams and Round 1987; Krammer and Lange-Bertalot 1991; Kawashima and Kobayasi 1992; Kobayasi et al. 2006), do not correspond to the typical features of F. elliptica by Schumann s type material observation (Edlund et al. 2006). F. elliptica in Schumann s type material has a wide axial area and Fig. 28. Distribution of Staurosira elliptica. areolae heavily occluded by 2 5 dichotomously branching volae (Edlund et al. 2006). And its striae are composed of two areolae: one the valve face, and the other on valve mantle (see, Edlund et al. 2006: f ). Thus, Edlund et al. (2006) made a nomenclatural combination of Pseudosturosira elliptica with the basionym, Fragilaria elliptica (Schumann 1867). Therefore, there is no valid taxonomical name based on International Code of Botanical Nomenclature for Fragilaria elliptica Schumann non sensu Schumann, sensu auct nonnull, or Staurosira elliptica (Schumann) Williams and Round, sensu auct nonnull. The specimens in this monograph are similar to Fragilaria construens var. subsalina Hustedt, which was transferred to the genus Pseudostaurosira, as P. subsalina (Hust.) Morales by Morales (2005). Especially Morales stated that P. subsalina has been broadly misidentified as Fragilaria elliptica Schumann, or Sturosira elliptica (Schumann) Williams and Round. 19. Staurosira venter (Ehrenberg) H. Kobayasi 2002 (Figs. 29, 30). Krammer and Lange-Bertalot 1991: 153. f. 132: 1 34; f. 129: 21 27; f. 131: 5, 6. Kawashima and Kobayasi 1992: 17. f. 8B F. Mayama et al. 2002: 90. Kobayasi et al. 2006: 76. pl. 94. Paull et al BASIONYM: Fragilaria venter Ehrenberg 1854: pl. 14. f. 50. SYNONYM: Fragilaria construens var. venter (Ehrenberg) Grunow 1881.

97 Araphidineae: Diatomaceae: Staurosira μm D E A B C F G H Fig. 29. Staurosira venter. A C. (LM. 2,000); D, E. (SEM); D. external valve view; E. internal valve view; F H. Staurosirella pinnata (LM. 2,000). Fragilaria construens var. pumila Grunow Fragilaria construens f. venter (Ehrenberg) Hustedt Cells variable in shape, linear-lanceolate, to rhombic, rostrate in apices, somewhat concave at the margins, tapering rounded at end. Rimoportulae absent. Pore fields at both poles. Sternum linear, widens towards the central area. Striae almost parallel but slightly radiate next to the pole, in 10 μm. Valve length μm. Width 5 7 μm. TYPE: Not clear. SEASONALITY: Present throughout the year. DISTRIBUTION: Occurring worldwide in temperate freshwaters. KOREA: This species occurs predominately in many rivers and streams in Korea-Nakdong River (Lee 2002; Cho et al. 1993; Shin 2004), Shinchun River in Daegu (Choi et al. 1993), Lake Imha (Kim et al. 1995; Kim et al. 1997; Kim 1993), Kwang River (Kwangchun) in Ulchin county (Lee et al. 1994), Kumho River (Lee and Chung 1992; Park 1995), Tongwha stream (Tongwhachun) in Daegu (Hong 1990) and Nam stream (Namchun) in Kyungsan city (Jung 2008). SPECIMEN EXAMINED: LJ995 (Nakdong river, Milyang city, Kyungnam 2008). ECOLOGY: The species is β-mesosaprobous, meso-eutraphentic, and alkaliphilous (Van Dam et al. 1994).

98 90 Algal Flora of Korea Freshwater Diatoms II REMARKS: This species is highly variable in shape, and includes a vast number of varieties. Staurosira venter differs from the nominated variety by its less swollen central valve margin and its almost parallel striae distribution (Kawashima and Kobayasi 1992). According to Williams and Round is especially Staurosirella pinnata (Ehrenb.) very similar to this species. Generally have fragilarioid diatoms been used as indicators for changes of nutrient levels, salinity, and ph (Lowe 1974; Van Dam et al. 1994; Watanabe 2006). Although Stuarosirella pinnata and Staurosira venter are often used as such environmental indicators, studies of their assemblages sometimes produce conflicting evidences by high negative correlations of both species abundances (Paull et al. 2008). Staurosira venter and Staurosirella pinnata have been classified as taxa of the genus Fragilaria, on the basis of colonial form and characteristic of attachment. Although enhanced observation technology and more detailed studies of the life cycle have provided much Fig. 30. Distribution of Staurosira venter. new information on the morphology of the genus Fragilaria (sensu lato), the identification of small fragilaroid forms, such as Staurosira venter, Pseudostaurosira elliptica (Schumann) Edlund, and Staurosirella pinnata, can still be problematic (Paull et al. 2008). Genus Staurosirella Williams and Round 1987: 274. Frustules are rectangular, attached and in chains either linking at corners or by valve face to face contact. Valves are elliptical to linear, occasionally cruciform. Striae are uniseriate and containing slit-like areolae. Sternum is wide. Apical pore fields are usually large. Rimoportulae are absent. Spines are often complexly branched. This genus encompasses the previous Fragilaria species. It differs from Staurosira in the nature of the areolae, apical pore plates, and the structure of spines. The thick ribs separating the areolae are a very distinctive characteristic of this taxon (Round et al. 1990). Type species: S. lapponica Williams and Round. 20. Staurosirella pinnata (Ehrenberg) Williams and Round 1987 (Fig. 31). Williams and Round 1987: 274. Kobayasi et al. 2006: 80. pl. 98. Krammer and Lange-Bertalot 1991: 156. f. 131: 3, 4; Tafel 133.

99 Araphidineae: Diatomaceae: Staurosirella 91 BASIONYM: Fragilaria pinnata Ehrenberg 1843: 415. pl. 3. f. 8. SYNONYM: Odontidium mutabile W. Smith Fragilaria pinnata var. lancettula (Schumann 1867) Hustedt in A, Schmidt et al Fragilaria pinnata var. subrotunda Mayer Odontidium martyi var. polymorpha (Jouravleva) Proschkina-Lavrenko Frustules linear-rectangular, almost square in girdle view. Valve elliptical to linear with rounded apices. Rimoportulae not present. Pseudoraphe narrow, sometimes widened to a small lanceolate space at the center of valve. Striae almost parallel, radiate near valve apices, crossed by fine lines. Striae in 10 μm. Valve length 5 17 μm. Width 3 6 μm. Apical pore fields variable in the degree of development, always composed of several rows of round areolae disposed in ordered rows, heteropolar in form, unevenly developed as one at the foot pole always being larger. Spines developed (single or paired) or absent, located on costae towards the valve face-mantle junction, spatulate, sometimes dichotomously branched, providing a firm grip between neighboring valves that allows the growth of ribbon-like colonies. Fig. 31. Distribution of Staurosirella pinnata. TYPE: Not clear, type locality: Montezuma Fluss, Mexico. SEASONALITY: Present throughout the year. DISTRIBUTION: Occurring worldwide in temperate freshwaters. KOREA: This species occurs in various rivers and streams; Nakdong River (Lee 2002; Cho et al. 1993), Lake Imha (Kim et al. 1995; Kim et al. 1997; Kim 1993), Kwang River (Kwangchun) in Ulchin county (Lee et al. 1994), Kumho River (Lee and Chung 1992; Park 1995), Tongwha stream (Tongwhachun) in Daegu (Hong 1990) and Nam stream (Namchun) in Kyungsan city (Jung 2008). SPECIMEN EXAMINED: LJ129 (Kwang river, Ulchin county, Kyungbuk 1992). ECOLOGY: The species is saproxenous (Kobayasi et al. 2006). REMARKS: The original description of the genus Staurosirella Williams and Round states that the genus should include species that form two types of colonies that are adapted to benthic habitats; (frustules attached end to end and ribbon like aggregations), and that have spines and a cingulum composed of open copulae (Morales and Manoylob 2006). One of the most conspicuous unifying features of all the species within Staurosirella is the punch-hole appearance of the striae, which becomes clearer in inner views of the valves (Morales and Manoylob 2006). Nevertheless, there are specimens of Staurosira venter (Ehrenberg) H. Kobayasi, for example, that are very difficult to separated from elliptical forms of S. pinnata. In such cases detailed population studies are required in order to assess phenotypic plasticities and variability (Morales 2001, 2006). This species is distinguished by its coarse striae which are crossed by fine lines like longitudinal slits.

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105 Bryozoa: Gymnolaemata: Cheilostmata: Inovicellata, Malacostega, Flustrina, Ascophora 97 Synedra Kyung Lee and Sook-Kyung Yoon

106 98 Algal Flora of Korea Freshwater Diatoms II

107 Bryozoa: Gymnolaemata: Cheilostmata: Inovicellata, Malacostega, Flustrina, Ascophora 99 List of Taxa Family Diatomaceae Dumortier 1822 Genus Synedra Ehrenberg 1830 Synedra acus Kützing 1844 Synedra delicatissima W. Smith 1853 Synedra delicatissima var. angustissima Grunow in Van Heurck 1881 Synedra goulardi Brébisson ex Cleve and Grunow 1880 Synedra pulchella (Ralfs ex Kützing) Kützing 1844 Synedra pulchella var. lacerata Hustedt 1913 Synedra rumpens var. meneghiniana Grunow in Van Heurck 1880 Synedra socia Wallace 1960 Synedra tabulata var. obtusa (Pantocsek) Hustedt 1959 Synedra tenera W. Smith 1856 Synedra tenuissima (Kützing) Kützing 1844 Synedra ulna (Nitzsch) Ehrenberg 1836 Synedra ulna var. amphirhynchus (Ehrenberg) Grunow 1862 Synedra ulna var. contracta Østrup 1901 Synedra ulna var. danica (Kützing) Van Heurck 1885 Synedra ulna var. obtusa Van Heurck 1885 Synedra ulna var. oxyrhynchus (Kützing) Van Heurck 1885 Synedra ulna var. oxyrhynchus f. mediocontracta (Forti) Hustedt 1959 Synedra ulna var. ramesi (Héribaud) Hustedt 1930 Synedra ulna var. spathulifera (Grunow in Van Heurck) Grunow in Van Heurck 1885

108 100 Introduction The genus Synedra was originally proposed by C.G. Ehrenberg in 1830, but no description of the species characters was provided, so therefore he reproposed it in 1832, while the genus Fragilaria was described by Lynbye in 1819 (VanLangdingham ). The key characters for classification of Synedra species were peculiarities of the apical pore field, striae and the basal siliceous layer, rimoportulae, cingulum, the exterior of the mantle margin, spines, and colony formation, while those of Fragilaria were the valve striation, linking spines, apical pore fields, rimoportulae, girdle, and plastids. Recently, were the generic characters of Fragilaria and Synedra reduced to the overall form, nature of sternum/central area, presence or absence of interlocking spines, arrangement and form of areolae, structure of cribrae, presence of rimoportulae and/or apical pore fields, and the nature of girdle bands (Williams and Round 1986). However, these two genera have caused a controversy between Round (1984) and Lange-Bertalot (1980). These two diatomists recognized the problem of complexity between two genera. However, each of two took a different approach to solve these problems. Lange-Bertalot (1980) emphasized colony formation and habitats differences other than many other characters between the two genera, and suggested a subgeneric division of them based on colonial and other ultrastructural features: rimoportulae, copulae, and striae formation. However, Williams and Round (1987) showed that colony formation is not sufficient in itself to justify a separation of the two genera. There are often differences in many other characters: valve striation, linking spines, apical pore field, rimoportulae, girdle structure, and plastids. These authors also pointed out that lumping genera was a much more serious matter than splitting since it can reduce the information content provided in a taxon like a genus, that in turn provides confusion for ecological interpretations since it e.g. greatly affects genus and species identification (Round 1984). Therefore, the establishment of 5 new genera (Catacombas, Hyalosynedra, Tabularia, Ctenophora, and Neosynedra) from the old genus Synedra (Williams and Round 1986) and another new 5 genera (Staurosira, Staurosirella, Pseudostaurosira, Punctastriata, and Neofragilaria) resulted from the old genus Fragilaria (Williams and Round 1987). Round et al. (1990) also proposed a new classification system for diatoms as a whole: Family Fragilariaceae, Order Fragilariales, Class Fragilariophyceae, Division Bacillariophyta. Otherwise Lange-Bertalot (1980) lumped genus Synedra with the genus Fragilaria with the emphasis of the above mentioned characters and also established new 5 subgenera within the combined genus Synedra/Fragilaria: Fragilaria, Alterasynedra, Ctenophora, Tabularia, and Staurosira (Krammer and Lange-Bertalot 1991). The amended genus Fragilaria belonged to the following classification system according to Simonsen (1979): Genus Fragilaria, Family Fragilariaceae, Suborder Araphidineae, Order Pennales, Class Bacillariophyceae, Phylum Chrysophyta. The objectives of this study are to examine the morphological variations of Synedra species among Korean material, to elucidate their taxonomic identity, and we followed the Simonsen s classification system (1979) because the controversy between two genera (Fragilaria and Synedra) has not been solved still now.

109 Bryozoa: Gymnolaemata: Cheilostmata: Inovicellata, Malacostega, Flustrina, Ascophora 101 Materials and Methods Samples were cleaned by nitric acid or hydrogen peroxide (Hendey 1974), and mounted in Naphrax. Permanent slides were examined with light microscope (Zeiss Axioplan-2, MC-80 Camera System), and identified at 1,000 magnification. The idenfitication of diatoms followed Hustedt (1930, 1959), Krammer and Lange-Bertalot (1991), Patrick and Reimer (1966), Sims (1996). The terminology used was following Anonymous (1975), Choi et al. (1995), Lee et al. (1995), Ross et al. (1979), and Von Stosch (1975).

110

111 103 Taxonomic Notes Genus Synedra Ehrenberg 1830: 40. Dae-ba-neul-dol-mal-sok ( ) Valves occur in planktonic or periphytic forms. Frustules may occur in single or colony form, never though forming long filaments. In valve view, they are very slender, usually linear, needlelike, and lanceolate, while in girdle view, linear and rectangular. Apices of the valve are rounded, capitate, and rostrate. Pseudoraphe is narrow-linear or lanceolate. A central area may or may not present and the shape is squarish or rectangular. Striae are composed of rows of simple round to elongated areolae with uniseriate or biseriate areolation. A jelly pore is present at the end of the valve. Chloroplasts are arranged either as many small plates or as two large plate-like structures. Lectotype: Synedra ulna (Nitzsch) Ehrenberg 1832: 40. SPECIES ca. 211 (43 in Korea). DISTRIBUTION: The species occur as free-living or benthic forms worldwidely in freshwater, brackish water, and sea water. KEY REFERENCE: Lange-Bertalot (1980), Round (1984), Williams and Round (1986), Round et al. (1990), Krammer and Lange-Bertalot (1991). Key to the species of genus Synedra 1. Valve very narrow, needle shaped 2 Valve not narrow, needle shaped 6 2. Central area present 3 Central area absent Length-to-width ratio, greater than 27:1 4 Length-to-width ratio, less than 27:1 S. acus 4. Valve, usually more than 200 μm in length 5 Valve, usually less than 200 μm in length S. tenuissima 5. 2/3 of the valve, 2 μm S. delicatissima v. angustissima 2/3 of the valve, not 2 μm S. delicatissima 6. Valve constricted in central portion 12 Valve not constricted in central portion 7 7. Central area present 8 Central area absent S. ulna v. amphirhynchus 8. Ends of the valve rostrate 9 Ends of the valve not rostrate Linear near the ends of the valve S. ulna Swollen near the ends of the valve S. ulna v. spathulifera 10. Ends of the valve round 11 Ends of the valve capitate S. ulna v. danica

112 104 Algal Flora of Korea Freshwater Diatoms II 11. Ends of the valve clubbed S. ulna v. obtusa Ends of the valve not clubbed Valve, more than 100 μm in length 13 Valve, less than 100 μm in length Central area, oval shaped 15 Central area, not oval shaped Central area, rectangular S. goulardi Central area, squarish S. ulna v. ramesi 15. Ends of the valve long wedge shaped S. ulna v. contracta Ends of the valve short wedge shaped S. ulna v. oxyrhynchus f. mediocontracta 16. Valve swollen at the central area 17 Valve not swollen at the central area S. ulna v. oxyrhynchus 17. Striae distinctly punctate 19 Striae not distinctly punctate Striae, more than 15 in 10 μm S. rumpens v. meneghiniana Striae, less than 15 in 10 μm S. socia 19. Striae, more than 14 in 10 μm S. pulchella v. lacerata Striae, less than 14 in 10 μm S. pulchella 20. Pseudoraphe, more than 3/2 of the valve S. tabulata v. obtusa Pseudoraphe, less than 3/2 of the valve S. tenera 1. Synedra acus Kützing 1844: 68 (Figs. 1, 2). Hustedt 1930: 155. f Kützing 1849: 46. Hustedt 1959: 188. f Patrick and Reimer 1966: 135. f. 1. Chung 1968: 249. f Krammer and Lange-Bertalot 1991: 144. f. 8. Mizuno 1993: 137. f. 12. Sims 1996: 580. f. 1. BASIONYM: Synedra acus Kützing 1844: 68. SYNONYM: Synedra acus Kützing 1849: 46. Synedra tenuis Kützing 1844: 65. Synedra ulna var. acus Mayer f. 30. Synedra acus var. genuina Mayer 1913: 51. f. 42. Syndera goulardi var. acus (Kützing) Frenguelli 1925: 168. f. 18. Fragilaria ulna var. acus (Kützing) Lange-Bertalot 1980: 144. f. 122: Fragilaria acus (Kützing) Lange-Bertalot in Krammer and Lange-Bertalot 2000: 144. f Ulnaria acus (Kützing) M. Aboal Valve lanceolate, with tapering to rounded to slightly capitate apices. Pseudoraphe narrow, becoming a little wider towards middle of valve. Central area distinct, a little longer than wide. Striae parallel, in 10 μm. Length, μm. Width, 4 7 μm. Length-to-width ratio 21:1 27:1. TYPE: Hamberger Moor: Binder. DISTRIBUTION: Spain (Margalef et al. 1976; Aboal and Llimona 1984; Aboal 1988; Aboal 1989;

113 Araphidineae: Diatomaceae: Synedra 105 Aboal et al. 1996), Iran (Jamaloo et al. 2006), China (Cao et al. 2005). New South Wales (Day et al. 1995), Queensland (Day et al. 1995), South Australia (Day et al. 1995), Tasmania (Day et al. 1995), Victoria (Day et al. 1995), Hawaiian Islands (Sherwood 2004). A B C Fig. 1. Synedra acus. A C. valve view ( 1,000, LM). Scales: 20 μm.

114 106 Algal Flora of Korea Freshwater Diatoms II Fig. 2. Distribution of Synedra acus ( : This study, : Previous study). KOREA: Cheongryangri (Skvorzow 1929a), Mid Han River (Chung et al. 1968; Cho 1968a; Chung 1969; Hong 1969; Chung and Kim 1970; Cho 1971b; Cho and Ra 1971; Chung 1972; Cho 1974; Cho et al. 1978; Chung and Lee 1978, 1981, 1984; Cho et al. 1979; Cho 1979; Lee and Chung 1983; Lee 1985; Cho et al. 1989; Lee and Cho 1994; Lee and Yoon 2001), Low Han River (Shim and Choi 1981; Shim et al. 1987), Geum River (Chung and Lee 1979; Park et al. 1984; Chung et al. 1985; Lee 1998; Kim et al. 1998; Kim et al. 2000; Shin and Cho 2001), Mid Nakdong River (Kim and Lee 1996; Lee and Yoon 2001), Low Nakdong River (Chung 1984; Chung et al. 1987; Kim and Lee 1991; Cho 1991; Cho et al. 1993; Seo and Chung 1994), Taehwa River (Lee and Kang 1985; Lee et al. 1997), Lake Songji (Cho et al. 1975; Mizuno et al. 1980), Gangweon-do Hoingseong (Cho 1978a), Lake Imgye (Cho 1978b; Chung and Lee 1978b), Dalcheon, Joryeongcheon (Chung 1979a), Chilgabsan, Gyeryeongsan (Chung and Lee 1980), Okdongcheon (Lee and Cho 1980), Namdaecheon (Chung and Lee 1983), Mt. Chiri (Chung and Lee 1983), Lake Imha (Chung et al. 1986; Kim et al. 1997; Park et al. 1999), Hamanneup (Chung and Noh 1987), Wooponeup, Junam Reservoir (The Administration of Environment 1987), Hongjecheon (Chang and Jin 1980), Hyeongsan River (Chung 1987), Dasajeongsujang (Kim et al. 1990), Namcheon (Choi and Chung 1990), Dongcheon, Isacheon (Noh et al. 1991), Donghwacheon (Hong and Chung 1990), Chojongcheon (Lee 1991), Hoeincheon (Lee 1992), Geumho River (Chung et al. 1993), Shincheon (Chung et al. 1985), Gwangcheon (Lee et al. 1994), Mankyeong River (Choi and Kim 1994), Lake Juam (Lee 1994; Lee and Song 1995), Lake Jinyang (Kim et al. 1995), Seom River (Shin 1996), North Korea (Lee et al. 1997; Cho 2000a, 2000b, 2000c, 2001), Lake Dongbok (Cho et al. 1998), Mid Seomjin River (Lee and Yoon 1999, 2001), Seokjeong Spring (Chung et al. 1999), Suoecheon (Lee and Yoon 2000), Yedang Reservoir (Lee and Yoon 2001). SPECIMEN EXAMINED: (Lake Chuncheon: v.1998), (Geumnamri: v.1998), (Jeonhori: v.1998), (Paldang: v.1998), (Lake Paro: v.1998), (Lake Uiam: v.1998), (Yeoju: v.1998), (Buyeo: vi.1998), (Lake Andong: vi.1998), (Hakpo: vii.1998), (Lake Imha: vi.1998), (Nakjeong: vi.1998), (Yeongho Bridge: vi. 1998), (Pyeongnamri: viii.1998), (Yedang Reservoir: v.1998). ECOLOGY: This species is widely distributed and seems to prefer circumneutral water, which does not have a very low conductivity (Patrick and Reimer 1966). They occur as a littoral form, mostly as tufted colonies on filamentous algae, commonly in freshwater over the entire area (Hustedt 1959). Free-living form usually occurs in water with a ph REMARKS: Synedra acus is characterized by its needle shape, and the terminal area shows rounded or slightly capitate in apices. This species is most closely related to Synedra delicatissima and Synedra radians. Hustedt (1930) has described these species as variety of Synedra acus. However, on careful examination, this species differs in its more lanceolate shape of the valve which gives it a coarser

115 Araphidineae: Diatomaceae: Synedra 107 appearance. Also, the central area is more squarish in being slightly longer than wide, and reaches the margins of the valve. The striae are coarser than in Synedra radians. According to Hustedt (1959), the species is only about 1.5 μm wide near weakly capitate apices, long in μm, wide in 5 6 μm. Transapical striae are fairly delicate, in 10 μm. Central area is usually present and rectangular in outline. Patrick and Reimer (1966) mentioned as length μm, width μm, striae in 10 μm, and length-to-width ratio 20:1 30:1. 2. Synedra delicatissima W. Smith 1853: 72 (Figs. 3, 4). Patrick and Reimer 1966: 136. f. 2. Mizuno 1993: 138. f. 13. SYNONYM: Synedra acus var. delicatissima (W. Simth) Grunow Synedra delicatissima var. mesoleia Grunow f. 6. Synedra acus var. radians Hustedt 1930: 155. f A B C Fig. 3. Synedra delicatissima. A C. valve view ( 1,000). Scales: 20 μm.

116 108 Algal Flora of Korea Freshwater Diatoms II Fragilaria delicatissima (W. Smith) Lange-Bertalot 1980: 129. f. 115: 11 13, 114: 1 8. Ulnaria delicatissima (W. Smith) M. Aboal and P.C. Silva 2004: 361. Valve linear, needle shaped, tapering a little to rounded ends. Pseudoraphe very narrow, but distinct. Central area distinct, longer than wide, not reaching margins of valve. On margins of valve in each side central area very short striae present. Striae throughout valve in parallel, 9 13 in 10 μm. Length, μm. Width, μm. Length-to-width ratio, 30:1 42:1. TYPE: Freshwater. Lough Neagh, D. Dickie, DISTRIBUTION: Victoria (Day et al. 1995). KOREA: Mid Han River (Lee and Yoon 2001), Mid Nakdong River (Lee and Yoon 2001), Mid Imjin River (Lee and Yoon 2002), Togyo Reservoir (Lee et al. 1997), Mid Seomjin River (Lee and Yoon 1999), Suoecheon (Lee and Yoon 2000), Yedang Reservoir (Lee and Yoon 2001). SPECIMEN EXAMINED: (Lake Chuncheon: v.1998), (Geumnamri: v.1998), (Paldang: v.1998), (Lake Uiam: v. Fig. 4. Distribution of Synedra delicatissima ( : This study, : Previous study). 1998), (Lake Andong: vi.1998), (Hakpo: vii.1998), (Hasanri: vii.1998), (Lake Imha: vi.1998), (Nakjeong: vi.1998), (Yeongho Bridge: vi.1998), (Myeongsanri: xi.1998), (Yedang Reservoir: v.1998). ECOLOGY: This species usually occurs free-living or epiphytic forms, found in water with a ph It seems to prefer circumneutral water. REMARKS: Synedra delicatissima is characterized by its needle shape, and its central area is longer than wide, and with a very short striae on each margin. This species is most closely related to Synedra acus, which is more nearly squarish and only slightly longer than wide. Patrick and Reimer (1966) mentioned as length μm, striae in 10 μm, and length-to-width ratio 30:1 50:1. 3. Synedra delicatissima var. angustissima Grunow in Van Heurck 1881: 515 (Figs. 5, 6). Patrick and Reimer 1966: 136. f. 3. Krammer and Lange-Bertalot 1991: 144. f. 21. Mizuno 1993: 138. f. 14. Hustedt 1923: 33. f. 17. Boyer 1926: 202. Hustedt 1930: 155. f Hustedt 1959: 189. f BASIONYM: Synedra delicatissima var. angustissima Grunow in Van Heurck 1881: 515. SYNONYM: Synedra acus var. angustissima (Grunow in Van Heurck) Van Heurck Fragilaria delicatissima var. angustissima (Grunow) Lange-Bertalot 1981: 144. f. 121: 15, 16; f. 114: 21. Ulnaria delicatissima var. angustissima (W. Smith) M. Aboal and P.C. Silva 2004: 361.

117 Araphidineae: Diatomaceae: Synedra 109 A A B B Fig. 5. Synedra delicatissima var. angustissima. A, A, B, B. valve view ( 1,000, LM). Scales: 20 μm.

118 110 Algal Flora of Korea Freshwater Diatoms II Valve very long, narrow by its delicate structure, often sinuous. Pseudoraphe very narrow but distinct. Central area much longer than wide, sometimes clearly rectangular in shape, or with irregular shape. Striae throughout valve parallel, in 10 μm. Length, μm. Width, μm. Length-to-width ratio 42:1 68:1. TYPE: Uncertain, Belgium. DISTRIBUTION: Spain (Aboal and Llimona 1984, Aboal 1988), New South Wales (Day et al. 1995), Victoria (Day et al. 1995), Tasmania (Day et al. 1995). KOREA: Mid Han River (Chung et al. 1968; Chung 1969; Chung and Kay 1969; Hong 1969; Chung and Kim 1970; Cho 1971b; Chung and Lee 1978, 1981, Kim 1996; Kim 1998; Lee and Yoon 2001), Low Han River (Shim et al. 1987), Mid Nakdong River (Kim and Lee 1996; Lee and Yoon 2001), Low Nakdong River (Lee 1973; Chung 1984; Chung et al. 1987; Cho 1991), Geum River (Chung and Lee 1979; Kim et al. 2000), Dalcheon, Joryeongcheon (Chung 1979a), Chilgabsan, Gyeryeongsan (Chung and Lee 1980), Namdaecheon (Chung and Lee 1983), Lake Imgye (Chung and Lee 1978), Lake Yeongcheon (Lee et al. 1992), Geumho River (Chung et al. 1993), Lake Imha (Kim et al. 1995; Kim Fig. 6. Distribution of Synedra delicatissima var. angustissima ( : This study, : Previous study). et al. 1997), Lake Jinyang (Kim et al. 1995), Lake Juam (Lee and Song 1995; Lee and Yoon 2001), Seom River (Shin 1996), Taehwa River (Lee et al. 1997), Mid Seomjin River (Lee and Yoon 1999). SPECIMEN EXAMINED: (Lake Chuncheon: v.1998), (Geumnamri: v.1998), (Paldang: v.1998), (Lake Soyang: v.1998), (Lake Uiam: v.1998), (Lake Andong: vi.1998), (Hakpo: vii.1998), (Hasanri: vii. 1998), (Lake Imha: vi.1998), (Nakjeong: vi.1998), (Ssangrim: vii.1998), (Myeongsanri: xi.1998). ECOLOGY: This species is a typical plankton form, quite commonly found among plankton of inland waters (Hustedt 1959). Usually they are free-living, found in water with a ph REMARKS: Synedra delicatissima var. angustissima is characterized by its needle shape, the central area longer than wide, and has very short striae on each margin. This taxon differs from the nominate variety by its more delicate valve structure and the shape and structure of the central area. Hustedt (1959) mentioned length is over 500 μm, width is 3 μm, and striae is in 10 μm. Patrick and Reimer (1966) mentioned their length to be usually over 200 μm, striae in 10 μm, lengthto-width ratio 55:1. According to Hustedt (1959), Synedra radians Kützing and Synedra delicatissima W. Smith are identical, and represent only one variety of Synedra acus. Patrick and Reimer (1966) have described this species as a variety of Synedra delicatissima. 4. Synedra goulardi Brébisson ex Cleve and Grunow 1880: 107 (Figs. 7, 8). De Toni 1892: 655. Hustedt 1927: 250. Okuno 1952: 39. f. 5. Patrick and Reimer 1966: 154. f. 8.

119 Araphidineae: Diatomaceae: Synedra 111 A B C Fig. 7. Synedra goulardi. A C. valve view ( 1,000, LM). Scales: 20 μm. SYNONYM: Synedra fluviatilis Lemmermann 1910: 317. f. 12, 13. Synedra goulardi var. fluviatilis (Lemmermann) Frenguelli 1941: 313. f. 12, 13. Fragilaria ulna var. goulardi (Brebisson ex Cleve and Grunow) Lange-Bertalot 1980: 745. Valve constricted in middle portion, somewhat swollen toward wedge-shaped ends to become rostrate or subcapitate. Pseudoraphe distinct, narrow. Central area almost as broad as long. Striae faint, present throughout central area; parallel or slightly radiate at ends. Striae radiate in apprearance, in some cases, to be due to whether the inside is examined rather than outside of valve. Striae in 10 μm. Length μm. Width of valve in widest portion about 8 μm. Length-to-width ratios 9:1. TYPE: Uncertain. DISTRIBUTION: Victoria (Day et al. 1995). KOREA: Mid Han River (Lee and Yoon 2001), Mid Nakdong River (Lee and Yoon 2001), Mid Imjin River (Lee and Yoon 2002), Togyo Reservoir (Lee et al. 1997), Fig. 8. Distribution of Synedra goulardi ( : This study, : Previous study).

120 112 Algal Flora of Korea Freshwater Diatoms II Mid Seomjin River (Lee and Yoon 1999), Suoecheon (Lee and Yoon 2000), Yedang Reservoir (Lee and Yoon 2001). SPECIMEN EXAMINED: (Nakjeong: vi.1998), (Yukgyetoseong: v.1998). ECOLOGY: The species often found in warm water (Patrick and Reimer 1966). They usually occur in free-living or epiphytic forms in water with a ph REMARKS: Synedra goulardi is characterized by its constricted at middle portion of valve and an long and widened central area. According to Patrick and Reimer (1966), length is μm, width 8 10 μm, and striae in 10 μm. This taxon is distinguished from Synedra ulna var. ramesi, which resembles in many characteristics by presence of faint striae throughout the central area. 5. Synedra pulchella (Ralfs ex Kützing) Kützing 1844: 68 (Figs. 9, 10). Kützing 1849: 46. Rabenhorst 1853: 56. f. 17. W. Smith 1853: 70. f. 84. Grunow 1862: 392. Hustedt 1930: 160. f Hustedt 1959: 176. f Patrick and Reimer 1966: 146. f. 10, 12. Krammer and Lange-Bertalot 1991: 148. f Mizuno 1993: 138. f. 16. BASIONYM: Exilaria pulchella Ralfs ex Kützing 1844: 68. SYNONYM: Synedra acicularis W. Smith 1853: 70. f. 86., non Kützing Synedra smithii Ralfs in Pritchard 1861: 786. Synedra pulchella var. smithii (Ralfs) Grunow in Van Heurck f. 4. A B C Fig. 9. Synedra pulchella. A C. valve view ( 1,000, LM). Scales: 20 μm.

121 Araphidineae: Diatomaceae: Synedra 113 Synedra pulchella var. abnormis Machiati 1889: 264. Ctenophora pulchella (Ralfs ex Kützing 1844) Schönfeldt 1907: 104. Fragilaria pulchella (Ralfs ex Kützing 1844) Lange-Bertalot 1980: 148. f. 136: 1 7. Ctenophora pulchella (Ralfs ex Kützing 1844) Williams and Round 1986: 330. f Valve in girdle view narrowed toward ends of valve. Valve liear to lanceolate with slightly attenuated rostrate or slightly capitate apices. Central area distinct, slightly swollen, reaching to margins of valve, rectangular to somewhat rounded in shape. Striae distinctly punctuate, parallel to sometimes slightly radiate at ends of valve. Striae parallel, in 10 μm. Length μm. Width 6 8 μm. Length-to-width ratio 13:1 17:1. TYPE: Conferven in England. Rabenhorst 662, Van Heurck 298, 300. Tempère and Peragallo 34, 123, 188, 280, 333. DISTRIBUTION: Spain (Aboal 1988), New South Wales (Day et al. 1995), Queensland (Day et al. 1995), Tasmania (Day et al. 1995), Victoria (Day et al. 1995). Fig. 10. Distribution of Synedra pulchella ( : This study, : Previous study). KOREA: Lake Seo (Skvortzow 1929b), Mid Han River (Chung et al. 1968; Chung and Lee 1978; Cho et al. 1979; Kim 1996), Low Han River (Chung et al. 1965; Sim and Choi 1981; Sim et al. 1987), Namdaecheon (Chung and Lee 1983), Gyeongju (Chung and Watanabe 1984), Geum River (Park et al. 1984; Shin and Cho 2001), Mount Woelchul (Lee 1989), Namcheon (Choi and Chung 1990), Yangsancheon (Chung et al. 1992), Geumho River (Chung et al. 1993), Low Nakdong River (Cho et al. 1993), Gwangcheon (Lee et al. 1994), Mankyung River (Choi and Kim 1994), North Korea (Cho 2000a, 2000b, 2000c, 2001), Mid Nakdong River (Lee and Yoon 2001). SPECIMEN EXAMINED: (Jeongyang: vii.1998), (Hakpo: vii.1998). ECOLOGY: This species are usually found in fresh water of high mineral content, or in slightly brackish water (Hustedt 1959; Patrick and Reimer 1966). They usually occur in free-living forms, found in water with a ph REMARKS: Synedra pulchella is distinguished by its shape: its distinctly punctuated striae, and its swollen central area. According to Hustedt (1959), length are μm, width are 5 8 μm and striae are in 10 μm. 6. Synedra pulchella var. lacerata Hustedt 1913 (Figs. 11, 12). Hustedt 1959: 178. f. 688c. Patrick and Reimer 1966: 147. f. 11.

122 114 Algal Flora of Korea Freshwater Diatoms II A B C Fig. 11. Synedra pulchella var. lacerata. A C. valve view ( 1,000, LM). Scales: 20 μm. SYNONYM: Synedra pulchella f. lacerata (Hustedt in Schmidt) Hustedt 1957: 235. Valve in girdle view narrow toward ends of valve. Valve linear to lanceolate with slightly attenuated rostrate or capitate apices. Central area distinct, swollen, reaching to margins of valve, rectangular to somewhat rounded in shape. Striae distinctly punctuate, parallel to sometimes slightly radiate at ends of valve. Striae parallel, in 10 μm. Length μm. Width 5 8 μm. Length-to-width ratio 13:1 18:1. TYPE: Lake Victoria. DISTRIBUTION: U.S. Florida, Nebraska (Patrick and Reimer 1966). KOREA: Gwangcheon (Lee et al. 1994), Seomjin River (Lee and Yoon 1999), Nakdong River (Lee and Yoon 2001), Imjin River (Lee and Yoon 2002). SPECIMEN EXAMINED: (Lake Imha: vi.1998), (Yeongho Bridge: vi.1998), (Jeongyang: vii.1998), (Yeukgyaetoseong: v.1998), (Jeongam: vii.1998). ECOLOGY: This variety is usually found in freshwater and brackish water with high conductivity (Patrick and Reimer 1966). They usually occur in free-living or epiphytic forms, found in water

123 Araphidineae: Diatomaceae: Synedra 115 with a ph REMARKS: Synedra pulchella var. lacerate is linear-lanceolate in girdle view, gradually decreasing in width from the middle toward the ends. This variety is distinguished by a width, somewhat irregularly formed axial area (Hustedt 1959; Patrick and Reimer 1966). Patrick and Reimer (1966) states striae are in 10 μm, length are about 100 μm, and width are 6 7 μm. 7. Synedra rumpens var. meneghiniana Grunow in Van Heurck 1880 (Figs. 13, 14). Hustedt 1930: 156. f Hustedt 1959: 193. Patrick and Reimer 1966: 145. f. 3. Krammer and Lange- Bertalot 1991: 124. f BASIONYM: Synedra rumpens var. meneghiniana Grunow in Van Heurck SYNONYM: Synedra rumpens var. meneghiniana f. curta A. Mayer 1916: 22. f. 31. Valve linear to linear-lanceolate, slightly narrowed from middle toward ends. Apices round to rostrate. Pseudoraphe very narrow, sometimes indistinct. Central area longer than wide, somewhat swollen in each side. Striae parallel, slightly radiated from middle toward ends. Striae coarse, in 10 μm. Length Fig. 12. Distribution of Synedra pulchella var. lacerata ( : This study, : Previous study). A B C Fig. 13. Synedra rumpens var. meneghiniana. A C. valve view ( 1,000). Scales: 20 μm.

124 116 Algal Flora of Korea Freshwater Diatoms II μm. Width 4 6 μm. Length-to-width ratios 5:1 11:1. TYPE: Battaglia DISTRIBUTION: Queensland (Day et al. 1995), Hawaiian Island (Sherwood 2004). KOREA: Mid Han River (Chung 1972), Gyeongju (Chung and Watanabe 1984), Donghwacheon (Hong and Chung 1990), Baekcheon (Chung and Kim 1991), Mangyeong River (Choi and Kim 1994), Mid Geum River (Chung 1995), Mid Seomjin River (Lee and Yoon 1999, 2001), Mid Nakdong River (Lee and Yoon 2001). SPECIMEN EXAMINED: (Yangsuri: iv.2000), (Ssangrim: vii.1998), (Yeongho Bridge: vi.1998), (Oeenjeong: i.1999), (Jangsan: iv.1999), (Pyeongnamri: viii.1998). ECOLOGY: This variety occurs in free-living or epiphytic forms, found in water with a ph of It seems to prefer fresh water of low mineral content (Patrick and Reimer 1966). REMARKS: Synedra rumpens var. meneghiniana is distinguished from the other varieties of the species by its valve shape showing lanceolate and coarser striae. According to the descriptions of Patrick and Reimer (1966) and Hustedt (1959), striae are in 10 μm. However, in this study, they are usually 11 Fig. 14. Distribution of Synedra rumpens var. meneghiniana ( : This study, : Previous study). in 10 μm. In morphological appearance it is very similar to Synedra socia, but differs in its finer striae, shorter length, wider width, and somewhat swollen central area on each side. This species is closely related to Synedra rumpens var. fragilarioides (Fukushima et al. 1991). 8. Synedra socia Wallace 1960: 1 (Figs. 15, 16). Patrick and Reimer 1966: 145. f. 4 6; Krammer and Lange-Bertalot 1991: 126. SYNONYM: Fragilaria socia (Wallace) Lange-Bertalot 1980: 126. Valve linear-lanceolate with capitate to rostrate apices. Pseudoraphe very narrow, but distinct. Valve slightly constricted, then swollen in middle portion in region of central area. Central area well developed, somewhat swollen in middle portion, reaching to margins of valve on both sides, sometimes short striae along margin on one side. Striae fine, parallel, in 10 μm. Length usually μm. Width μm. Length-to-width ratios 7:1 11:1. TYPE: U.S. Georgia, Screven County, Savannah River. A G.C. 4036a, Wallace DISTRIBUTION: Hawaiian Islands (Sherwood 2004), Southeastern States (Patrick and Reimer 1966).

125 Araphidineae: Diatomaceae: Synedra 117 A B C D Fig. 15. Synedra socia. A D. valve view ( 1,000). Scales: 20 μm. KOREA: Mid Han River (Kim 1996), Gyeungju (Chung and Watanabe 1984), Sincheon (Chung et al. 1985), Hyeongsan River (Chung 1987), Namcheon (Choi and Chung 1990), Yangsancheon (Kim et al. 1992; Chung et al. 1992), Donghwacheon (Hong and Chung 1990), Lake Yeongcheon (Lee et al. 1992), Baekcheon (Chung and Kim 1991). SPECIMEN EXAMINED: (Guhwanggyo: i.1999), (Hwangjukgyo: i.1999), (Jangsan: iv.1999). ECOLOGY: This species occurs in free-living or epiphytic forms, found in water with a ph of This species prefers the circumneutral water of low conductivity (Patrick and Reimer 1966). REMARKS: Synedra socia is characterized by its lanceolate valve shape, slightly swollen central area, and finer striae. According to the description of Patrick and Reimer (1966), length is μm, width 3 4 μm, and striae 17 in 10 μm. This taxon is typically much smaller in length-to-width ratio and more lanceolate than Synedra rumpens. The shape of the valve is similar to that of Fragilaria vaucheriae, except for the structure of Fig. 16. Distribution of Synedra socia ( : central area and much coarse striae. This species is This study, : Previous study). very similar to Synedra rumpens var. familiaris (Kützing) Hustedt. However, in the variety familiaris the striae are finer and the length-to-width ratio is greater (Patrick and Reimer 1966). 9. Synedra tabulata var. obtusa (Pantocsek) Hustedt 1959: 219 (Figs. 17, 18). Hustedt 1930: 160. f Hustedt 1959: 203. f. 710h. Patrick and Reimer 1966: 141. f. 17. Chung

126 118 Algal Flora of Korea Freshwater Diatoms II A B Fig. 17. Synedra tabulata var. obtusa. A, B. valve view ( 1,000). Scales: 20 μm. 1968: 257. f Mizuno 1993: 139. f. 28. BASIONYM: Synedra fasciculata var. obtusa Pantocsek 1889: 64. f SYNONYM: Synedra affinis var. obtusa Arnott ex Van Heurck f. 12. Synedra tabulata var. pantocsekii Cleve-Euler 1953: 70. f. 392r. Synedra affinis var. obtusa sensu Halden 1929: 362. Synedra affinis var. obtusa Hustedt f Fragilaria tabulata var. obtusa (Pantocsek) Lange-Bertalot Catacombas obtusa (Pantocsek) Snoeijs in Snoeijs 1991: 157. f. 1, 4, 8, 9, Valve linear to linear-lanceolate, widen slightly toward middle portion, somewhat narrow towards rounded to slightly subcapitate apices. Pseudoraphe distinct, 80% in width of valve. Central area absent. Jelly pore distinct near end of one terminal striae. Striae parallel, very short, in 10 μm. Length μm. Width μm. Length-to-width ratios 18:1 21:1.

127 Araphidineae: Diatomaceae: Synedra 119 TYPE: Øresund (Sweden). DISTRIBUTION: Northern Baltic Sea, Forsmark (Snoeijs 1991). KOREA: Mid Han River (Lee and Cho 1994), Gwangcheon (Lee et al. 1994), Low Nakdong River (Seo and Chung 1994), Lake Juam (Lee 1994; Lee and Song 1995), Taehwa River (Lee et al. 1997), North Korea (Cho 2000b, 2000c, 2001). SPECIMEN EXAMINED: (Lake Imha: vi.1998), (Nakjeong: vi.1998), (Hwangjookgyo: i.1999), (Jangsan: iv. 1999), (Hwaseokjeong: vii.2000), (Geumsan: ix. 2001). ECOLOGY: This variety occurs in free-living or epiphytic forms, found in water with a ph of It seems to prefer circumneutral water. This taxon is found throughout the year as an epiphyte on filamentous macroalgae (especially, Cladophora glomerata) (Snoeijs et al. 1991). REMARKS: Synedra tabulata var. obtusa is characterized by its very short striae, wide pseudoraphe, and round and subcapitate valves. According to the description of Hustedt (1959), length is over 250 μm, and width 7 8 μm, while to Patrick and Reimer (1966), length is μm, width 2 7 μm, and striae in 10 μm. This taxon is closely related to Synedra fasciculata, but differs mainly in number of finer striae, in in 10 μm Fig. 18. Distribution of Synedra tabulata var. obtusa ( : This study, : Previous study). (Patrick and Reimer 1966). This variety is very similar to Synedra fasciculata, but differs in its finer striae, and shape of the pseudoraphe. Williams and Round (1986) placed Synedra tabulata in the genus Tabularia as Tabularia fasciculata, and mentioned that Synedra tabulata var. obtusa might also belong to the genus Catacombas. Snoeijs et al. (1991) suggest transfer Synedra tabulata var. obtusa to the genus Catacombas. 10. Synedra tenera W. Smith 1856: 98 (Figs. 19, 20). Hustedt 1930: 158. f Hustedt 1959: 196. f Patrick and Reimer 1966: 137. f. 5. Krammer and Lange-Bertalot 1991: 129. f. 1 5, f Sims 1996: 582. f. 9. SYNONYM: Synedra tenera var. genuina Cleve-Euler 1953: 57. f. 375a c. Fragilaria tenera (W. Smith) Lange-Bertalot 1980: 129. f. 115: 1 5, 6, 7; F. 114: Valve very narrow, linear, tapering to rounded slightly inflated ends. Apices not clearly differentiated from rest of valve, sometimes slightly rostrate-capitate. Pseudoraphe indistinct. Central area generally absent. Striae fine, parallel, usually in 10 μm. Length μm. Width μm. Length-to-width ratio 26:1 43:1.

128 120 Algal Flora of Korea Freshwater Diatoms II A B B Fig. 19. Synedra tenera. A, B, B. valve view ( 1,000, LM). Scales: 20 μm. TYPE: Uncertain. DISTRIBUTION: Spain (Varela 1982). New South Wales (Day et al. 1995), Queensland (Day et al. 1995), Victoria (Day et al. 1995).

129 Araphidineae: Diatomaceae: Synedra 121 KOREA: Mid Han River (Chung and Kim 1970; Chung and Lee 1984), Low Han River (Shim and Choi 1981), Lake Imgye (Chung and Lee 1978), Seom River (Shin 1996), Lake Yedang (Lee and Yoon 2001). SPECIMEN EXAMINED: (Yukgyetoseong: v.1998), (Lake Yedang: v.1998). ECOLOGY: This species usually occurs in free-living forms, found in water with a ph They are found in the littoral standing waters (Hustedt 1959), and seem to prefer slow-moving water of low conductivity (Patrick and Reimer 1966). REMARKS: Synedra tenera is characterized by its needle shape, lack of the central area, and very fine striae. Valve is narrow-lanceolate, gradually narrowed from middle toward the ends. Patrick and Reimer (1966) mentioned the length to be usually μm, width μm, and striae in 10 μm. According to Hustedt (1959), they exhibit the length μm, width μm, and striae in 10 μm. This species is closely related to Synedra delicatissima, but differs mainly in finer striae. Fig. 20. Distribution of Synedra tenera ( : This study, : Previous study). 11. Synedra tenuissima (Kützing) Kützing 1844: 64 (Figs. 21, 22). Hustedt 1930: 155. f Hustedt 1959: 189. f. 693 b. Patrick and Reimer 1966: 137. f. 4. BASIONYM: Frustulia tenuissima Kützing 1833: 64. f. 6. SYNONYM: Synedra radians Kützing 1844: 64. f. VII, 1 4. Synedra radians var. tenuissima (Kützing) Rabenhorst Synedra acus var. tenuissima (Kützing) Schumann Valve narrow-linear, narrowing towards slightly rostrate rounded apices. Pseudoraphe very narrow-linear. Central area rectangular in shape, a little longer than wide, or absent. Striae throughout valve parallel, fine, in 10 μm. Length μm. Width μm. Length-to-width ratio 27:1 40:1. TYPE: An Cladophora fracta bei Tennstädt. DISTRIBUTION: Romania (Caraus 2002). KOREA: Mid Geum River (Chung 1995), Seokjeong Spring (Chung et al. 1999). SPECIMEN EXAMINED: (Hwang River: vii.1998), (Lake Imha: vi.1998), (Nakjeong: vi.1998), (Nam River: vii.1998), (Ssangrim: vii.1998), (Yeongho Bridge: vi.1998), (Yukgyetoseong: v.1998), (Yedang Reservoir: v.1998).

130 122 Algal Flora of Korea Freshwater Diatoms II A B B C Fig. 21. Synedra tenuissima. A, B, B, C. valve view ( 1,000, LM). Scales: 20 μm. ECOLOGY: This species is often found in plankton in slightly alkaline water, often with fairly high conductivity (Patrick and Reimer 1966). They usually occur in free-living form, found in water with a ph It seems to prefer circumneutral water. REMARKS: Synedra tenuissima is characterized by its linear needle shape. It differs from Synedra acus by its finer striae and more linear narrow shape. It is similar to Synedra delicatissima in shape, but differs in fineness of the striae. According to the description of Patrick and Reimer (1966), the length is usually μm, width μm, and striae in 10 μm, while according to Hustedt (1959), the length is μm, and width is μm. This taxon is previously identified

131 Araphidineae: Diatomaceae: Synedra 123 as Synedra radians Kützing. According to Patrick and Reimer (1966), this spieces is intermediate in shape between Synedra acus and Synedra delicatissima. It is also closely related to Exilaria tenuissima Brébisson. 12. Synedra ulna (Nitzsch) Ehrenberg 1838: 211 (Figs. 23, 24). Ehrenberg 1841: 389. f. 23. Kützing 1844: 66. f. 28. Hustedt 1930: 151. f Hustedt 1959: 181. f Patrick and Reimer 1966: 148. f. 1, 2. Krammer and Lange-Bertalot 1991: 143. pls Mizuno 1993: 137. f Sims 1996: 582. f. 10. BASIONYM: Bacillaria ulna Nitzsch 1817: 99. SYNONYM: Frustulia splendens Kützing 1833: 553. f. 23. Synedra lanceolata Kützing 1844: 66. f. 31. Fig. 22. Distribution of Synedra tenuissima ( : This study, : Previous study). Synedra ulna var. undulata Grunow Synedra ulna var. notata Grunow in Van Heurck Synedra ulna var. subaequalis Grunow in Van Heurck f. 13. Synedra ulna var. splendens (Kützing) Van Heurck Synedra ulna var. curta A. Mayer 1912: 4. Synedra ulna var. longirostris (Grunow in Van Heurck) A. Cleve-Euler 1948: 10. Synedra ulna var. notata f. crassa (Østrup) A. Cleve-Euler 1953: 63. f Synedra ulna var. balatoneis f. pantocsekii A. Cleve-Euler 1953: 63. f Synedra ulna (Nitzsch) Ehrenberg Fragilaria ulna (Nitzsch) Lange-Bertalot 1980: 143. f. 122: 1 8. Ulnaria ulna (Nitzsch) Compère in Jahn et al. 2001: 100. Valve linear, very gradually attenuated to rostrate or sometimes somewhat rostrate-wedge-shaped ends. Pseudoraphe narrow, but distinct. Central area not much longer than wide, often almost squarish, sometimes with very short striae on margins. Striae parallel, usually 8 10 in 10 μm. Length usually μm. Width μm. Length-to-width ratio 13:1 28:1. TYPE: Grundschlamme des Wittenberger Stadtgrabens, Germany. Kützing 1. Rabenhorst 6, 67. Van Heurck 15, 107, 284, 286, 312. Tempère and Peragallo 13, 37, 113, 198, 227, 280, 290, 308, 390, 470, 485, 596, 599, 686, 789, 910. DISTRIBUTION: Germany (Ettl and Gärtner 1995; Bahulikar and Kroth 2007), Iceland (Ettl and Gärtner 1995), Spain (Alvarez Cobelas 1982; Aboal 1988, 1989; Aboal et al. 1996), Iran (Jamaloo et al. 2006). New South Wales (Day et al. 1995), Queensland (Day et al. 1995), South Australia (Day et al. 1995), Tasmania (Day et al. 1995), Victoria (Day et al. 1995), Western Australia (Day et al. 1995),

132 124 Algal Flora of Korea Freshwater Diatoms II A B C Fig. 23. Synedra ulna. A C. valve view ( 1,000, LM). Scales: 20 μm. Hawaiian Islands (Sherwood 2004). KOREA: Cheonryangri (Skvortzow 1929a), Boseong River, Seomjin River, Milyang River (Ueda and Okada 1935), Hamnam Anbyeon (Okuno 1948), Han River (Choi and Shin 1976; Cho 1967, 1968a, 1968b; Chung et al. 1965; Chung et al. 1968; Chung 1969; Chung and Kay 1969; Hong 1969; Chung and Kim 1970; Cho 1971b, 1974, 1979a; Cho and Ra 1971; Cho et al. 1978b, 1979; Cho et al. 1989; Chung and Lee 1978, 1981, 1984; Lee 1985; Chung 1972; Lee et al. 1994; Lee and Cho 1994; Kim 1996; Lee and Yoon 2001), Low Han River (Chung et al. 1965; Kim 1972; Shim and Choi 1981; Shim et al. 1987; Yoo and Lim 1990; Lee and Chang 1997), Sangdongcheon (Cho 1971a), Lake Sanjeong

133 Araphidineae: Diatomaceae: Synedra 125 (Kang 1966), Lake Yeongrang (Cho and Park 1969; Cho et al. 1975), Lake Songji (Cho et al. 1975), Mid Nakdong River (Kim and Lee 1996; Lee and Yoon 2001), Low Nakdong River (Chung 1970, 1984; Chung et al. 1987; Kim and Lee 1991; Cho 1991; Cho et al. 1993; Seo and Chung 1994), Lake Imgye (Chung and Lee 1978), Geum River (Cho et al. 1978a; Chung and Lee 1979; Shim and Yang 1982; Park et al. 1984; Chung et al. 1985; Kim et al. 1998; Kim et al. 2000; Lee and Yoon 2001), Dalcheon, Joryeongcheon (Chung 1979a), Juwangsan (Chung 1979b; Chung et al. 1985), Balwangsan (Chung 1979b), Chilgabsan, Gyeoryeongsan (Chung and Lee 1980), Okdongcheon (Lee and Cho 1980), Mushimcheon (Lee 1980), Gyebangsan (Chung and Lee 1982a), Odaesan (Chung et al. 1982b), Namdaecheon (Chung and Lee 1983; Chung et al. 1987), Bogildo (Chung 1983), Chirisan (Chung and Lee 1983), Gyeongju (Chung and Watanabe 1984), Shincheon (Chung et al. 1985), Taehwa River (Lee and Kang 1985; Lee et al. 1997), Lake Imha (Chung et al. 1986; Kim et al. 1997), Hamanneup (The Administraion of Environment 1984; Chung and Noh 1987), Wooponeup, Lake Junam (The Administraion of Environment 1987), Hongjecheon (Jang and Jin 1980), Hyeongsan River Fig. 24. Distribution of Synedra ulna ( : This study, : Previous study). (Chung 1987), Youngsan River (Kim and Choi 1988), Wolchulsan (Lee 1989), Muju Seolcheonmyeon (Lee et al. 1990; Lee 1990), Namcheon (Choi and Chung 1990), Jejudo (Lee and Chang 1991), Dongcheon, Isacheon (Noh et al. 1991), Yangsancheon (Kim et al. 1992; Chung et al. 1992), Donghwacheon (Hong and Chung 1990), Lake Yeongcheon (Lee et al. 1992), Jojeongcheon (Lee 1991), Hwoiincheon (Lee 1992), Mangyeong River (Chung et al. 1992; Choi and Kim 1994; Chung et al. 1996), Geumho River (Chung et al. 1993), Baekcheon (Chung and Kim 1991), Gwangcheon (Lee et al. 1994), Lake Juam (Choi et al. 1994; Lee 1994; Lee and Song 1995), Lake Imha (Kim et al. 1995; Park et al. 1999), Lake Jinyang (Kim et al. 1995), Seom River (Shin 1996), Boryeong (Boo et al. 1997), Unjangsan (Kim et al. 1998), Mid Seomjin River (Lee and Yoon 1999, 2001), Seokjeong Spring (Chung et al. 1999), Suoecheon (Lee and Yoon 2000), North Korea (Cho 2000a, 2000b, 2000c, 2001), Yedang Reservoir (Lee and Yoon 2001). SPECIMEN EXAMINED: (Geumnamri: v.1998), (Ipo: v.1998), (Paldang: v.1998), (Yeoju: v.1998), (Buyoe: vi.1998), (Gongju: vi.1998), (Lake Andong: vi.1998), (Hakpo: vii.1998), (Lake Imha: vi.1998), (Nakjeong: vi.1998), (Yeongho Bridge: vi.1998), (Lake Damyang: xi.1998), (Lake Jangseong: xi.1998), (Myeongsanri: xi.1998), (Yeongsan Bridge: xi.1998), (Pyeongnamri: viii.1998), (Yukgyetoseong: v. 1998), (Yedang Reservoir: v.1998). ECOLOGY: This species is widely distributed in fresh water, in free-living or epiphytic forms, found in water with a ph REMARKS: There are many varieties described in Synedra ulna, but are distinguished from them by the shape of apices and central area of the species. Valve is linear, very gradually attenuated to the rostrate or sometimes somewhat rostrate-wedge-shaped ends. Central area is not much longer

134 126 Algal Flora of Korea Freshwater Diatoms II than wide, often almost squarish. Striae are parallel. Hustedt (1959) states sometimes 8 12 striae in 10 μm, and Patrick and Reimer (1966) mentioned usually 9 11 in 10 μm. Boyer (1927) designated Synedra ulna as the type species of the genus. However, this is not technically correct as in Ehrenberg s 1830 paper, when Synedra ulna was listed as Navicula ulna (Bory) Ehrenberg. This would leave Synedra balthica Ehrenberg [=Synedra gaillonii (Bory) Ehrenberg] as the type of the genus. Williams (1986) showed that S. gaillonii would not be included within a genus typified by S. ulna, and Synedra would be better conserved for species having the type of structure in Synedra ulna. 13. Synedra ulna var. amphirhynchus (Ehrenberg) Grunow 1862: 397 (Figs. 25, 26). Brun 1880: 126. f. 25. Van Heurck : 151. f. 5. Hustedt 1930: 154. f Hustedt 1959: 186. f Cleve-Euler 1953: 62. f Patrick and Reimer 1966: 149. f. 6, 7. Mizuno 1993: 137. f. 10. BASIONYM: Synedra amphirhynchus Ehrenberg 1843: 425. f. 25. SYNONYM: Synedra vitrea Bory ex Kützing 1844: 66. f. 17. Synedra ulna var. vitrea (Bory ex Kützing) Van Heurck 1885: 151. Navicula affinis var. amphirhynchus (Ehrenberg) Grunow. Ulnaria ulna var. amphirhynchus (Ehrenberg) M. Aboal 2003: 113. Valve linear, suddenly constricted to become attenuated toward rostrate or sometimes slightly capitate ends. Pseudoraphe very narrow, but distinct. Central area absent. Striae parallel, about 10 in 10 μm. Length usually μm. Width 8 10 μm. Length to width ratio 22:1 28:1. TYPE: Uncertain. DISTRIBUTION: Romania (Caraus 2002). Hawaiian Islands (Sherwood 2004), Spain (Aboal et al. 2003). KOREA: Lake Seo (Skvortzow 1929b), Han River (Chung et al. 1968; Chung 1969; Chung and Kay 1969; Hong 1969; Chung and Kim 1970, 1983; Lee 1985; Chung 1972; Sim and Choi 1981), Lake Imgye (Chung and Lee 1978), Chilgabsan, Gyeryeongsan (Chung and Lee 1980), Gyebangsan (Chung and Lee 1982a), Namdaecheon (Chung and Lee 1983), Hamanneup (Chung and Noh 1987), Nakdong River (Lee and Yoon 2001), Mid Imjin River (Lee and Yoon 2002). SPECIMEN EXAMINED: (Hakpo: vii.1998), (Nakjeong: vi.1998), (Yeongsan Bridge: xi.1998), (Yukgyetoseong: v.1998). Fig. 25. Distribution of Synedra ulna var. amphirhynchus ( : This study, : Previous study).

135 Araphidineae: Diatomaceae: Synedra 127 ECOLOGY: This variety is found in circumneutral, usually mesotrophic to eutrophic fresh water (Patrick and Reimer 1966), in free-living or epiphytic forms, found in water with a ph A B Fig. 26. Synedra ulna var. amphirhynchus. A, B. valve view ( 1,000, LM). Scales: 20 μm.

136 128 Algal Flora of Korea Freshwater Diatoms II REMARKS: Synedra ulna var. amphirhynchus is characterized by its size, the structure of valve ends, and the lack of central area. Valve is linear, capitate or subcapitate in ends. According to Patrick and Reimer (1966), length are μm, width are 4 7 μm, and striae are in 10 μm. 14. Synedra ulna var. contracta Østrup 1901: 281 (Figs. 27, 28). Hustedt 1930: 152. f Hustedt 1959: 199. f Hustedt 1959: 185. f Okuno 1952: 39. f. 8. Patrick and Reimer 1966: 150. f. 3. SYNONYM: Synedra ulna var. oxyrhynchus f. contracta Hustedt 1930: 152. f Valve linear to linear-capitate with concave margins, appearing to be somewhat constricted in middle portion of valve. Valve wedge-shaped in end with attenuate-rostrated apices. Pseudoraphe distinct, abruptly widening into central area. Central area almost as wide as long, reaching to margins of valve. Striae parallel throughout most of valve, slightly radiate in ends. Striae in 10 A B C Fig. 27. Synedra ulna var. contracta. A C. valve view ( 1,000, LM). Scales: 20 μm.

137 Araphidineae: Diatomaceae: Synedra 129 μm. Length μm. Width 8 9 μm. Length-towidth ratio 12:1 15:1. TYPE: Found in a gathering from the Faeroes labeled Green algae from a swamp on Nolsö. DISTRIBUTION: Romania (Caraus 2002), New South Wales (Day et al. 1995), Victoria (Day et al. 1995). KOREA: Mid Han River (Chung et al. 1968; Chung 1969; Chun and Kay 1969; Chung and Kim 1970; Chung 1972; Lee and Chung 1983; Chung and Lee 1984; Lee 1985), Low Han River (Yoo and Lim 1990), Boseong River (Ueda and Okada 1935), Okdongcheon (Lee and Cho 1980), Gyebangsan (Chung and Lee 1982a), Odaesan (Chung et al. 1982b), Namdaecheon (Chung et al. 1987), Hyeongsan River (Chung 1987), Dasajeongsujang (Kim et al. 1990), Namcheon (Choi and Chung 1990), Mujoo Seolcheonmyeon (Lee 1990), Donghwacheon (Hong and Chung 1990), Mid Nakdong River (Lee and Yoon 2001), Mid Imjin River (Lee and Yoon 2002). SPECIMEN EXAMINED: (Nakjeong: vi.1998), (Jeongam: vii.1998), (Ssangrim: vii.1998), (Yukgyetoseong: v.1998). ECOLOGY: This variety occurs in circumneutral Fig. 28. Distribution of Synedra ulna var. contracta ( : This study, : Previous study). fresh water, in spite of small amounts of salt (Patrick and Reimer 1966). They are free-living or epiphytic forms, found in water with a ph REMARKS: Synedra ulna var. contracta is distinguished by the shape of valve and the central area. This taxon is most closely related to Synedra ulna var. ramesi, from which it differs by the size, greater length-to-width ratio, and the shape of central area. This variety as well as Synedra ulna var. ramesi is closely related to Synedra goulardi (Patrick and Reimer 1966). Patrick and Reimer (1966) states that length μm, width 7 8 μm, and striae about 10 in 10 μm. 15. Synedra ulna var. danica (Kützing) Van Heurck 1885: 151 (Figs. 29, 30). Van Heurck 1896: 311. f Hustedt 1930: 154. f Hustedt 1959: 187. f Cleve-Euler 1953: 62. f Patrick and Reimer 1966: 151. f. 10. Krammer and Lange-Bertalot 1991: 144. f. 9. Sims 1996: 582. f. 12. BASIONYM: Synedra danica Kützing 1844: 66. f. 13. SYNONYM: Syndea splendens var. danica Grunow 1862: 396. Synedra chasei Thomas ex Wolle f. 1. Synedra longissima var. acicularis Meister 1912: 75. f. 2. Synedra danica f. typica Cleve-Euler 1953: 62. f. 382.

138 130 Algal Flora of Korea Freshwater Diatoms II Valve linear to lanceolate, swollen, somewhat capitate at ends. Pseudoraphe narrow. Central area transverse, usually not reaching to margins of valve. Striae parallel to sometimes slightly radiate A B C Fig. 29. Synedra ulna var. danica. A C. valve view ( 1,000, LM). Scales: 20 μm.

139 Araphidineae: Diatomaceae: Synedra 131 at ends of valve, 9 11 in 10 μm. Length μm. Width 5 8 μm. Length-to-width ratio 26:1 51:1. TYPE: In süssem Wasser der jütischen Halbinsel: Hb. Binder. DISTRIBUTION: Spain (Aboal and Llimona 1984), New South Wales (Day et al. 1995), Queensland (Day et al. 1995), Victoria (Day et al. 1995), Hawaiian Islands (Sherwood 2004). KOREA: Mid Han River (Chung 1972), Kyungju (Chung and Watanabe 1984), Hamanneup (Chung and Noh 1987), Mid Geum River (Chung 1995), Mid Seomjin River (Lee and Yoon 1999, 2001), Mid Nakdong River (Lee and Yoon 2001), Mid Imjin River (Lee and Yoon 2002). SPECIMEN EXAMINED: (Lake Andong: vi.1998), (Lake Imha: vi.1998), (Yeongho Bridge: vi.1998), (Nakjeong: vi.1998), (Ssangrim: vii.1998), (Jeongyang: vii.1998), (Hakpo: vii.1998), (Myeongsanri: xi.1998), (Yoengsan (Bridge: xi.1998), (Yukgyetoseong: v.1998), (Pyeongnamri: viii.1998). ECOLOGY: This variety is especially common and distributed as a plankton form in eutrophic lakes (Hustedt 1959) and also, often found in the plankton of circumneutral ph fresh water, being in sensitive to Fig. 30. Distribution of Synedra ulna var. danica ( : This study, : Previous study). small amounts of salts (Patrick and Reimer 1966) in free-living or epiphytic forms, with a ph REMARKS: Synedra ulna var. danica is distinguished by the shape of valve and central area. Valve is very narrow, barely over 5 μm width in middle, almost needle-shaped, and gradually narrowing towards the weakly capitate ends, but not remarkably constricted near the ends (Hustedt f. 691A, f). Patrick and Reimer (1966) mentioned length μm, width 5 8 μm, and striae 9 10 in 10 μm, while Hustedt (1959) mentioned length to be μm, width 5 8 μm, and striae in 10 μm. 16. Synedra ulna var. obtusa (W. Smith) Van Heurck 1885: 151 (Figs. 31, 32). Patrick and Reimer (1966) p. 152, pl. 7. f. 9. BASIONYM: Synedra obtusa W. Smith 1853: 71. f. 92. SYNONYM: Frustulia aequalis Kützing 1833: 546. f. 30., non. Synedra aequalis Kützing f. 14. Valve very long, linear with round to slightly swollen apices. Pseudoraphe narrow, but distinct. Central area variable in size; squarish, rectangular, or absent. Jelly pore well developed. Striae

140 132 Algal Flora of Korea Freshwater Diatoms II A B C Fig. 31. Synedra ulna var. obtusa. A C. valve view ( 1,000, LM). Scales: 20 μm.

141 Araphidineae: Diatomaceae: Synedra 133 parallel, 9 10 in 10 μm. Length μm. Width 5 6 μm. Length-to-width ratios 34:1 44:1. TYPE: Uncertain. DISTRIBUTION: U.S. East Central States; Montana (Patrick and Reimer 1966). KOREA: Mid Nakdong River (Kim 1999). SPECIMEN EXAMINED: (Hakpo: vii.1998), (Lake Imha : vi.1998), (Nakjeong: vi.1998), (Yeongho Bridge: vi. 1998), (Lake Gwangpo: x.1986). ECOLOGY: According to Patrick and Reimer (1966), this variety seems to prefer cool water. However, in this study, their water temperature varies between C. They are free-living or epiphytic forms, found in water with a ph REMARKS: Synedra ulna var. obtusa is distinguished by its broad apices, which are scarcely differentiated from main part of cells. Our varieties have a greater length-to-width ratio than the original variety, while its central area is less well developed. According to the description of Patrick and Reimer (1966), lengths are μm, widths are 6 8 μm, and striae are 8 11 in 10 μm. Smith (1853) states this taxon is same as the specimens of Synedra aequalis Kützing (1849) sent to him by Brébisson. However, they are not the same Fig. 32. Distribution of Synedra ulna var. obtusa ( : This study, : Previous study). as Synedra aequalis illustrated by Kützing (1844) although Kützing considered them as the same as the species listed in Species Algarum. Although Smith s name may have been superfluous when published, it is the oldest one at the variety rank (Patrick and Reimer 1966). 17. Synedra ulna var. oxyrhynchus (Kützing) Van Heurck 1885: 151 (Figs. 33, 34). Hustedt 1930: 152. f Hustedt 1959: 184. f. 691B. q., Krammer and Lange-Bertalot 1991: 144. f. 10. Mizuno 1993: 137. f. 5, 6. Sims 1996: 582. f. 13. BASIONYM: Synedra oxyrhynchus Kützing 1844: 66. f. 8, f. 2, IX, X, XI. SYNONYM: Synedra oxyrhynchus var. amphicephala Grunow 1862: 399. f. 14. Synedra oxyrhynchus var. genuina Grunow 1862: 399. Synedra oxyrhynchus var. undulata Grunow 1862: 400. f. 13. Valve is linear to linear-lanceolate or constricted in middle portion, usually somewhat widened toward ends. Valve ends rostrate, somewhat swollen toward the ends, wedge-shaped in apices. Pseudoraphe distinct. Central area usually squarish or rectangular. Striae 8 10 in 10 μm. Length

142 134 Algal Flora of Korea Freshwater Diatoms II A B C D Fig. 33. Synedra ulna var. oxyrhynchus. A D. valve view ( 1,000, LM). Scales: 20 μm μm. Width 7 10 μm in central area. Length-to-width ratios 9:1 14:1. TYPE: Rabenhorst Tempère and Peragallo 470, 489, 638. DISTRIBUTION: New South Wales (Day et al. 1995), Queensland (Day et al. 1995), Victoria (Day et al. 1995), Spain (Aboal et al. 2003). KOREA: Upper Han River (Choi and Shin 1976), Mid Han River (Chung et al. 1968; Chung 1969; Chung and Kay 1969; Hong 1969; Chung and Kim 1970; Chung 1972), Low Han River (Chung et al. 1965; Shim and Choi 1981; Chung and Lee 1978, 1981, 1984; Lee 1985), Geum River (Chung and Lee 1979; Lee 1998; Kim et al. 2000; Shin and Cho 2001), Dalcheon, Joryeongcheon (Chung 1979a), Chilgabsan, Gyeryongsan (Chung and Lee 1980), Ulleungdo, Dokdo (Chung 1981), Gyebangsan (Chung and Lee 1982a), Odaesan (Chung and Lee 1982b), Taehwa River (Lee and Gang 1985; Lee et al. 1997), Namdaecheon (Chung and Lee 1983; Chung et al. 1987), Okdongcheon (Lee and Cho 1980), Lake Imha (Chung 1986), Hamanneup (The Administration of Environment 1984; Chung and Noh

143 Araphidineae: Diatomaceae: Synedra ), Wolchulsan (Lee 1989), Dongcheon, Isacheon (Noh et al. 1991), Muju Seolcheonmyeon (Lee 1990), Yangsancheon (Kim et al. 1992), Jojongcheon (Lee 1991), Hoeincheon (Lee 1992), Mid Seomjin River (Lee and Yoon 1999, 2001), Mid Nakdong River (Lee and Yoon 2001), Mid Imjin River (Lee and Yoon 2002). SPECIMEN EXAMINED: (Lake Andong: vi.1998), (Hakpo: vii.1998), (Hwang River: vii.1998), (Lake Imha: vi.1998), (Nakjeong: vi.1998), (Ssangrim: vii.1998), (Myeongsanri: xi.1998), (Youngsan Bridge: xi.1998), (Goohwanggyo: i.1999), (Jangsan: iv.1999), (Pyeongnamri: viii.1998), (Hwaseokjeong: vii.2000), (Jangnamgyo: x.2000), (Yukkeytoseong: v.1998), (Hantan River: xi.1998). ECOLOGY: This variety usually occurs in free-living or epiphytic forms, found in water with ph This taxon seems to prefer circumneutral water. REMARKS: Synedra ulna var. oxyrhynchus is distinguished by the shape of long rostrate and wedgeshape ends, and appears to be somewhat contricted or wideen in middle portion of the valve. This variety is similar to Synedra ulna var. contracta and Synedra ulna var. oxyrhynchus f. mediocontracta. Although Synedra ulna var. oxyrhynchus resembles in many taxonomic Fig. 34. Distribution of Synedra ulna var. oxyrhynchus ( : This study, : Previous study). characters, it is distinguished from them by the shape of apices, shape of central area, and length-towidth ratio. Hustedt (1930) described the striae are in 10 μm. However, in this study, 8 10 striae were observed in 10 μm. 18. Synedra ulna var. oxyrhynchus f. mediocontracta (Forti) Hustedt 1959: 199 (Figs. 35, 36). Hustedt 1959: 199. f. 691B. r., Patrick and Reimer 1966: 152. f. 4. BASIONYM: Synedra oxyrhynchus var. mediocontracta Forti 1910: SYNONYM: Synedra oxyrhynchus var. contracta Hustedt 1911: 273. f. 25. Synedra ulna f. mediocontracta Komarenco Valve linear to linear-lanceolate, somewhat constricted in middle portion, swollen towards wedgeshaped ends to become rostrate or sucapitate. Pseudoraphe distinct, somewhat wideened near central area. Central area oval to rounded square-shaped with short striae. Striae in 10 μm. Length μm. Width 8 10 μm in central area. Length-to-width ratios 6:1 13:1. TYPE: nel sedimento più leggero del saggio raccoloto sulle colline dei Soddo.

144 136 Algal Flora of Korea Freshwater Diatoms II A B C D Fig. 35. Synedra oxyrhynchus f. mediocontracta. A D. valve view ( 1,000). Scales: 20 μm. DISTRIBUTION: U.S. Tennessee (Patrick and Reimer 1966). KOREA: Mid Imjin River (Yoon 2002). SPECIMEN EXAMINED: (Jangnamgyo: x.2000), (Yukgyetoseong: v.1998). ECOLOGY: They occur in free-living or epiphytic forms, found in water with a ph This taxon seems to prefer circumneutral water of low conductivity (Patrick and Reimer, 1966). In this study, they are usually found in water with a ph 7.4. REMARKS: Synedra oxyrhynchus f. mediocontracta is characterized by short rostrate and wedge-shaped ends, somewhat constricted middle portion, with oval shaped central areas showing short striae. This form is similar to Synedra ulna var. contracta and Synedra ulna var. oxyrhynchus, but it differs from Synedra ulna var. oxyrhynchus by abrupt constriction of the valve in central area. According to the description of Patrick and Reimer (1966), length is μm, width 6 8 μm, and striae 11 in 10 μm. Fig. 36. Distribution of Synedra ulna var. oxyrhynchus f. mediocontracta ( : This study, : Previous study).

145 Araphidineae: Diatomaceae: Synedra Synedra ulna var. ramesi (Héribaud) Hustedt 1930: 152 (Figs. 37, 38). Patrick and Reimer 1966: 153. f. 9. Chung 1968: 260. f Mizuno 1993: 137. f. 8. BASIONYM: Synera ramesi Héribaud 1903: 80. f. 28. Valve linear to lanceolate with margins concave in middle portion of valve, slightly swollen near narrow attenuated rostrate apices. Pseudoraphe narrow, distinct. Central area usually longer than wide in square shaped. Striae parallel, in 10 μm. Length μm. Width 7 8 μm. Length-to-width ratios 6:1 7:1. TYPE: Uncertain. DISTRIBUTION: U.S. New England States, Middle Atlantic States, Southeastern States, Gulf Coast States, Montana, New Mexico, California (Patrick and Reimer 1966). KOREA: Mid Nakdong River (Kim 1999), Mid Imjin River (Kim 1999), Mid Seomjin River (Kim 1999). SPECIMEN EXAMINED: (Nakjeong: vi.1998), (Ssangrim: vii.1998), (Ganjeongyo: x.1998), (Jewolri: viii.1998), (Yukgyetoseong: v.1998). ECOLOGY: This variety occurs in free-living or epiphytic forms, found in water with a ph of It seems to prefer circumneural water. Fig. 37. Distribution of Synedra ulna var. ramesi ( : This study, : Previous study). A B C Fig. 38. Synedra ulna var. ramesi. A C. valve view ( 1,000, LM). Scales: 20 μm.

146 138 Algal Flora of Korea Freshwater Diatoms II REMARKS: Synedra ulna var. ramesi is characterized by its rostrate or wedge shaped apices, valve with margins concave in middle portion, and central area of widened square shape. This taxon is distinguished from Synedra ulna var. contracta by the bigger, difference of length-to-width ratio, and the different shape of the central area (Patrick and Reimer, 1966). Hustedt (1959: 199) has synonymized this taxon with Synedra ulna var. contracta Østrup. Also, this taxon is most closely related to Synedra goulardi. However, it differs in its larger shape of the valve, finer striae, and more thick-lanceolated apices. 20. Synedra ulna var. spathulifera (Grunow in Van Heurck) Grunow in Van Heurck 1885: 151 (Figs. 39, 40). Hustedt 1930: 152. f Hustedt 1959: 186. f. h, k p. Patrick and Reimer 1966: 153. f. 8. Mizuno 1993: 137. f. 9. BASIONYM: Synedra spathulifera Grunow in Van Heurck 1881: 151. f. 4. SYNONYM: Synedra balatonis Pantocsek 1902: 76. f Synedra rostata Pantocsek 1902: 76. f Synedra joursacensis Héribaud 1903: 23. f. 8. Valve linear, somewhat swollen near ends of valve. Apices wedge-shaped, rounded to be ends of valve, as spatulate in shape. Pseudoraphe narrow. Central area variable in size, squarish to rectangular, somewhat swollen. Striae parallel throughout valve or sometimes slightly radiate at apices, 8 10 in 10 μm. Length μm. Width 5 7 μm. Length-to-width ratio 20:1 37:1. TYPE: Deure près d Anvers. DISTRIBUTION: Romania (Caraus 2002), New South Wales (Day et al. 1995), Queensland (Day et al. 1995). KOREA: Yoon (2002). SPECIMEN EXAMINED: (Jangsan: iv.1999). ECOLOGY: According to Patrick and Reimer (1966), this variety seems to prefer cool water. However in this study, they are usually observed at more than 20 C in freshwater with a ph about 7.0, and seem to prefer circumneurtral water. REMARKS: Synedra ulna var. spathulifera is characterized by the linear round cells ending rounded spatulate to wedge-shaped. Central area is slightly swollen, sometime occurring with faint striae. According to the description of Patrick and Reimer (1966), length is μm, width μm, and striae in 10 μm. Fig. 39. Distribution of Synedra ulna var. spathulifera ( : This study, : Previous study).

147 Araphidineae: Diatomaceae: Synedra 139 A B C Fig. 40. Synedra ulna var. spathulifera. A C. valve view ( 1,000, LM). Scales: 20 μm.