Algal Flora of Korea

Transcription

1 Algal Flora of Korea Volume 3, Number 9 Chrysophyta: Bacillariophyceae: Pennales: Raphidineae: Naviculaceae: Biremis, Caloneis I, Pinnularia I Freshwater Diatoms VII 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 9 Chrysophyta: Bacillariophyceae: Pennales: Raphidineae: Naviculaceae: Biremis, Caloneis I, Pinnularia I Freshwater Diatoms VII 2012 National Institute of Biological Resources Ministry of Environment

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5 Algal Flora of Korea Volume 3, Number 9 Chrysophyta: Bacillariophyceae: Pennales: Raphidineae: Naviculaceae: Biremis, Caloneis I, Pinnularia I Freshwater Diatoms VII Gyeongje Joh Inje University

6 Copyright c 2012 by the National Institute of Biological Resources Published by the National Institute of Biological Resources Environmental Research Complex, Hwangyeong-ro 42, 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 : Sang Pal Lee Project Staff : Youn-Bong Ku, Ga Youn Cho, Jin Hee Kim Published on December 24, 2012 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 The adoption of the Nagoya protocol on access to genetic resources and the fair and equitable sharing of benefits arising from their utilization in 2010 led to the realization of international standardization in the fulfillment of the biological sovereignty and the exploration and preservation of indigenous biological organisms will play a critical role in enhancing the national development and the international competitiveness. Most developed countries had already organized the information of species inhabiting in their territories to claim their severeignty over those biological resources, but in this respect Korea was outpaced by these countries. In order to effectively secure, use and manage the indigenous biological organisms it is imperative to systematically understand them at the national level and to organize them to publish in the Flora and Fauna of Korea. Recognizing the importance of its securement and management in taking the initiative in bioindustry in future, National Institute of Biological Resources of the Ministry of Environment has been publishing the Flora and Fauna of Korea for systematic and efficient management of biological resources of our own. For the last 5 years, professional research groups consisting of relevant professors and the like conducted systematic surveys and organizations for a variety of and wide range of taxa. As a result, 65 issues of Flora and Fauna of Korea, both in Korean and in English, covering 7,709 species and two issues of world monograph covering 216 species were published and 25 issues of Flora and Fauna of Korea, both in Korean and in English, covering 1,313 species are published this year. These efforts serve not only to identify indigenous species living in Korea and to provide the scientific evidences and certifications to claim the sovereign rights over indigenous biological resources in Korea, but also provide the opportunity to prepare the framework for the biotechnological industrialization of biological resources. Finally I would like to express sincere appreciation for Professor Gyeongje Joh of Inje University who did not spare his efforts to publish Biological Magazine. Sang Pal Lee President NIBR

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9 1 Contents List of Taxa 3 Introduction 5 Materials and Methods 7 Taxonomic Notes 9 1. Biremis ambigua (Cleve) D.G. Mann 9 2. Biremis lucens (Hustedt) Sabbe, Witkowski et Vyverman Caloneis africana (Giffen) Stidolph Caloneis amphisbaena (Bory) Cleve Pinnularia acrosphaeria W. Smith Pinnularia angulosa Krammer Pinnularia borealis Ehrenberg Pinnularia borealis var. scalaris (Ehrenberg) Rabenhorst Pinnularia borealis var. sublinearis Krammer Pinnularia brauniana (Grunow) Mills Pinnularia crucifera Cleve-Euler Pinnularia divergens W. Smith Pinnularia divergentissima (Grunow) Cleve Pinnularia eifelana (Krammer) Krammer Pinnularia episcopalis Cleve Pinnularia gibba Ehrenberg Pinnularia gigas Ehrenberg Pinnularia globiceps var. linearis Krammer Pinnularia graciloides Hustedt Pinnularia graciloides var. triundulata (Fontell) Krammer Pinnularia grunowii Krammer Pinnularia karelica Cleve Pinnularia lata (Brébisson) Rabenhorst Pinnularia macilenta Ehrenberg Pinnularia microstauron (Ehrenberg) Cleve Pinnularia microstauron var. nonfasciata Krammer Pinnularia neomajor var. inflata Krammer Pinnularia nobilis var. regularis Krammer Pinnularia obscura Krasske Pinnularia peracuminata Krammer Pinnularia pluvianiformis Krammer Pinnularia pseudogibba Krammer Pinnularia pulchra Østrup Pinnularia quadratarea (A. Schmidt) Cleve Pinnularia rabenhorstii var. franconica Krammer Pinnularia rhombarea Krammer 79

10 2 Algal Flora of Korea Freshwater Diatoms VII 37. Pinnularia schoenfelderi Krammer Pinnularia similiformis var. koreana Metzeltin et Krammer Pinnularia sinistra Krammer Pinnularia stomatophora var. salina Krammer Pinnularia subbrevistriata Krammer Pinnularia subcapitata var. subrostrata Krammer Pinnularia subgibba var. sublinearis Krammer Pinnularia subgibba var. undulata Krammer Pinnularia subgigas Krammer Pinnularia subinterrupta Krammer et Schroeter Pinnularia transversa (A. Schmidt) Mayer Pinnularia viridiformis Krammer Pinnularia viridiformis var. minor Krammer Pinnularia viridis (Nitzsch) Ehrenberg 105 Literature Cited 108 Index to Korean Names 116 Index to Korean Names as Pronounced 117 Index to Scientific Names 118

11 3 List of Taxa Class Bacillariophyceae Order Pennales Suborder Raphidineae Family Naviculaceae Kützing 1944 Genus Biremis D.G. Mann et Cox in Round, Crawford and D.G. Mann 1990 Biremis ambigua (Cleve) D.G. Mann 1990 Biremis lucens (Hustedt) Sabbe, Witkowski et Vyverman 1995 Genus Caloneis Cleve 1894 Caloneis aerophila Bock 1963 Caloneis africana (Giffen) Stidolph 1995 Caloneis amphisbaena (Bory) Cleve 1894 Genus Pinnularia Ehrenberg 1843 Pinnularia acrosphaeria W. Smith 1853 Pinnularia angulosa Krammer 2000 Pinnularia borealis Ehrenberg 1843 Pinnularia borealis var. scalaris (Ehrenberg) Rabenhorst 1864 Pinnularia borealis var. sublinearis Krammer 2000 Pinnularia brauniana (Grunow) Mills 1934 Pinnularia crucifera Cleve-Euler 1934 Pinnularia divergens W. Smith 1853 Pinnularia divergentissima (Grunow) Cleve 1895 Pinnularia eifelana (Krammer) Krammer 2000 Pinnularia episcopalis Cleve 1891 Pinnularia gibba Ehrenberg 1843 Pinnularia gigas Ehrenberg 1843 Pinnularia globiceps var. linearis Krammer 2000 Pinnularia graciloides Hustedt 1937 Pinnularia graciloides var. triundulata (Fontell) Krammer 2000 Pinnularia grunowii Krammer 2000 Pinnularia karelica Cleve 1891 Pinnularia lata (Brébisson) Rabenhorst 1853 Pinnularia macilenta Ehrenberg 1843 Pinnularia microstauron (Ehrenberg) Cleve 1891 Pinnularia microstauron var. nonfasciata Krammer 2000 Pinnularia neomajor var. inflata Krammer 2000 Pinnularia nobilis var. regularis Krammer 2000 Pinnularia obscura Krasske 1932 Pinnularia peracuminata Krammer 2000 Pinnularia pluvianiformis Krammer 2000 Pinnularia pseudogibba Krammer 1992 Pinnularia pulchra Østrup 1897 Pinnularia quadratarea (A. Schmidt) Cleve 1895 Pinnularia rabenhorstii var. franconica Krammer 2000

12 4 Algal Flora of Korea Freshwater Diatoms VII Pinnularia rhombarea Krammer 1998 Pinnularia schoenfelderi Krammer 1992 Pinnularia similiformis var. koreana Metzeltin et Krammer 2000 Pinnularia sinistra Krammer 1992 Pinnularia stomatophora var. salina Krammer 2000 Pinnularia subbrevistriata Krammer 2000 Pinnularia subcapitata var. subrostrata Krammer 1992 Pinnularia subgibba var. sublinearis Krammer 1992 Pinnularia subgibba var. undulata Krammer 1992 Pinnularia subgigas Krammer 2000 Pinnularia subinterrupta Krammer et Schroeter 1992 Pinnularia transversa (A. Schmidt) Mayer 1939 Pinnularia viridiformis Krammer 1992 Pinnularia viridiformis var. minor Krammer 2000 Pinnularia viridis (Nitzsch) Ehrenberg 1843

13 5 Introduction 1. Taxonomy of Pinnularia diatoms Diatoms of the genus Pinnularia and related genera are characterized by transapically chambered striae on valves among the bilateral naviculoid diatoms, while other naviculoids have punctate striae. The outer surface of the strial chamber is ornamented with multiple rows of small pores and the inner surface with elongate openings. Additionally, the outer fissures of raphes in species of the genus Pinnularia are straight, curved and undulated (double twisted), and raphe forms are very complex and variable. The internal openings of the striae reflect the appeareance of longitudinal lines or bands across the striae under the light microscopy. In his synopsis of naviculoid diatoms published in 1895, P.T. Cleve had already divided Pinnularia into eight sections, Parallelistriatae, Capitatae, Divergentes, Distantes, Tabellarieae, Brevistriatae, Majores, Complexae. Hustedt (1930) and Patrick and Reimer (1966) followed the Cleve s system and splitted the genus into eight and seven sections, respectively, and the morphology of both the axial areas and central area are especially important in their systems. The forms of the central area change during the life cycle of diatoms and larger forms of a certain species often have more extensively developed area than small forms (Krammer 1992). The presence of the fascia in central area and the forms of the raphe is considered by Krammer (2000) to be particularly important in the classification of Pinnularia. A wide range of morphological variability is seen within the species of Pinnularia, and many infraspecific taxa will be separated from their nominate taxa when the fine-grained species concepts proposed by Mann (1999) are adopted (Kulikovskiy et al. 2010a). Revisions of the Pinnularia taxa were intensively done by K. Krammer in 1992, but dealing only European taxa and later by Krammer (2000, 2002) with the taxa collected from the temperate regions. In Krammer s revisions, many infraspecies or morphotypes in the old classification are converted to species and the complexes containing many forms are splitted into higher taxonomic unit of species or varieties. Caloneis has more delicate and denser striae than the diatoms of Pinnularia and the ranges overlap between two groups (Mann 2001), but has different striae orientation and dissimilar curvatures of the terminal raphe fissures. There are moderate differences in the ultrastructure of the stria chambers between two groups. However, the smaller forms of Caloneis show indistinct longitudinal lines like small Pinnularia. Although the two genera are considered as independent taxa in history, combining the genera, Pinnularia and Caloneis, was proposed by many authors such as Patrick and Reimer (1966), Krammer and Lange-Bertalot (1986), Round et al. (1990) and Mann (2001). Despite the evidence from molecular phylogeny, the two genera are separately described in this monograph. The traditional separation of Caloneis from Pinnularia is challenged by many authors. Recently, a multi-gene approach was adopted to reconstruct a molecular phylogeny or diversification for the geuns Pinnularia and Caloneis (Souffreau et al. 2010). These results produce three major clades and some subclades, but not universally, delimited by valve morphology. This does not yet support the view that either Pinnularia or Caloneis, or both is monophyletic and the sibling genus is to be combined with Pinnularia. The Pinnularia-Caloneis complex is considered to have diverged between the Upper Cretaceous and the early Eocene, implying a ghost range of at least 10 million year in the fossil record. In addition, Lange-Bertalot and Metzeltin (1996) proposed a new genus, Chamaepinnularia, pre-

14 6 Algal Flora of Korea Freshwater Diatoms VII Table 1. The number of Pinnularia series taxa (species and variety) reported in freshwater, brackish or coastal water of Korea Family Freshwater Genus Coastal water Naviculaceae* Sum 94 *Naviculaceae as defined by Simonsen (1979). Caloneis (12) Pinnularia (69) Biremis (1) 81 Caloneis (4) 13 Pinnularia (8) viously belonging to Navicula and Pinnularia, and approximately 44 species have been described to date (Guiry and Guiry 2012). Pulchella Krammer and Hygropetra Krammer were separated from the genus Pinnularia and Alveovallum was newly created by Krammer (2000), and Biremis D.G. Mann et E.J. Cox was also removed from Pinnularia in Round et al. (1990). 2. Flora of Pinnularia diatoms The genus Pinnularia and two other related genera, Caloneis and Biremis, are widely distributed worldwide. They have been reported with approximately 2700 taxa of Pinnularia series, of which 500 are accepted taxonomically as valid taxa (Guiry and Guiry 2012). Although the diatoms of the genus Pinnularia are widespread throughout a variety of water bodies, they never reaches a high abundances, but are patchy in their distributions in freshwaters. These diatoms generally prefer oligotrophic and low electrolyte freshwater, but a few are marine forms. In Korea, Pinnularia species were first reported from the study of ponds near Seoul and Suwon by Skvortzow (1929a, 1929b). Chung (1968) reviewed 39 taxa of Pinnularia in his monograph of freshwater diatoms in Korea. Chung (1993) published the second edition of freshwater diatoms in Korea and included nine species in the treatment. Noh (1991) collected Pinnularia diatoms from 39 sites of various types in Korean freshwaters and reported the 44 taxa in detailed descriptions in her monograph. The flora of Pinnularia was thus considered to be 41 species and 52 infraspecies in the diatom inventory summarized comprehensively by KNCCN (1996) and Lee (1997), and Caloneis to be 9 and 13 taxa, respectively. Summing up the Pinnularia taxa in Korea, the number of species is currently 77.

15 7 Materials and Methods Diatoms of the genus Pinnularia are found mainly in mountain peatlands or bogs, which are oligotrophic, dystrophic or acidic environments. The sampling locations were situated at five regions: so called Yeongnam Alps, mountains in the southeastern area of Korea, Mountain Jiri in Sancheong, Mountain Daeam in Inje and Mountain Odae in Pyeongchang, Mountain Halla in Jeju Island. Wetlands form peatlands or bogs around the summits of mountains from 600 to 1100 m elevations. Peatlands near the Mountain Jeongjok are on a small-area from 0.1 to 0.9 ha, whilst the wetlands in the other regions are relatively large to 78 ha (NWC 2010). Wetlands are distributed around the mountain summit and the hydrology of these areas depends mainly on sudden or irregular rainfall. The mountain wetlands are usually covered with moor grass and surrounded by oak and pine trees at their margins. These grass plants are the major contributors to the peat deposit and formation, and the mosses Sphagnum are not every considerable in the coverage of the wetland. Small pools or water holes are scattered in wetlands. In the pools, the slurrys or suspension in the water holes are collected to sample the benthic diatoms in the bogs. The top sediments or deposited peats represent a temporally and spatially integrated sample of recent years. Sampling locations from a variety of wetlands are summarized as follow, 1. The first Mujechineup, the second Mujechineup and the third Mujechineup in Mountain Jeongjok, Ulsan 2. The Daeseongdwitneup, the Daeseongkunneup and a small water pool near the Daeseongkunneup, near Mountain Jeongjok, Yangsan 3. The Daeseongneup A near Mountain Jeongjok, Yangsan 4. The Anjeokneup and the Anjeokneup C near Mountain Jeongjok, Yangsan 5. The Hwaeomneup in Mountain Wonhyo, Yangsan 6. The Sinbulsanneup A, B and C in Mountain Sinbul, Yangsan 7. The Danjoneup in Mountain Yeongchuk, Yangsan 8. The Sandeulneup in Mountain Jaeyak, Miryang 9. The Wandeungjaeneup and the Oegogyeneup in Mountain Jiri, and the Duncheolneup in Mountain Duncheol, Sancheong 10. The Sumeunmulbaengdwineup and the 1100 wetland in Mountain Halla, Jeju Island 11. The Yongneup in Mountain Daeam, Inje 12. The Sohwangbyeongsanneup in Mountain Odae, Pyeongchang 13. An unnamed wetland in Daegwanryeong Pasture, Pyeongchang 14. The Eoseungsaengak and the Saraoreum, crater lakes in Halla Mountain, Jeju Island 15. The Jangdoneup, Jangdo Island, Sinan Preparations of diatom samples followed the standard procedures of APHA (1995). The materials are oxidized with HNO 3 and potassium dichromate (K 2 Cr 2 O 7 ) on a sand bath, a hotplate filled with coarse sands. The oxidized materials are washed twelve times with distilled water to remove chemicals. Clean diatom frustules are mounted on slide glasses with Pleurax resin to make permanent slides. Diatoms are observed under oil immersion using an Olympus microscope (model, Provis AX2) equipped with differential interference contrast (DIC) optics and a Zeiss microscope (Axioplan model). All photographs of diatoms are made with 2000 magnifications.

16 8 Algal Flora of Korea Freshwater Diatoms VII For the identification of the genus Pinnularia, many taxonomic works were consulted, particularly Hustedt (1930), Patrick and Reimer (1966), Krammer and Lange-Bertalot (1986), Noh (1991) and Krammer (2000). Krammer (2000), however, was exclusively referred to for the classification of Pinnularia diatoms in the present monograph. The key index for the classification of Pinnularia diatoms is adopted from 21 subgroups of Krammer (2000) and their keys. To provide synonyms and basionyms of diatom species, we refer to Catalogue of Diatom Names of academic website, (CAS 2012), which has been held by California Academy of Sciences (CAS) for world life s diversity. Informations about diatom flora over the world came from the global database, AlgaeBase ( (Guiry and Guiry 2012). The description by Noh (1991) and check-lists authorized by Lee (1995), KNCCN (1996) and Lee (1997) are source documents to sum up the Pinnularia flora reported or described in Korea. The terminology for diatom frustules of Ross et al. (1979) is adopted to describe the species.

17 9 Taxonomic Notes Genus Biremis D.G. Mann et E.J. Cox in Round, Crawford and D.G. Mann 1990: 664. Cells solitary, usually lying in the girdle view. Valves linear in outline, bilaterally symmetrical, occassionaly dorsiventral or curved. The forms of dorsivental valves like those of the genus Amphora. Girdle with many porous bands. The ends of the valve moderately rounded, not protracted and not differentiated from the body. Raphe central to eccentric, straight or biarcuate in the wide axial area. The central fissures of the raphe simple or expanded, and the terminal fissures curved smoothly to turn towards the margins. Axial area wide and central area not developed. Striae complex, visible as transversely broad lines in the light microscopy, and, but in the electron microscopy, elongated chambers with internally porous cribrums and externally two circular foramina near the valve margin and axial area. Two plastids distributed on both sides of median transapical plane. Lectotype: Biremis ambigua (Cleve) D.G. Mann in Round, Crawford and D.G. Mann 1990 (= Pinnularia ambigua Cleve 1895) SPECIES: There are 24 species and infraspecific names in the database at present, of which 14 have been accepted as taxonomically valid (Guiry and Guiry 2012). DISTRIBUTION: These diatoms are common and frequent in the coastal and brackish sediment off the sea as attached forms to sands and epipelic forms on the muds. Although Biremis was originally described from marine habitats, these diatoms are also encountered in freshwater, oligotrophic lakes (Spaulding and Edlund 2009). KEY REFERENCE: Round et al. (1990). REMARKS: Diatoms belonging to the genus Biremis are similar to the genus Pulchella Krammer, but differ in the curvature of the valve mantle, many intercalary bands and other some characteristics (Krammer 2000). The genus diatoms have the characteristics of three groups, Amphora, Pinnularia and Mastogloia. 1. Biremis ambigua (Cleve) D.G. Mann in Round, Crawford and D.G. Mann 1990: 664 (Figs. 1, 2). Hendey 1964: 233. pl. 34. f. 5. Witkowski et al. 2000: 158. pl f. 2. BASIONYM: Pinnularia ambigua Cleve 1895: 94. Valves linear in outline, the margins of the valve convex or concave in the middle, the ends of the valve broadly or acutely rounded, or asymmetrical and sublunate in outline. Raphe slightly eccen-

18 10 Algal Flora of Korea Freshwater Diatoms VII B C D E 10 μm A Fig. 1. Biremis ambigua. A D. view of valves; E. view of a girdle ( 2000, LM). tric, the central fissures of the raphe straight and close each other, terminal fissures curved smoothly in one direction. Axial area wide and central area not developed. Striae 8 9 in 10 μm, shortly distributed on the margins of the valve in symmetrical forms, longer on the dorsal side and reduced to a line of puncta on the ventral side in asymmetrical forms. Valves μm in length and 8 10 μm in breadth. SYNTYPE LOCALITIES: Oldenburg in Germany; Cape Wankarema (Vankarem) in Russia; Lysekil, Bohüslan in Sweden. ECOLOGY AND DISTRIBUTION: This species is frequently widespread on the coastal and brackish sediments from the high Arctic to the tropical region. It was recorded with high density as planktons in a productive water body of the Balayan Bay, Philippines Fig. 2. Distribution of Biremis ambigua.

19 Pennales: Naviculaceae: Biremis 11 (Evangelista 2009). This species was reported as Pinnularia ambigua Cleve from the western coast of Korea (Choi and Shim 1986), the sandflats of the Nakdong River Estuary (Cho 1989) and the Incheon Dock (Lee and Byun 1991). REMARKS: It seems that Biremis ambigua is the most frequent and common among the Biremis taxa in the coastal environments. 2. Biremis lucens (Hustedt) Sabbe, Witkowski et Vyverman 1995: 38 (Figs. 3, 4). Sabbe et al. 1995: 380. f. 4. Witkowski et al. 2000: 159. pl f. 9. BASIONYM: Navicula lucens Hustedt 1942: 69. Valves linear-elliptical in outline, the margins of the valve convex narrowing towards the rounded ends, sometimes slightly constricted in the middle. Raphe straight, the central fissures of the raphe straight and expanded, the terminal fissures of the raphe curved in one direction. Axial area widely lanceolate narrowing towards the ends, 3/4 of the valve breadth, central area not developed. Striae in 10 μm, short along the margins of the valve, nearly parallel. Valves 7 25 μm in length and μm in breadth. TYPE LOCALITY: Miang Besar, Borneo. ECOLOGY AND DISTRIBUTION: This species is com- Fig. 3. Distribution of Biremis lucens. A B C D 10 μm E F G Fig. 4. Biremis lucens. A G. view of valves ( 2000, LM).

20 12 Algal Flora of Korea Freshwater Diatoms VII mon and widespread in brackish and marine sediments. It usually forms small colonies firmly attached to sand grains (Sabbe et al. 1995). Biremis lucens is found as benthic diatoms in the estuaries of Westerschelde in the southwestern Netherlands, Hopkins in Australia, the Gulf of Gdansk in Poland and in a coastal lake of Papua New Guinea (Sabbe et al. 1995). In Korea, this species is first found from the sandflats in the estuary of the Nakdong River. REMARKS: The characteristic features of this species is characterized by the structure of the striae. Genus Caloneis Cleve 1894: 46. O-i-dol-mal-sok ( ) Valves naviculoid with two isopolar ends, linear to broadly lanceolate in outline and sometimes swollen or undulate in the middle parts. The terminal ends of the valve cuneate, rounded, rostrate to subcapitate. Raphe straight and the terminal ends of the raphe curved to one side. Axial area narrowly linear, often broad, central area developed in various forms, sometimes widening up to the margins of the valve. Central nodules, lunate or irregular thickenings, developed in the center. Striae visible as transverse lines under light microscopy, but with electron microscopy, finely chambered or alveolate with porous plates in the outer wall. One or two longitudinal lines occasionally running across the striae. Lectotype: Caloneis amphisbaena (Bory) Cleve 1894: 58. SPECIES: Although 250 species and infraspecific names have been reported, only 50 taxa have been currently accepted as taxonomically valid (Guiry and Guiry 2012). Some 13 species were reported in Korea until 1997 (Lee 1997), but only 10 taxa are accepted here. DISTRIBUTION: Most species of this genus are distributed in the inland waters, and a few are in coastal zones. These diatoms inhabit the β- or α- mesosaprobic water of the nine pollution degrees proposed by Fjerdingstad (1964), namely partially or moderately polluted area (Krammer and Lange- Bertalot 1986). KEY REFERENCE: Hustedt (1930), Patrick and Reimer (1966), Krammer and Lange-Bertalot (1986), Round et al. (1990). Key to the species of genus Caloneis 1. Longitudinal line marginally across striae on the each side of the raphe 2 Longitudinal line across striae not developed marginally, but developed near the raphe Tranverse fascia absent in the central area 3 Tranverse fascia developed in the central area 5 3. Valve linear in outline C. tenuis Valve triundulate in outline 4 4. Valve distinctly triundulate in margins C. ventricosa Valve weakly triundulate in margins C. tenuis 5. The fascia not completely or asymmetrically develped C. tenuis The fascia completely developed 6 6. The fascia shallow rectangular in outline C. ventricosa var. truncatula

21 Pennales: Naviculaceae: Caloneis 13 The fascia more or less deep rectangular in outline 7 7. Valve linear in outline 8 Valves linear in outline, but more or less triundulate The central area symmetrical apically C. bacillum The central area asymmetrical along apical axis 9 9. Raphe strongly curved and lateral C. lauta Raphe moderately curved C. leptosoma 10. Valve above 9 μm in breadth C. ventricosa var. subundulata Valve below 9 μm in breadth C. ventricosa var. minuta 11. A single longitudinal line or a narrow band on each side of the raphe 12 Two longitudinal lines or broad band on each side of the raphe The ends of the valve broadly rounded to obtusely protracted C. africana The ends of the valve broadly rounded to obtuse C. crassa 13. The central area more or less circular C. permagna The central area large rhomboidal C. amphisbaena REMARKS: The valve face of this genus resembles that of the genus Pinnularia. The genus Caloneis is closely similar to the related genus, Pinnularia, in many morphological features. The two genera are too closely related and, in some cases, discrimination is difficult. Delimitation may be only possible with electron microscopy. Traditionally, this genus has been separated from the genus Pinnularia by the fine striae and the shape of their alveoli. However, this can not be resolved under the best conditions of light microscopy. Many authors have suggested the further study and the combining two groups into a single genus. Round et al. (1990) have included this genus within the genus Pinnularia. The diatoms of this genus are classified into two representative groups, Caloneis amphisbaena and C. ventricosa (Krammer and Lange-Bertalot 1986). Two groups are distinctly discriminated in the shape and size of the valves, the strongness and alveoli of the striae, and longitudinal bands across the striae. The former resembles more closely to the Pinnularia than the latter. 3. Caloneis africana (Giffen) Stidolph 1995: 167 (Figs. 5, 6). Stidolph 1995: 167. f. 16. Witkowski et al. 2000: 162. pl f. 1. BASIONYM: Caloneis brevis var. distoma f. africana Giffen 1967: 256. pl. 2. f. 23. Valves elliptical in outline, with broadly rounded to Fig. 5. Distribution of Caloneis africana.

22 14 Algal Flora of Korea Freshwater Diatoms VII C D B 10 μm A E Fig. 6. Caloneis africana. A E. view of valves ( 2000, LM).

23 Pennales: Naviculaceae: Caloneis 15 obtusely protracted ends. Raphe straight, the central fissures of the raphe distinctly expanded and distant, and terminal fissures hooked to one side. Axial area narrow at ends, widening towards the middle and central area large circular and occasionally asymmetrical. Striae rows in 10 μm, radiate throughout most of the valve and parallel at the center. Striae crossed by a distinct longitudinal line, placed 1/4 to 1/3 distance from the margins, more or less parallel with the valve margin. Valves μm in length and μm in breadth. ECOLOGY AND DISTRIBUTION: This species is widely distributed in the southern hemisphere (Giffen 1967; Stidolph 1995), and is found in tidal flat of the North Sea and the White Sea (Witkowski et al. 2000). In Korea, it occurs commonly on the sediments in the Nakdong River Estuary. Caloneis africana was recorded in the estuary under the name of C. fenzlii (Grunow) Patrick (Cho 1989). 4. Caloneis amphisbaena (Bory) Cleve 1894: 58 (Figs. 7, 8). Patrick and Reimer 1966: 579. pl. 53. f. 2. Krammer and Lange-Bertalot 1986: 385. pl f. 4. BASIONYM: Navicula amphisbaena Bory SYNONYM: Schizonema amphisbaenum (Bory) Kuntze 1898: 551. Valves broadly lanceolate or elliptical in outline with capitate to rostrate-capitate ends. Raphe filiform, central pores of the raphe distinctly elongated and terminal fissures of the raphe distinct. Axial area widening into a large rhomboidal central area. Striae rows in 10 μm, radiate throughout most of the valve and parallel or convergent towards the ends. Striae crossed by two distinct longitudinal lines between the margin of the valve and the axial area. Valves μm in length and μm in breadth. TYPE LOCALITY: Dans la Marne, sous le pont de Charenton in France. ECOLOGY AND DISTRIBUTION: This species is frequently observed in freshwater to slightly brackish water, and is sometimes classified as a euryhaline species. It was regularly recorded in the estuarine mudflat of a river (Aykulu 1982). Caloneis amphisbaena also was found irregularly in the phytoplanktons collected from 40 reservoirs in Spain (Negro and de Hoyos 2005). In Korea, it was rarely found in epiphytic diatom assemblages of lakes near the Bukcheongnamdaecheon Stream, North Korea (Cho 2000c), and from Ulleung Island (Kang 1973). Fig. 7. Distribution of Caloneis amphisbaena.

24 16 Algal Flora of Korea Freshwater Diatoms VII A B 10 μm C Fig. 8. Caloneis amphisbaena. A C. view of valves ( 2000, LM). REMARKS: This species is characterized by the shape of the ends and the axial area, and the presence of the longitudinal bands. Genus Pinnularia Ehrenberg 1843: 45, nom. et. typ. cons. Bit-sal-dol-mal-sok ( ) Cells solitary or unicellular, very rarely forming band-shaped colonies. Typical naviculoid forms, elongated and bilaterally symmetrical in valve view. Valves usually linear, lanceolate or rarely elliptical in outline, with bluntly rounded, sometimes slightly subrostrate to subcapitate ends. The

25 Pennales: Naviculaceae: Pinnularia 17 valve face flat or curved to form deep mantles. The inner fissures of the raphe usually straight, but the outer fissures of the raphe not superimposed and curved or undulate laterally, resulting in sinuous curves, which sometimes cross over each other along the raphe (see Fig. 9). Some crossings along the raphe are referred to as semicomplex or complex. Some raphes are straightly central in a hyaline axial area, the central ends of the raphe usually expanded, the terminal ends bent to one side with hook shapes resembling question marks, bayonets or commas. Circular thickening called central nodules at the centre and similar thickenings with inwardly projecting lingular structures (helictoglossa) at terminal nodule also present. The forms of terminal ends of the raphe are very varied. Striae basically multiseriate and chambered. Each chamber (alveolus) having a porous plate as its outer wall, containing many rows of small round poroids occluded by hymens. However, these poroids of the striae are invisible in the light microscopy and the striae are only apparent as broad lines. The chambered striae occasionally giving the appearance of one to two longitudinal lines across the striae. The striae sometimes interrupted in the middle parts to form a broad hyaline central area, asymmetrical in many cases, frequently up to the margins of the valve to form a hyaline fascia. In addition, characteristic markings on the central area irregularly present or less frequently in the axial area, bilaterally symmetrical (see Fig. 10). Girdle bands few, the valvocopula with a row of elongate pores, and intercalary bands and septa absent. Two large plate-like plastids lying along the girdle or two plate plastids connecting the center to form a single H-shaped plastid. Lectotype: Pinnularia viridis (Nitzsch) Ehrenberg 1843: 305. SPECIES: Pinnularia has a large number of species and their numbers increased from 221 in the 19th century to the above 1100 during 20th century (Krammer 2000). There are 2465 species and infraspecific names in AlgaeBase at present, of which approximately 400 are currently accepted as taxonomically valid (Guiry and Guiry 2012), but the list is incomplete (Guiry, personal communication). Through the literature survey of Noh (1991), KNCCN (1996) and Lee (1997), 69 taxa of Pinnularia have been described in the inland water and even the brackish water of Korea. DISTRIBUTION: Most Pinnularia species are encountered in freshwater, moist soil or spring, and ponds, and some taxa, up to 10, are brackish or marine. They are abundant in low conductivity and acidic environments of forests and mountainous area (Krammer and Lange-Bertalot 1986; Ciniglia et al. 2007). Their life forms are littoral, benthic or meroplanktonic as relatively dense cells. In the acidic, low conductivity sites overgrown by moss Sphagnum and in the acidic bogs of alpine mountains, epiphytic or benthic diatoms are represented by Eunotia and Pinnularia, and Pinnularia is one of the most important genera in cold regions such as the subantarctic and arctic area (Van de Vijver and Beyens 1997; Van de Vijver et al. 2004; Nováková and Poulíčková 2004; Kulikovskiy 2009a, 2000b). In subantarctic regions, Pinnularia was the second dominant group in soil diatom assemblages of Crozet Archipelago (Van de Vijver et al. 2002). On the other hand, a larger number of tropical Pinnularia species were described by Metzeltin and Lange-Bertalot (1998). Diatoms of the genus Pinnularia are significant components of the benthic diatom assemblages of acidic peatlands (Krammer and Lange-Bertlaot 1986) and occasionally the epilithic diatoms of some streams (Salomoni et al. 2006). Pinnularia was very scarce in planktons collected from lake or reservoirs (Negro and de Hoyos 2005). They were commonly encountered in mudflats off the coast, however, they are never abundant (Aykulu 1982). Pinnularia diatoms are widespread in Korean freshwaters, but are often not abundant; their frequency and abundance are, however, high in low electrolyte environments such as mountain wetlands and swamps. They are unusually the dominant planktons in a swamp of Motjae Mountain in Hapcheon (Lee 1997).

26 18 Algal Flora of Korea Freshwater Diatoms VII 10 μm J G D B H A I C E F Fig. 9. The forms of the raphe in Pinnularia diatoms. A, J. The raphe is filiform, straight and central in the axial area; B F. lateral; B. narrowly lateral; D, E. broadly lateral; G, H. semicomplexly lateral; C. complexly lateral; I. undulately lateral; F. strongly lateral.

27 Pennales: Naviculaceae: Pinnularia μm A B C D E Fig. 10. The forms of markings on the central area. A, B. four large markings or flecks on the central area; C. the scattered puncta; D. the arranged puncta; E. crescent-shaped markings on the central area and small flecks on the axial area. KEY REFERENCE: Hustedt (1930), Patrick and Reimer (1966), Krammer and Lange-Bertalot (1986), Round et al. (1990), Noh (1991). Key to the species of genus Pinnularia 1. The fascia, the hyaline central area up to the valve margin, developed on the central parts of the valve 2 The fascia not developed on the central parts of the valve The ends of the valve differentiated from the body 3 The ends of the valve not differentiated from the body Raphe filiform 4 Raphe not filiform in its fashion Axial area relatively narrow along the axial area 5 Axial area not narrow Transapical striae more than 14 rows in 10 μm (Subgroup 1) 6 Transapical striae less than 14 rows in 10 μm 9 6. Valves more or less linear to slightly lanceolate 7 Valves broadly lanceolate P. metzeltinii 7. Valves linear or linear-lanceolate in outline with slightly convex margins 8

28 20 Algal Flora of Korea Freshwater Diatoms VII The valve linear or linear-lanceolate in outline, but the margins of the valve strongly triundulate P. grunowii 8. Striae on the valve moderately radiate in center and convergent P. sinistra Striae on the valve strongly radiate in the middle and convergent P. divergentissima 9. Valves less than 7 μm in breadth (Subgroup 2) 13 Valves more than 6 μm in lowest breadth (Subgroup 3) Valves more than 9 μm in breadth 11 Valves less than 9 μm in breadth The margins of the valve parallel or slightly concave P. biceps The margins of the valve slightly convex P. subfalaiseana 12. Larger forms, the striae more than 11 in 10 μm P. grunowii Smaller forms, the striae less than 11 in 10 μm P. saprophila 13. The margins of the valve triundulate 14 Different from the above Larger forms with more than 7 μm in breadth P. grunowii Smaller forms with less than 7 μm in breadth P. pluvianiformis 15. The ends of the valve distinctly rostrate or capitate 16 The ends of the valve slightly protracted The ratios of the length and breadth more than 8 P. subcapitata var. subrostrata The ratios of the length and breadth less than 8 P. metzeltinii 17. Striae in the middle strongly radiate and convergent towards the ends P. divergentissima Striae moderately radiate and convergent Striae radiate in the middle and strongly convergent towards the ends P. obscura Striae slightly radiate and convergent P. sinistra 19. Raphe straightly filiform 20 Raphe curved or undulate, not filiform in its fashion Transapical striae more than 14 rows in 10 μm (Subgroup 4) 21 Transapical striae less than 14 rows in 10 μm (Subgroup 5) Valves lanceolate in outlines P. pseudosimilis Valves more or less linear in outlines Striae radiate in the middle parts and convergent towards the ends P. schoenfelder Striae parallel to slightly radiate in the middle parts and slightly convergent towards the ends P. julma 23. Longitudinal costae or irregular flecks on the central and axial area P. subundulata No costae or fleks on the axial area The raphe filiform type, but very strong P. sakurajimensis Not so strong raphe The ends of the valve not protracted and cuneate 26 The ends of the valve not protracted and rounded Striae more than 12 in 10 μm P. brebissonii var. bicuneata Striae less than 12 in 10 μm P. dornii 27. The ends of the valve not protracted and broadly rounded P. obscura The ends of the valve not protracted and narrowly rounded P. similiformis 28. Valves more than 13 μm in lowest breadth (Subgroup 6) P. episcopalis Valves less than 13 μm in maximum breadth (Subgroup 7) Axial area very wide, more than 1/2 the breadth of the valve 30

29 Pennales: Naviculaceae: Pinnularia 21 Axial area less than 1/2 the breadth of the valve The margins of the valve parallel and the ends of the valve braosdly round P. lokana The margins of the valve weakly undulate, and the ends smoothly to weakly protracted and broadly rounded P. cruxarea 31. Valve linear in outline 32 Valve linear with weakly convex margins The valves linear in outline, but the margins of the valve slightly triundulate P. stomatophora var. irregularis The valves linear in outline without triundulate margins Large markings on the central area P. spitsbergensis No markings on the central area Central area broadly developed and extended beyond the central ends of the raphe as a fascia type P. quadratarea Central area not so broad Valves less than 75 μm in length P. eifelana Valves more than 75 μm in length Striae parallal or weakly radiate in the middle P. crucifera Striae radiate in the middle P. subgibba var. sublinearis 37. The terminal ends of the raphe very long P. stomatophora var. salina The terminal ends of the raphe moderate in length P. pseudogibba 38. Some markings or thickenings in axial area, central area or the margin of the central area (Subgroup 8) 39 No markings or thickenings in axial or central area (Subgroup 9) The thickening distinctly developed in the margin of the central area 40 The thickening absent in the margin of the central area Valves lanceolate in outline 41 Valves more linear in outline Thickenings on the margin of the central area P. divergens Markings on the central area P. gibba 42. Valves linear or slightly lanceolate, but narrowing towards the ends P. divergens var. media Valves linear and broadly capitate P. divergens var. undulata 43. Valve linear in outline 44 Valve more or less lanceolate in outline Smaller forms P. lange-bertalotii Larger forms Valve linear in outline and weakly swollen in the middle P. macilenta Valve triundulate in the margin P. graciloides var. triundulata 46. The margins of the valve undulate 47 The margins of the valve not undulate, slightly convex P. parvulissima 47. The margins of the valve slightly triundulate P. graciloides The margins of the valve weakly undulate P. subgibba var. undulata 48. The ends of the valve distinctly capitate 49 Differnet from the above Valves linear to slightly lanceolate and not undulate P. polyonca var. sumatrana Valves linear and triundulate P. perumbrosa 50. The margin of the valve parallel 51

30 22 Algal Flora of Korea Freshwater Diatoms VII The margin of the valve not parallel The ends of the valve broadly rostrate P. rhombarea The ends of the valve cuneate rounded P. eifelana 52. The ratios of valve length and breadth almost 5.8 P. subbrevistriata The ratios of valve length and breadth above Striae radiate in the center and convergent towards the ends P. subgibba Striae strongly radiate in the center and convergent towards the ends P. similiformis var. koreana 54. Valves more than 7 μm in lowest breadth (Subgroup 10) 55 Valves less than 7 μm in maximum breadth (Subgroup 11) Valves strongly triundulate in margins P. nodosa var. robusta Not triundulate The ends of the valve narrowly protracted or capitate 57 Different from the above Broad shoulders on the margins of the valve P. latarea No distinct shoulders on the margins Valves more than 9 μm in breadth P. lundii Valves less than 9 μm in breadth Striae more delicate 60 Striae strong and more radiate in the middle P. certa 60. Valves lanceolate and striae partly curved P. brauniana Valves broadly linear and the margins of the valve parallel P. valdetolerans 61. The ends of the valve not protracted or slightly protracted P. microstauron The ends of the valve distinctly protracted The ends of the valve broadly protracted or constricted near the ends 63 The ends of the valve not capitate The ends of the valve capitate and constricted beneath the ends P. globiceps More elongated forms P. globiceps var. linearis 64. Axial area 1/4 1/3 of the valve breadth P. pulchra Axial area more than 1/2 of the valve breadth P. scotica 65. Valve broadly linear in outline P. globiceps Valves narrowly linear or lanceolate The margins of the valve weakly triundulate P. pulchra The margins of the valve linear to elliptic-lanceolate P. marchica 67. Longitudinal bands cross the striae 68 No longitudinal bands cross the striae Transapical striae less than 7 rows in 10 μm 69 Transapical striae more than 7 rows in 10 μm Raphe lateral (Subgroup 12) 70 Raphe semicomplex (Subgroup 13) 76 Raphe complex (Subgroup 14) Valves linear and slightly swollen in the middle and the ends P. neomajor var. inflata Valve linear to slightly lanceolate in outline Axial area very broad, more than 1/3 breadth 72 Axial area more or less broad, less than 1/3 breadth The ratio of the length and breadth more than 5 P. brevicostata

31 Pennales: Naviculaceae: Pinnularia 23 The ratio of the length and breadth less than 5 P. pseudacuminata 73. Valves slightly lanceolate, narrowing towards the ends P. rhomboelliptica Valves more or less linear in outline Axial area broad, 1/3 the valve breadth, and central area moderately wide P. gigas Axial area moderately broad, but 1/5 1/4 the valve breadth, and central area not distinct Striae more coarse, 6 7 rows in 10 μm and valve more than 21 μm in breadth P. viridis Striae less coarse, 7 9 rows in 10 μm and valve less than 21 μm in breadth P. viridiformis 76. Valve less than 21 μm in breadth 77 Valve more than 21 μm in breadth Valves slightly lanceolate narowing towards the ends P. rhomboelliptica Valves more or less linear The ends of the valve not swollen P. viridis The ends of the valve swollen P. gentilis 79. Larger form and axial area more wide P. viridiformis Smaller form and axial area less wide P. viridiformis var. minor 80. Valves linear in outline, with broadly rounded ends, and weakly swollen in the middle and at the ends P. nobilis var. regularis Valves linear in outline with cuneately rounded ends, and slightly swollen in the middle P. subgigas 81. Valves less than 16 μm in lowest breadth of the valve (Subgroup 15) 82 Valves more than 13 μm in maximum breadth The axial area very broad, 1/2 or more than the valve breadth P. cruxarea The axial area moderately broad or narrow Valves lanceolate P. subrhombica Valves more or less linear Valves narrowly linear 85 Valves broadly linear Central area very narrow fascia up to the margins of the valve P. trifonovae No fascia form in the central area The ends of the valve broadly rounded P. subcommutata The ends of the valve cuneately rounded P. peracuminata 87. Larger form and axial area more wide P. viridiformis Smaller form and axial area less wide P. viridiformis var. minor 88. Raphe semicomplex to complex in its fashion (Subgroup 16) 89 Raphe lateral (Subgroup 17) Valves less than 130 μm in length P. viridiformis Valves more than 130 μm in length The valves less than 25 μm in breadth P. viridis The valves more than 25 μm in breadth P. gentilis 91. Valves linear in outline and the ends of the valve rounded 92 Valves linear or slightly lanceolate, but swollen in the middle and ends of the valve Axial are very wide, 1/3 breadth of the valve P. pseudacuminata Axial are narrow or moderately wide, less than 1/3 breadth of the valve The valves weakly lanceolate, narrowing towards the ends P. rhomboelliptica The valves more or less linear P. viridiformis

32 24 Algal Flora of Korea Freshwater Diatoms VII 94. Large swollen in the middle and ends P. transversa Weakly swollen in the middle and ends P. neomajor var. frequentis 95. The ends of the valve differentiated from the body 96 The ends of the valve not differentiated from the body Raphe filiform (Subgroup 18) 97 Raphe not filiform in its type (Subgroup 19) The ends of the valve not protracted P. subrabenhorstii The ends of the valve protracted The ends of the valve capitate 99 The ends of the valve more and less protracted, not capitate Hyaline fascia of central area reaching the valve margins 100 Central area not fascia form, but rhombic in the middle Hyaline fascia of central area widely developted up to the margins P. subcapitata Hyaline fascia narrowly developted without one or two striae P. angliciformis 101. Smaller form, the valves less than 42 μm in length P. subinterrupta Larger form, the valves more than 42 μm in length P. angliciformis 102. The breadth of the valve more than 8 μm 103 The breadth of the valve less than 8 μm P. pulchra 103. Axial area broad P. subgibba Axial area narrowly linear P. microstauron var. nonfasciata 104. Striae extremely strong and wide P. rabenhorstii Striae moderately wide Valves linear, and the margins of the valve parallel and weakly convex P. subgibba Valves rhombic-lanceolate and the margins of the valve convex P. erratica 106. Valves more than 10 μm in breadth (Subgroup 20) 107 Valves less than 10 μm in lowest breadth of the valve (Subgroup 21) Irregular puncta or flecks on the axial and central area P. acrosphaeria Puncta not present on the hyaline area of the valve The axial area very wide, about 1/2 the breadth of the valve P. cruxarea The axial area narrow or moderately wide Striae more and less delicate 110 Striae very strongly broad and distant between two striae Central area large rhombic to rhombic-circular 111 Central area irregularly round P. islandica 111. Valves linear to lanceolate P. karelica Valves linear in outline P. mormonorum 112. Valves less than 14 μm in breadth and striae less strong P. rabenhostii var. franconica Valves more than 14 μm in breadth and striae more strong P. lata 113. Axial area more than 1/2 breadth of the valve P. cruxarea Axial area less than 1/2 breadth of the valve Striae very wide and strong, 3 6 rows in 10 μm 115 Striae not so wide and general typed Valves linear to linear-elliptical in outline P. borealis Valves linear in outline Valves linear with bluntly rounded ends P. borealis var. scalaris Valves linear or rectangular in outline with bluntly rounded 117

33 Pennales: Naviculaceae: Pinnularia The striae more than 4 in 10 μm P. borealis var. sublinearis The striae less than 4 in 10 μm P. angulosa 118. The ends of the valve broadly rounded P. subcommutata The ends of the valve cuneately rounded P. peracuminata REMARKS: Pinnularia is characterized by chambered striae and longitudinal lines, the expanding central ends of the raphe and the hooked terminal ends of the raphe. However, species boundaries are often poorly defined, especially with Caloneis. This genus is closely related to Caloneis and, as discussed earlier, they are often considered as the same genus. Great confusion has existed in the various aspects in the taxonomy of diatom Pinnularia. For example, in taxa related to Pinnularia viridis, the stretch of the raphe fissure has been considered as an important feature, but the extent of the variation was not noticed. As the raphe is not consistent with other features of the valve, some authors described many hybrids, such as P. maior P. viridis, P. viridis P. hemiptera, P. gentilis P. maior. Mayer (1936) also included such hybrids in his sudy Pinnularia diatoms collected in Bavaria region of Germany. However, we have no firm evidence that Pinnularia diatoms form hybrids, although they are certainly possible (Krammer and Lange- Bertalot 1986). In addition, the type of plastids is potentially useful for the taxonomy of the genus to discover the relationships between both plastids and valve features (Cox 1988). 5. Pinnularia acrosphaeria W. Smith 1853: 58 (Figs. 11, 12). Hustedt 1930: 330. f Patrick and Reimer 1966: 623. pl. 60. f. 2. Krammer and Lange-Bertalot 1986: 409. pl f. 1. Noh 1991: 61. f. 19. Chung 1993: 288. f Krammer 2000: 54. pl. 19. f. 1. BASIONYM: Frustulia acrosphaeria Brébisson 1838: 19. SYNONYM: Pinnularia acrospharia Rabenhorst 1853: 45. pl. 6. f. 36. NON SYNONYM: Frustulia acrosphaeria Brébisson 1838: 19. Valves linear in outline, the margins of the valve parallel swollen in the middle, and the ends of the valve broadly rounded to swollen, not protracted. Raphe strongly lateral, the outer fissures of the raphe straight in the middle, the inner fissure of the raphe strongly lateral, but difficult to see under the light microscope, the central ends of the raphe expanded and strongly curved in one direction, central pores small round and close together, the terminal ends of the raphe large shaped like a sickle. Axial area broad and linear in outline and slightly widening in the central area, usually asymmetrical. Large puncta or gra- Fig. 11. Distribution of Pinnularia acrosphaeria.

34 26 Algal Flora of Korea Freshwater Diatoms VII A B 10 μm C D E F Fig. 12. Pinnularia acrosphaeria. A F. view of valves ( 2000, LM). nules densely scattered in the axial and central area. Striae parallel to weakly radiate in the middle and weakly convergent towards the ends, rows in 10 μm. Longitudinal lines marginal across the striae, but hardly visible in the light microscopy. Valves μm in length and μm in breadth. TYPE LOCALITY: Freshwater near Lewes, W. Smith. Premnay Peat, Dolgelly Earth (Smith 1953: 58 [England]). ECOLOGY AND DISTRIBUTION: The main distribution area of this species is tropical regions and it is most common in freshwaters of mountains (Krammer and Lange-Bertalot 1986). Pinnularia acros-

35 Pennales: Naviculaceae: Pinnularia 27 phaeria seldom occurs in plains and is distributed the littoral areas of circumneutral lakes and ponds. In Korea, this species has a wide distributions in freshwater and is found in mountains, streams, reservoirs and lakes and swamps in the lowlands near rivers (Noh 1991; Chung 1993). This species was particularly dominant in the planktons of Motjaeneup, a pond in the mountainous area at Hapcheon (Lee 1997). It is collected recently from the epiphyton algal assemblages of the Seonakdong River, but was very rare. REMARKS: Frustulia acrosphaeria Brébisson and its synonym Navicula acrosphaeria sensu Kützing represent a species separable to Pinnularia acrosphaeria (Krammer 2000). 6. Pinnularia angulosa Krammer 2000: 27 (Figs. 13, 14). Hustedt 1930: 326. f Krammer 2000: 27. pl. 6. f. 11. REPLACED SYNONYM: Pinnularia borealis var. brevicostata Hustedt 1914: 82. SYNONYM: Navicula borealis var. brevicostata (Hustedt) Prochazka 1923: 80. Valves linear and rectangular in outline, the margins of the valve parallel with broadly rounded ends. Raphe moderately lateral, the central fissures of the raphe somewhat reverse-lateral, central pores distinctly rounded, and the terminal fissures of the raphe sickle-shaped. Axial area wide and central area not distinctly developed. Striae broad and distant between the Striae 3 4 rows in 10 μm, and parallel over the valve. Valves μm in length and about 10 μm in breadth. 10 μm A B C Fig. 13. Pinnularia angulosa. A C. The morphology of valves ( 2000, LM).

36 28 Algal Flora of Korea Freshwater Diatoms VII TYPE LOCALITY: Sudeten: Oder Valley, Eulengebirge, Heinrichau, Silesia in Poland, Central Europe. ECOLOGY AND DISTRIBUTION: Pinnularia angulosa is found in the type locality of the Central Europe, lakes in the North Finland (Krammer 2000), and from 476 localities of various water bodies on Mountain Shara and Nidze, Macedonia (Levkov et al. 2005). In Korea, it is rarely found in two peatlands, the first Mujechineup and Anjeokneup, in Mountain Jeongjok, Ulsan and from a channel of rice paddies in Gochang. This species is the first report from Korea. REMARKS: Pinnularia angulosa is differentiated from P. borealis series species to show the quasi rectangular outline of the valve, the marginal striae and the missing central area, and the reverse lateral raphe (Krammer 2000). Fig. 14. Distribution of Pinnularia angulosa. 7. Pinnularia borealis Ehrenberg 1843: 420 (Figs. 15, 16). Patrick and Reimer 1966: 618. pl. 58. f. 11. Krammer and Lange-Bertalot 1986: 405. pl f. 1. Noh 1991: 50. f. 17. Chung 1993: 289. f Krammer 2000: 24. pl. 6. f. 5. SYNONYM: Navicula borealis (Ehrenberg) Kützing 1844: 96. pl. 28. f. 68. Pinnularia chilensis Bleisch 1859: 28. pl. 4. f. 1. Schizonema boreale (Ehrenberg) Kuntze 1898: 552. Valves linear to linear elliptical in outline, with parallel margins and bluntly rounded ends. Raphe filiform to weakly lateral, the outer fissures of the raphe lateral in the middle, the central ends of the raphe slightly expanded and curved in the same direction, central pores distinctly round, and the terminal ends of the raphe resembling a large sickle. Axial area narrow and central area roundly developed with one or two short striae in middle. Striae coarsely broad, distant between the striae, 5 6 rows in 10 μm, parallel or slightly radiate at centre. Valves μm in length and μm in breadth. SYNTYPE LOCALITY: Chile (?); New York: Kotzebue Sound, Alaska. ECOLOGY AND DISTRIBUTION: This species is widely distributed in the moist environments of lowlands and mountain. As a well known aerophilic or xerophytic diatom, it is dominant in moss epi-

37 Pennales: Naviculaceae: Pinnularia 29 A B C D 10 μm E F G Fig. 15. Pinnularia borealis. A G. view of valves ( 2000, LM). phytons in the cold subantarctic or subarctic regions (Vande Vijver and Beyens 1997; Kulikovskiy 2009a, 2009b). In these cases, the major ecological importance seems to be the moisture content of the samples and the size of Pinnularia borealis decreases significantly depending on the dryness of the habitat (Vande Vijver and Beyens 1997). It is one of a few taxa tolerating desiccation and freezing stress experimentally (Souffreau et al. 2010). However, in a study of diatom assemblages on the sediment collected from 30 clear Florida lakes, it is considered as eutrophic and circumneutral species (Whitmore 1989). Furthermore, Pinnularia borealis is frequently encountered in the sediment in river estuaries and adjacent regions, never reaching high abundance (Obrezkova 2009). In certain assemblages, the nominal variety and other varieties are dominant in the palaeological records (Razjigaeva et al. 2004). Fig. 16. Distribution of Pinnularia borealis.

38 30 Algal Flora of Korea Freshwater Diatoms VII In Korea, it is one of the most frequent species of the Pinnularia and was found frequently in mountain streams (Chung and Lee 1979, 1982), occasionally rivers (Chung et al. 1968) and lakes (Chung and Kay 1969; Kim et al. 1995). Pinnularia borealis occurs abundantly in mosses attached to rocks in Mountain Juwang, Jiri and Odae (Noh 1991), and to trees of Mountain Halla in Jeju Island, August REMARKS: More than 50 infraspecific taxa are listed in the AlgaeBase database (Guiry and Guiry 2012), but in many cases, diagnoses and observations conducted on a single specimen are difficult to characterize (Krammer 2000). The morphological scopes of P. borealis and its varieties are not clear, and it is difficult to find populations with a sufficient numbers of specimens for the interpretation of the morphological range of a taxon (Kulikovskiy et al. 2010a). 8. Pinnularia borealis var. scalaris (Ehrenberg) Rabenhorst 1864: 216 (Figs. 17, 18). Krammer 2000: 25. pl. 8. f. 10. BASIONYM: Stauroptera scalaris Ehrenberg 1843: 423. f. 4/2. SYNONYM: Stauroneis scalaris (Ehrenberg) Kützing 1844: 106. pl. 29. f. 37C. Navicula borealis var. scalaris (Ehrenberg) Grunow 1860: 518. pl. 2. f. 15. Navicula scalaris (Ehrenberg) O Meara 1875: 354. pl. 30. f. 39. Valves linear to linear elliptical in outline with bluntly rounded ends. Raphe filiform to weakly lateral, the outer fissures of the raphe lateral in the middle, the central ends of the raphe slightly expanded and curved in the same direction, central pores distinctly round, and the terminal ends of the raphe resembling a large sickle. Axial area narrow and central area roundly developed. Striae coarsely broad, distant between the striae, 4 5 rows in 10 μm, parallel or slightly radiate at the center. Valves (56) μm in length and 8 9 μm in breadth. TYPE LOCALITY: Okak, Labrador, Canada. ECOLOGY AND DISTRIBUTION: This variety is similar to the nominal variety in the ecological characteristics, for example in the aerophytic diatom assemblages. Pinnularia borealis var. scalaris was abundant in diatom assemblages attached to mosses from Heard Island (Vande Vijver et al. 2004) and in soil diatom assemblages of Crozet Archipelago (Van de Vijver et al. 2002) in the Subantarctic. In Korea, it is found less commonly from the Yongneup Bog of Mountain Daeam in Inje and this is the first report from Korea. Fig. 17. Distribution of Pinnularia borealis var. scalaris.

39 Pennales: Naviculaceae: Pinnularia 31 F A B C D E 10 μm Fig. 18. Pinnularia borealis var. scalaris. A F. view of valves ( 2000, LM). REMARKS: The variety is similar to the nominal variety, however, the size of the valve is larger and the ends are more broadly rounded. 9. Pinnularia borealis var. sublinearis Krammer 2000: 25 (Figs. 19, 20). Krammer 2000: 25. pl. 7. f. 14. Valves linear or rectangular in outline with bluntly rounded. Raphe filiform to weakly lateral, the outer fissures of the raphe lateral in the middle, the central ends of the raphe slightly expanded and curved in the same direction, central pores distinctly round, and the terminal ends of the raphe resembling a large sickle. Axial area narrow and central area developed by attenuation of one or two striae in the center. Striae coarsely broad, distant between the striae, 4 6 rows in 10 μm, parallel or slightly radiate at the center. Valves μm in length and 7 8 μm in breadth. TYPE LOCALITY: Erlangen in Germany, town wall, on moss.

40 32 Algal Flora of Korea Freshwater Diatoms VII A B 10 μm C D E F G Fig. 19. Pinnularia borealis var. sublinearis. A G. view of valves ( 2000, LM). ECOLOGY AND DISTRIBUTION: This variety is frequent on wet and nearly dry walls (Krammer 2000) and a mountain system, the Galician Central Macizo, located in the northwestern regions of Spain (Rodríguez et al. 2007). In Korea, it was first observed rarely from a peatland of Anjeokneup-C Bog near Mountain Jeongjok, Yangsan, the first report from Korea. REMARKS: The variety is more linear and narrow in the valve outline than the nominal variety. Fig. 20. Distribution of Pinnularia borealis var. sublinearis.

41 Pennales: Naviculaceae: Pinnularia Pinnularia brauniana (Grunow) Mills 1934: 1273 (Figs. 21, 22). Patrick and Reimer 1966: 594. pl. 55. f. 3. Krammer and Lange-Bertalot 1986: 416. pl f. 1. Noh 1991: 132. f. 48. Krammer 2000: 112. pl. 86. f. 10. SYNONYM: Navicula brauniana Grunow in A. Schmidt et al pl. 45. f. 77. Navicula braunii Grunow in VanHeurck 1880: 79. f. 6. Pinnularia braunii (Grunow) Cleve 1895: 75. A B C D E F G H 10 μm I J K L M N O Fig. 21. Pinnularia brauniana. A O. view of valves ( 2000, LM).

42 34 Algal Flora of Korea Freshwater Diatoms VII Valves lanceolate in outline, magins of the valve convex with distinctly capitate ends. Raphe straight to slightly lateral, the central ends of the raphe slightly curved to one side, central pores small round, the terminal ends of the raphe shaped like question marks. Axial area linear or linear-lanceolate and central area broadly rhombic to rectangular reaching the valve margins. Striae, rows in 10 μm on valves, radiate at the centre and convergent over the ends. Longitudinal lines absent across the striae along the apical axis. Valves μm in length and 7 9 μm in breadth. TYPE LOCALITY: Bath mud, Loka in Sweden. ECOLOGY AND DISTRIBUTION: Pinnularia brauniana is widespread and generally distributed in low electrolyte waters and dystrophic environments from the plains to mountain heights (Krammer and Lange- Bertalot 1986). It is mostly scattered in distribution, but may occasionally be found in large numbers. In a study of diatom assemblages on the sediment collected from 30 clear Florida lakes, Whitmore (1989) concluded that it is a mesotrophic, acidophilous and acidobiontic taxon. On the other hand, Pinnularia brauniana has been frequently considered as species typical Fig. 22. Distribution of Pinnularia brauniana. of polluted environments or α-mesosaprobic (Van Dam et al. 1994) and occurs abundantly from the epilithic diatoms of the Gravataí River, Brazil (Salomoni et al. 2006). It occurs commonly in Honghe Wetland of China (Zhang et al. 2011). This taxon is dominant in palaeological sediment profile from Laguna Zoncho in southern Pacific Costa Rica (Haberyan and Horn 2005). In Korea, this species is found primarily from streams (Noh 1991) and other water bodies, rivers (Yoo and Lim 1990), swamps in lowland (Chung and Noh 1987). It is recently found from peatlands of Mountain Jeongjok in Yangsan and Mountain Daeam in Inje, and from a rice paddy in Wando, July REMARKS: The following varieties do not belong to this nominal variety, P. brauniana (Krammer 2000). One is a more linear-lanceolate outline and smaller axial area, P. braunii var. amphicephala sensu Hustedt (Hustedt f. 578), elliptic-lanceolate specimens with broad capitate to truncate ends (Kobayashi 1963). Another variety is perhaps represented by two specimens illustrated by Krammer and Lange-Bertalot (1986. pl f. 4, 5). 11. Pinnularia crucifera Cleve-Euler 1934: 48 (Figs. 23, 24). Krammer and Lange-Bertalot 1986: 410. pl f. 4. Krammer 2000: 144. pl f. 1. REPLACED SYNONYM: Pinnularia brevicostata var. elongata Cleve-Euler in Brander 1933: 27. f. 4f.

43 Pennales: Naviculaceae: Pinnularia μm A B C D E Fig. 23. Pinnularia crucifera. A E. view of valves ( 2000, LM). Pinnularia crucifera var. elongata (Cleve-Euler) Cleve-Euler 1955: 36. f. 1044a. SYNONYM: Pinnularia brevicostata var. leptostauron Cleve 1891: 25. Cleve 1895: 86. Pinnularia brevicostata sensu Krammer 1992a. pl. 50. f. 1. Valves broadly linear in outline and the margins of the valve parallel, sometimes slightly convex in the center, with broadly rounded, not protracted ends. Raphe broadly lateral, commonly three lon-

44 36 Algal Flora of Korea Freshwater Diatoms VII gitudinal lines visible, the outer fissures of the raphe slightly undulate, the central ends of the raphe reverselaterally bent, the central pores small and round, the terminal ends of the raphe large sickle-shaped. Axial area linear, 1/3 of the valve breadth, and central area rectangular up to the margin to form a fascia. Striae 8 10 rows in 10 μm on valves, nearly parallel in the middle and parallel to slightly convergent towards the ends. A longitudinal line present along the valve. Valves μm in length and μm in breadth. TYPE LOCALITY: Frederiksbergsmossen near Helsinki, Finland. Finnish Lapland: Käkkälö District; L. Kuolajärvi, L. Aapajärvi; southern Sweden. ECOLOGY AND DISTRIBUTION: Pinnularia crucifera is cosmopolitan in oligotrophic to dystrophic and low electrolyte water of the Nordic and alpine regions (Krammer and Lange-Bertalot 1986). This species is primarily a benthic form and scattered in distribution. In Korea, this species is rarely observed in a peatland, the Yongneup Bog of Mountain Daeam in Inje and this is the first report from Korea. REMARKS: This taxon is closely related to P. brevicostata Cleve 1891, P. isostauron (Ehrenberg) Cleve 1895, P. Fig. 24. Distribution of Pinnularia crucifera. conifera (Brun and Héribaud) Krammer 2000 and P. hustedtii Mölder 1951, but distinguishable on the basis of valve outline, size, axial area, the structure of striae and central area (Krammer 2000). 12. Pinnularia divergens W. Smith 1853: 57 (Figs. 25, 26). Patrick and Reimer 1966: 603. pl. 56. f. 1. Krammer and Lange-Bertalot 1986: 407. pl f. 3. Noh 1991: 83. f. 28. Krammer 2000: 60. pl. 28. f. 1. BASIONYM: Pinnularia divergens W. Smith 1853: 57. pl. 18. f SYNONYM: Navicula divergens (W. Smith) Grunow 1860: 523. Schizonema schweinfurthii Kuntze 1898: 550. Valves broadly lanceolate, linear-elliptical in outline with slightly convex and triundulate margins and broadly rounded, subrostrate to rostrate ends. Raphe lateral, the outer fissure of the raphe undulate, the central ends of the raphe bent in the same direction, central pores with lateral appendages, the terminal ends of the raphe broadly bayonet-shaped. Axial area linear or linear-lanceolate, usually 1/4 of the valve breadth, and central area broadly rhombic to roundedrhombic, reaching the valve margins. Round thickenings developed at the margins of the central area and distinctly visible with careful focusing under the light microscopy. Striae 9 11 rows in 10

45 Pennales: Naviculaceae: Pinnularia 37 A B 10 μm C D Fig. 25. Pinnularia divergens. A D. view of valves ( 2000, LM). μm on valves, strongly radiate at center and extremely convergent over the ends. Longitudinal lines absent across the striae along the apical axis. Valves μm in length and μm in breadth. TYPE LOCALITY: Premnay Peat, Aberdeenshire, Scotland. ECOLOGY AND DISTRIBUTION: Pinnularia divergens occurs as benthic forms in various types of water bodies, primarily in mountainous regions, but relatively rare in lowlands (Krammer and

46 38 Algal Flora of Korea Freshwater Diatoms VII Lange-Bertalot 1986; Krammer 2000). This is one of the most common diatoms and dominant in autumn in Honghe Wetland, China (Zhang et al. 2011), and is distributed in oligotrophic, less acidic and low electrolyte water. In Korea, this species is first reported from a mountain reservoir (Noh 1991), and after then from rivers (Lee and Baik 1994), lakes (Lee 1994), and Yongneup Bog of Mountain Daeam in Inje, May REMARKS: Some 58 infraspecific taxa of Pinnularia divergens are listed in AlgaeBase database (Guiry and Guiry 2012), and are classified largely by the conical wall thickening in the central area and the terminal fissures of the raphe (Krammer 2000). Fig. 26. Distribution of Pinnularia divergens. 13. Pinnularia divergentissima (Grunow) Cleve 1895: 77 (Figs. 27, 28). Patrick and Reimer 1966: 616. pl. 58. f. 8. Krammer and Lange-Bertalot 1986: 419. pl f. 3. Krammer 2000: 44. pl. 11. f. 1. BASIONYM: Navicula divergentissima Grunow in Cleve and Möller 1879: 186. SYNONYM: Schizonema divergentissimum (Grunow) Kuntze 1898: 552. Pinnularia martinii f. martinii Krasske 1939: 394. pl. 11. f. 29. Pinnularia fottii Bily et Marvan 1959: 66. pl. 1. f. 4. Valves linear to linear-lanceolate in outline with slightly convex margins, obtusely rounded and broadly subrostrate ends, occasionally with subrostrate to subcapitate ends. Raphe straight, the central ends of the raphe slightly curved in the same direction, the central pores small, the terminal ends of the raphe shaped like question marks. Axial area narrowly linear and central area with broad fascia reaching the margins. Striae rows in 10 μm on valves, strongly radiate at centre and extremely convergent over the ends with an acute angle between the directions of the two striae. Valves μm in length and about 6 μm in breadth.

47 Pennales: Naviculaceae: Pinnularia 39 A B C D E F G H I J K 10 μm Fig. 27. Pinnularia divergentissima. A K. view of valves ( 2000, LM). TYPE LOCALITY: Fogstuen, Dovre in Norway. ECOLOGY AND DISTRIBUTION: Pinnularia divergentissima is cosmopolitan in northern alpine area, particularly in aerophytic mosses of forest and meadow, and low electrolyte waters (Krammer and Lange-Bertalot 1986; Krammer 2000). In Korea, this species is found mainly in streams (Chung and Kim 1970; Hong and Chung 1990), a mountain reservoir (Noh 1991), and is recently observed in peatlands in mountains, bogs. REMARKS: Pinnularia divergentissima is characterized by the strongly divergent striae. Fig. 28. Distribution of Pinnularia divergentissima.

48 40 Algal Flora of Korea Freshwater Diatoms VII 14. Pinnularia eifelana (Krammer) Krammer 2000: 133 (Figs. 29, 30). Lange-Bertalot and Metzeltin pl. 54. f. 5. Krammer 2000: 133. pl f. 9. BASIONYM: Pinnularia esoxiformis var. eifeliana Krammer 1992: 146. pl. 58. f. 5. Valves linear in outline and the margins of the valve parallel with cuneate rounded ends. Raphe moderately broad lateral, the outer fissures of the raphe curved, the central ends of the raphe bent to one side, central pores large, drop-shaped and not adjacent, the terminal ends of the raphe shaped like question marks. Axial area narrowly linear to moderately broad, 1/5 1/3 of the valve breadth, central area rectangular up to the margin of the valve. Striae 7 10 rows in 10 μm on valves, parallel to slightly radiate in the middle, parallel to slightly convergent towards the ends. A longitudinal band distinct along the apical axis. Valves μm in length and 8 11 μm in breadth. TYPE LOCALITY: Kleine Kyll, Eifel, Rheinland, Germany. ECOLOGY AND DISTRIBUTION: Pinnularia eifelana is most commonly found in oligotrophic and A B 10 μm C Fig. 29. Pinnularia eifelana. A D. view of valves ( 2000, LM). D

49 Pennales: Naviculaceae: Pinnularia 41 moderate electrolyte water in northern Europe and is not rare (Krammer 2000). In Korea, this species is found only from Yongneup Bog of Mountain Daeam in Inje, May It is the first report from Korea. REMARKS: This species can be differentiated from related taxa by the size and outline, and the distinct longitudinal bands. Fig. 30. Distribution of Pinnularia eifelana. 15. Pinnularia episcopalis Cleve 1891: 27 (Figs. 31, 32). Hustedt 1930: 323. f Krammer and Lange-Bertalot 1986: 408. pl f. 1. Krammer 2000: 66. pl. 42. f. 2. SYNONYM: Navicula episcopalis (Cleve) M. Peragallo 1903: 551. Schizonema episcopale (Cleve) Kuntze 1898: 552. Valves broadly linear in outline and the margins of the valve parallel with broadly rounded ends. Raphe lateral and broad, the outer fissures of the raphe hardly curved, the inner fissures of the raphe lateral, the central ends of the raphe straight, central pores with terminal appendages or perpendicular tails, the terminal ends of the raphe shaped like a bayonet. Axial area broadly linear, 1/3 of the valve breadth, central area rectangular up to the margin of the valve. In the central area and adjacent area, irregular markings or structures distinct. Striae 7 10 rows in 10 μm on valves, radiate in the middle, convergent towards the ends. A longitudinal band absent and a longitudinal line running near the axial area. Valves μm in length and μm in breadth. TYPE LOCALITY: Lake Wener, Sweden. ECOLOGY AND DISTRIBUTION: Pinnularia episcopalis is widely distributed, but rarely found in oligo-

50 42 Algal Flora of Korea Freshwater Diatoms VII 10 μm C D A B E Fig. 31. Pinnularia episcopalis. A E. view of valves (A: 750, B E: 1500, LM).

51 Pennales: Naviculaceae: Pinnularia 43 trophic, low electrolyte and moderately acidic freshwaterss of mountains (Krammer 2000) and in a lake and swamp of mountains in China (Liu et al. 2007). Frequently, it is found in fossil in the palaeological study. In Korea, this species is found only a mountain peatland and is the first report from Korea. However, P. cardinaliculus Cleve as collected from an oligotrophic reservoir by Noh (1991) may be identical with this species. REMARKS: Pinnularia novaezealandica Krammer is closely similar to P. episcopalis, but is distinguished by the cuneate ends and the broad longitudinal band (Krammer 2000). Fig. 32. Distribution of Pinnularia episcopalis. 16. Pinnularia gibba Ehrenberg 1843: 384 (Figs. 33, 34). Hustedt 1930: 327. f Krammer and Lange-Bertalot 1986: 423. pl f. 1. Noh 1991: 108. f. 38. Chung 1993: 289. f Krammer 2000: 93. pl. 68. f. 1. SYNONYM: Stauroptera gibba Ehrenberg 1843: 423. pl. 1/2. f. 3. Navicula stauroptera Grunow 1860: 516. f. 2. Navicula abaujensis Pantocsek 1889: 41. pl. 3. f. 54. Valves rhombic-lanceolate in outline, gradually tapering towards the broadly rounded and subcapitate to capitate ends. Raphe lateral and narrow, the outer fissures of the raphe slightly undulate, the central ends of the raphe expanded and bent in one direction, central pores rounded and close together, the terminal ends of the raphe shaped like large question marks. Axial area 1/2 2/3 of the valve breadth, widening from the ends towards the center to form a large rhombic hyaline area and a fascia up to the margins. Four large markings usually apparent in the hyaline central area. Striae 8 11 rows in 10 μm, radiate in the middle and convergent in the ends. Longitudinal lines absent across the striae. Valves μm in length and μm in breadth.

52 44 Algal Flora of Korea Freshwater Diatoms VII A B C 10 μm D E F G Fig. 33. Pinnularia gibba. A G. view of valves ( 2000, LM). TYPE LOCALITY: Chile. ECOLOGY AND DISTRIBUTION: Pinnularia gibba is widespread in freshwaters with low to average electrolyte from lowlands to mountainous area (Krammer and Lange-Bertalot 1986). This species is most abundant and widespread in Korean freshwaters, ponds and lakes, streams and rivers, swamps in lowlands and peatlands of mountains (Noh 1991; Chung 1993). It was found with very low frequency in epiphytons collected from a polluted water body, the Seonakdong River in

53 Pennales: Naviculaceae: Pinnularia REMARKS: This species is variable in valve forms and Krammer (2000) defined P. gibba with a narrow sense. It is best distinguished from the neighboring taxa by characteristic axial area and the capitate ends. Fig. 34. Distribution of Pinnularia gibba. 17. Pinnularia gigas Ehrenberg 1843: 421 (Figs. 35, 36). Patrick and Reimer 1966: 632. pl. 62. f. 4. Krammer and Lange-Bertalot 1986: 430. pl f. 1. Krammer 2000: 178. pl f. 2. SYNONYM: Pinnularia dactylus Ehrenberg pl. 2/2. f. 4. NON SYNONYM: Pinnularia dactylus Ehrenberg 1843: 421. pl. 4/1. f. 3. Valves broadly linear in outline, the middle parts of the valve slightly swollen and the ends of the valve smoothly to cuneiform rounded. Raphe broadly lateral, the outer fissures of the raphe slightly S-shaped, the central ends of the raphe bent to one side, the central pores small round, the terminal ends of the raphe shaped like question marks. Axial area moderately broad, 1/3 the valve breadth, and central area moderately widening, commonly asymmetrical and irregular. Striae strong, 4 5 in 10 μm, radiate in the middle, slightly convergent towards the ends. A longitudinal line across the striae along apical axis. Valves μm in length and μm in breadth. TYPE LOCALITY: New York, USA. ECOLOGY AND DISTRIBUTION: Pinnularia gigas is probably cosmopolitan in low electrolyte water of northern alpine area, particularly in mountain (Krammer and Lange-Bertalot 1986). In Korea, this species is rarely found in mountain wetlands, Duncheolneup Bog of Mountain Duncheol in San-

54 46 Algal Flora of Korea Freshwater Diatoms VII A B 10 μm C D Fig. 35. Pinnularia gigas. A D. view of valves (A, B: 600, C, D: 800, LM).

55 Pennales: Naviculaceae: Pinnularia 47 cheong and an unnamed bog in Daegwanryeong Pasture. It is the first report from Korea. REMARKS: Ehrenberg described two species, P. gigas and P. dactylus, in Although further studies are required, A. Cleve confused both taxa and his description was not of P. dactylus, but of P. gigas in 1895 (Krammer 2000). Subsequent authors followed the incorrect taxonomic concept of Cleve. 18. Pinnularia globiceps var. linearis Krammer 2000: 97 (Figs. 37, 38). Krammer 2000: 97. pl. 74. f. 10. SYNONYM: Pinnularia lundii sensu Krammer pl. 17. f. 7. Valves linear in outline, margins of the valve slightly triundulate through the entire valve with rounded ends. Raphe straight or slightly lateral in the center, Fig. 36. Distribution of Pinnularia gigas. A B C D E F 10 μm Fig. 37. Pinnularia globiceps var. linearis. A F. view of valves ( 2000, LM).

56 48 Algal Flora of Korea Freshwater Diatoms VII the outer fissures of the raphe weakly curved, the central ends of the raphe straight, central nodules large and round, terminal ends of the raphe straight or slightly bent to one side. Axial area very narrow, however, central area large rhombic to rectangular reaching the valve margins. Striae rows in 10 μm, radiate in the middle and slightly convergent towards the ends. Valves μm in length and 5 7 μm in breadth. TYPE LOCALITY: Puck Bay, Gulf of Gdansk, Poland, salt meadow. ECOLOGY AND DISTRIBUTION: This taxon is mainly found in mesotrophic habitats with moderately or highly electrolyte water (Krammer 2000). In Korea, this species is observed rarely as epiphytons near Bukcheongnamdaecheon Stream of North Korea in 1997 (Cho 2000c) and in the estuary of the Nakdong River. REMARKS: This species is easily distinguished by the outline of the valve, the narrow axial area and the delicate forms of the striae (Krammer 2000). Fig. 38. Distribution of Pinnularia globiceps var. linearis. 19. Pinnularia graciloides Hustedt 1937: 293 (Figs. 39, 40). Krammer 2000: 127. pl. 99. f. 6. SYNONYM: Pinnularia gracilis Hustedt 1934 in A. Schmidt et al. pl f. 2. Pinnularia franconica Mayer 1946: 131. pl. 6. f. 5. Valves moderately linear in outline with delicate triundulate margins and broadly protracted or subrostrate ends. The central inflation broader than the other two. Raphe lateral, the outer fissures of the raphe curved, the central ends of the raphe expanded and bent in the same direction, central pores rounded, the terminal ends of the raphe large and shaped like bayonet. Axial area 1/4 1/3 of the valve breadth, widening from the ends to the center, and central area large rhombic, sometimes reaching the margins to form fascia. Markings or flecks always present in the hyaline central area. Striae 8 13 rows in 10 μm, strongly radiate in the middle and convergent towards the ends. Longitudinal lines absent across the striae. Valves μm in length and 8 14 μm in breadth. TYPE LOCALITY: Spring, R. Lamongan, Java. ECOLOGY AND DISTRIBUTION: Pinnularia graciloides is widely distributed in low electrolyte

57 Pennales: Naviculaceae: Pinnularia 49 A B C D 10 μm Fig. 39. Pinnularia graciloides. A D. view of valves ( 2000, LM).

58 50 Algal Flora of Korea Freshwater Diatoms VII waters and common in tropical regions (Krammer 2000). It is designated as low ph indicator species in the study for estimating palaeoecological conditions from East Africa (Gasse and Tekaia 1983). In Korea, this species is found rarely from Anjeokneup-C Bog and the third Mujechineup of Mountain Jeongjok in Ulsan, in December 2009 and is the first report from Korea. REMARKS: Further considerations are required to decide whether or not P. substomatophora Hustedt 1934 (Patrick and Reimer 1966: 610. pl. 57. f. 6; Krammer 2000) represents this species. Fig. 40. Distribution of Pinnularia graciloides. 20. Pinnularia graciloides var. triundulata (Fontell) Krammer 2000: 127 (Figs. 41, 42). Krammer 2000: 127. pl. 99. f. 5. BASIONYM: Pinnularia stomatophora f. triundulata Fontell 1917: 38. pl. 1. f. 11. Valves moderately linear in outline with delicate triundulate margins and broadly protracted or subrostrate ends. The middle inflation equal in breadth or the middle inflation a little smaller, but undulations in small specimens nearly absent. Raphe lateral, the outer fissures of the raphe curved, the central ends of the raphe expanded and bent in the same direction, central pores rounded, the terminal ends of the raphe large shaped like bayonet. Axial area 1/4 1/3 of the valve breadth, widening from the ends to the center, and central area large rhombic, usually reaching the margins to form fascia. Markings or flecks always present in the hyaline central area. Striae rows in 10 μm, strongly radiate in the middle and convergent towards the ends. Longitudinal lines absent across the striae. Valves (119) μm in length and (17) μm in breadth. TYPE LOCALITY: Gren, Jämtland, Sweden. ECOLOGY AND DISTRIBUTION: This variety has probably similar ecological characteristics to that of

59 Pennales: Naviculaceae: Pinnularia 51 E 10 μm A B C D Fig. 41. Pinnularia graciloides var. triundulata. A E. view of valves (A: 1250, B: 1500, C E: 2000, LM).

60 52 Algal Flora of Korea Freshwater Diatoms VII the nominal variety. In Korea, this species is found only from Duncheol Bog of Mountain Duncheol in Sancheong, April 2010 and it is the first report from Korea. REMARKS: This variety is characterized by a nearly parallel outline in the margins of the valve. Fig. 42. Distribution of Pinnularia graciloides var. triundulata. 21. Pinnularia grunowii Krammer 2000: 100 (Figs. 43, 44). Hustedt 1930: 317. f. 573b. Patrick and Reimer 1966: 600. pl. 55. f. 17. Krammer and Lange-Bertalot 1986: 424. pl f. 1. Krammer 2000: 100. pl. 77. f. 7. SYNONYM: Pinnularia interrupta sensu Hustedt 1930: 317. f. 573b. Pinnularia mesolepta (Ehrenberg) W. Smith, morphotype 2 sensu Krammer 1992: 119. pl. 44. f. 1. Valves linear in outline, margins triundulate, the inflation of the middle parts equal or narrower than the other two, the ends capitate and distinctly protracted from the body. Raphe straight and filiform, the central ends of the raphe bent laterally, the central pores small, the terminal ends of the raphe shaped like question marks. Axial area narrowly linear and central area large rhombic to rectangular reaching the valve margins. Striae rows in 10 μm, radiate in middle and strongly convergent towards the ends. Valves μm in length and 7 9 μm in breadth. TYPE LOCALITY: Reichenbach, Frankenwald, Bavaria, Germany. ECOLOGY AND DISTRIBUTION: Pinnularia grunowii is widespread in freshwater, in standing or flowing water, and is found both in lowlands and mountainous area (Krammer and Lange-Bertalot 1986). It is most common in low electrolyte water with low calcium concentration in lakes of boreal

61 Pennales: Naviculaceae: Pinnularia μm A B C D E F G H I J K L Fig. 43. Pinnularia grunowii. A L. view of valves ( 2000, LM).

62 54 Algal Flora of Korea Freshwater Diatoms VII forest (Moser et al. 2004). In a study of planktonic and benthic diatoms, Catalan et al. (2009) concluded that this taxon is an indicator in a cluster group from the 235 alpine lakes of the Pyrenees, the Alps, the Tatras and of four other regions. In Korea, this species was recorded as P. interrupta W. Smith from streams (Choi et al. 1993), rivers (Cho 1967), swamps in DMZ area (Cho et al. 1987). It is observed recently from a wetland of Daegwanryeong Pasture in Pyeongchang, May in 2011 and a rice paddy in Wando, July in REMARKS: This species can be separated from the related species, such as P. biceps Gregory, P. angusta (Cleve) Krammer, P. pluvianiformis Krammer, P. mesolepta (Ehrenberg) W. Smith, by the outline, size of the valve and the arrangement of the striae (Krammer 2000). Fig. 44. Distribution of Pinnularia grunowii. 22. Pinnularia karelica Cleve 1891: 28 (Figs. 45, 46). Hustedt 1930: 322. f Krammer and Lange-Bertalot 1986: 422. pl f. 9. Noh 1991: 116. f. 41d. Krammer 2000: 79. pl. 57. f. 1. SYNONYM: Schizonema karelicum (Cleve) Kuntze 1898: 553. Valves moderately broadly linear in outline and margins weakly convex with flatly rounded ends. Raphe weakly lateral, the central ends of the raphe bent laterally, the central pores relatively large round, the terminal ends of the raphe simply bent to the one side. Axial area very narrow and linear, central area large rhombic to rhombic-circular. Striae rows in 10 μm, slightly radiate in middle and moderately convergent towards the ends. Valves μm in length and μm in breadth. TYPE LOCALITY: Tana Elf, Lapland, Finland. ECOLOGY AND DISTRIBUTION: Widespread in northern alpine waters, but rare, and it is mostly found in low to moderate electrolyte concentration (Krammer 2000). In Korea, it was first reported from a swamp in Haman and a tributary of the Namhan River (Noh

63 Pennales: Naviculaceae: Pinnularia 55 E C D 10 μm A B Fig. 45. Pinnularia karelica. A E. view of valves ( 2000, LM).

64 56 Algal Flora of Korea Freshwater Diatoms VII 1991), from planktons collected in the Nakdong River Estuary (Lee et al. 1995) and from epilithic diatom assemblages of the Bukcheongnamdaecheon Stream in North Korea (Cho 2000b). REMARKS: This species can be easily recognized by the broad valve and the delicate form of the striae. Fig. 46. Distribution of Pinnularia karelica. 23. Pinnularia lata (Brébisson) Rabenhorst 1853: 42 (Figs. 47, 48). Krammer and Lange-Bertalot 1986: 403. pl f. 3. Krammer 2000: 20. pl. 3. f. 1. BASIONYM: Frustulia lata Brébisson 1838: 18. SYNONYM: Pinnularia pachyptera Ehrenberg 1843: 421. pl. 4/2. f. 9. Pinnularia hebridensis Gregory 1854: 28. pl. 4. f. 13. Valves broadly linear to linear-elliptical in outline with parallel or weakly convex margins and obtusely rounded ends. Raphe broadly lateral, the outer fissures of the raphe curved, the central ends of the raphe expanded and bent in one direction, the central pores distinctly rounded, and the terminal ends of the raphe shaped like large sickle. Axial area linear and central area large rounded sometimes nearly reaching the valve margins. Central area occasionally asymmetrical in outline. Striae very strongly broad and distant between two striae, rows in 10 μm, radiate in middle and weakly convergent towards the ends. Valves μm in length and μm in breadth. TYPE LOCALITY: Falaise, Calvados, northern France. ECOLOGY AND DISTRIBUTION: Pinnularia lata is widespread in the cool and oligotrophic water of northern alpine regions, and often found in aerophytic diatom assemblages of moss (Krammer

65 Pennales: Naviculaceae: Pinnularia 57 A 10 μm B Fig. 47. Pinnularia lata. A, B. view of valves (Two are photographes of one specimen, but different in modes, DIC mode in A and general mode in B) ( 2000, LM).

66 58 Algal Flora of Korea Freshwater Diatoms VII 2000). Blooms of the species can occur in favorable conditions. In Korea, this species was observed in the Baekrokdam, a crater pond of the Mountain Halla in Jeju Island (Lee 1987) and the Sinbulsanneup B Bog of Mountain in Yangsan. REMARKS: The diagnostic characters of this species are the outline of the valve and the form of the striae, and this is different to P. rabenhorstii (Grunow) Krammer and P. alpina W. Smith by the weakly subrostrate ends and the elliptic-lanceolate outline, respectively (Krammer 2000). Fig. 48. Distribution of Pinnularia lata. 24. Pinnularia macilenta Ehrenberg 1843: 421 (Figs. 49, 50). Hustedt 1930: 331. f Krammer and Lange-Bertalot 1986: 429. pl f. 1. Krammer 2000: 86. pl. 62. f. 1. BASIONYM: Navicula macilenta Ehrenberg 1837: 45. SYNONYM: Navicula oblonga var. macilenta (Ehrenberg) Schumann 1867: 102. Navicula macilenta (Ehrenberg) Pantocsek 1889: 50. Pinnularia macilenta sensu Cleve 1891: 24. pl. 1. f. 7. Pinnularia macilenta sensu Cleve 1895: 88. Schizonema macilentum (Ehrenberg) Kuntze 1898: 553. Navicula oblonga var. macilenta (Ehrenberg) M. Peragallo 1903: 601. Pinnularia macilenta sensu Hustedt 1930: 331. f Pinnularia subgibba var. hustedtii Krammer 1992: 127. f. 47. Pinnularia macilenta sensu Krammer 1992: 156. pl. 66. f. 3. Valves linear in outline, the margins of the valve parallel or weakly convex, rhombic or triundulate, but slightly swollen in the middle and terminal portion, with broadly rounded ends. Raphe broadly lateral, the outer fissures of the raphe slightly curved, the central ends of the raphe

67 Pennales: Naviculaceae: Pinnularia 59 A B C 10 μm D E F Fig. 49. Pinnularia macilenta. A F. view of valves (A E: 1500, F: 2000, LM).

68 60 Algal Flora of Korea Freshwater Diatoms VII slightly expanded and bent in one direction, the terminal ends of the raphe large and shaped like question marks. Central pores large round and closely located each other. Axial area broad, 1/4 1/2 of the valve breadth, and central area irregularly rhombic. Striae 8 10 rows in 10 μm, strongly radiate in the middle and becoming convergent over the ends. Longitudinal bands absent. Valves μm in length and μm in breadth. TYPE LOCALITY: Cayenne, French Guyana. ECOLOGY AND DISTRIBUTION: Pinnularia macilenta is widely distributed from tropical to temperate regions and from oligotrophic to moderately eutrophic water (Krammer 2000). This species has a wide range of ecological characteristics. In Korea, it was collected from three localities by Noh (1991) and was found in a peatland, Yongneup of Mountain Daeam in Inje, May REMARKS: Cleve (1891, 1895) has confused Pinnularia macilenta Ehrenberg 1843 with other taxa, he regarded P. macilenta as a taxon similar to P. neomajor. Other authors followed Cleve (Krammer 2000). The investigations of Reichardt (1995: 10. pl. 11. f pl. 12. f. 15) on Ehrenberg s type material from Cayenne, Guyana Fig. 50. Distribution of Pinnularia macilenta. shows P. macilenta as a taxon identical with the largest morphotype of P. subgibba var. linearis (Krammer 2000). It is also convergent with P. gibba complex. 25. Pinnularia microstauron (Ehrenberg) Cleve 1891: 28 (Figs. 51, 52). Patrick and Reimer 1966: 597. pl. 55. f. 12. Krammer and Lange-Bertalot 1986: 425. pl f. 1. Krammer 2000: 73. pl. 50. f. 1. BASIONYM: Stauroptera microstauron Ehrenberg p. 1/4. f 1. SYNONYM: Stauroptera parva Ehrenberg pl. 3/1. f 19. Stauroneis microstauron (Ehrenberg) Kützing 1844: 106. pl. 29. f. 13 Navicula microstauron (Ehrenberg) O Meara 1875: 354. pl. 30. f. 36. Navicula bicapitata var. hybrid Grunow in Van Heurck 1880: 6. f. 9. Schizonema microstauron (Ehrenberg) Kuntze 1898: 553. Navicula microstauron (Ehrenberg) Héribaud-Joseph 1903: 62. Valves linear-lanceolate to linear-rhombic in outline with slightly convex or concave sides in the center and distinctly protracted ends in large forms and wedge-shaped ends in small forms. Raphe filiform to narrowly lateral, the outer fissures of the raphe weakly curved, the central fis-

69 Pennales: Naviculaceae: Pinnularia 61 C B A 10 μm D E F G H I J K L Fig. 51. Pinnularia microstauron. A L. view of valves ( 2000, LM).

70 62 Algal Flora of Korea Freshwater Diatoms VII sures of the raphe slightly expanded and bent in one direction, the central pores large tear-drop shaped, the terminal ends of the raphe small and indistinct in shape under light microscopy. Axial area relatively linear and central area rectangular up to the margins. Striae 9 14 rows in 10 μm on valves, slightly radiate at centre becoming convergent towards the ends. Striae interrupted at the center to form a broad hyaline fascia. Valves μm in length and 7 15 μm in breadth. TYPE LOCALITY: Rio de Janeiro in Brazil, coating plant roots. ECOLOGY AND DISTRIBUTION: Pinnularia microstauron is cosmopolitan over various types of water bodies and is most common in cold waters and low electrolyte conditions (Krammer 2000). In the study for plankton and benthic diatoms, this taxon and the following varieties are the indicative in cluster groups from the 235 alpine lakes of the Pyrenees, the Alps, the Tatras and four other regions (Catalan et al. 2009). It is the most common and dominant in the epilithic diatoms from rocks in the alpine stream and lakes of the Swiss Alps (Robinson and Kawecka 2005), and the most widespread in Sphagnum bogs (Kulikovskiy 2009a, Fig. 52. Distribution of Pinnularia microstauron. 2009b). This is designated as low ph indicator species in the study for estimating palaeoecological conditions from East Africa (Gasse and Tekaia 1983), and is considered to be moderately tolerant to heavy organic pollution and eutrophication using the epilithic diatoms in the Gravataí River, Brazil (Salomoni et al 2006). In Korea, this species is the most frequently occurring taxon in the freshwater of Korea. It was observed as periphyton in streams (Chung and Watanabe 1984; Noh 1991; Chung et al. 1992) and swamps (Cho et al. 1987; Chung and Noh 1987; Chung and Kim 1987; Noh 1991), as plankton and periphyton in rivers (Noh 1991; Choi and Kim 1994) and reservoirs and lakes (Lee 1985; Noh 1991). But P. microstauron described by Noh (1991) has morphologically varied forms of valves. REMARKS: Some 72 forms or varieties of P. microstauron are listed in the AlgaeBase database (Guiry and Guiry 2011), but most of these are typological forms within their variability or belong to other taxa (Krammer 2000). 26. Pinnularia microstauron var. nonfasciata Krammer 2000: 74 (Figs. 53, 54). Krammer 2000: 74. pl. 52. f. 1. Valves linear-lanceolate to linear-rhombic in outline with slightly convex or concave sides in the

71 Pennales: Naviculaceae: Pinnularia 63 A B C D E F 10 μm G H I J K Fig. 53. Pinnularia microstauron var. nonfasciata. A K. view of valves ( 2000, LM).

72 64 Algal Flora of Korea Freshwater Diatoms VII center and distinctly protracted ends in large forms and wedge-shaped ends in small forms. Raphe filiform to narrowly lateral, the outer fissures of the raphe weakly curved, the central fissures of the raphe slightly expanded and bent in one direction, the central pores large tear-drop shaped, the terminal ends of the raphe small and indistinct in shape under light microscopy. Axial area relatively linear and central area always rounded to rhomboid. Striae rows in 10 μm on valves, slightly radiate at centre becoming convergent towards the ends. Striae interrupted at the center to occasionally form a hyaline fascia. Valves μm in length and 8 11 μm in breadth. TYPE LOCALITY: Frankenwald, Reichenbach on the Spitzberg in Germany. ECOLOGY AND DISTRIBUTION: This variety is probably most common in cold and low electrolyte water like the nominal variety. In Korea, Pinnularia microstauron var. nonfaciata was frequently or dominantly found from a mountain bog, Sumeunmulbaengdwineup, in Mountain Halla of Jeju Island. This is the first report in Korea. REMARKS: This variety can be differentiated from nominal variety in a distinctly rhomboid central area (Krammer 2000). Fig. 54. Distribution of Pinnularia microstauron var. nonfasciata. 27. Pinnularia neomajor var. inflata Krammer 2000: 166 (Figs. 55, 56). Krammer 2000: 166. pl f. 1. Valves linear in outline, slightly swollen in the middle and at the rounded ends. Raphe lateral, the outer fissures of the raphe moderately undulate, the central ends of the raphe expanded and bent in one direction, the central pores round and close together, the terminal ends of the raphe shaped like question marks. Axial area 1/4 1/3 of the valve breadth, broadly linear in outline and tapering towards the ends. Central area a little wide than axial area and usually asymmetrical. Striae 6 7 rows in 10 μm, radiate in the middle and weakly convergent in the ends. Longitudinal lines distinct across the striae. Valves (322) μm in length and (35) μm in breadth. TYPE LOCALITY: Lake Grand in Canada. ECOLOGY AND DISTRIBUTION: Pinnularia neomajor var. inflata is most often found in oligotrophic to dystrophic waters with low electrolyte (Krammer 2000). This variety is abundant on sediments of large lakes in Canada and probably common in the freshwaters like the nominal vari-

73 Pennales: Naviculaceae: Pinnularia 65 A B C D 10 μm E Fig. 55. Pinnularia neomajor var. inflata. A E. view of valves (A: 600, B: 800, C: 1000, D, E: 2000, LM).

74 66 Algal Flora of Korea Freshwater Diatoms VII ety, Pinnularia neomajor var. neomajor. In Korea, it was found in epiphytons collected in the Seonakdong River, but its abundance was very low. This is the first report in Korea. REMARKS: Pinnularia major Rabenhorst 1853 is different from P. neomajor Krammer 2000 and its synonym P. major sensu Cleve 1895 by outline, size and axial area (Krammer 2000). P. neomajor var. inflata is different from P. gentilis by the raphe structure, the former has lateral or semicomplex, and wide raphe, but the latter has complex and some crossed raphe. Fig. 56. Distribution of Pinnualria neomajor var. inflata. 28. Pinnularia nobilis var. regularis Krammer 2000: 184 (Figs. 57, 58). Krammer 2000: 184. pl f. 1. Valves linear in outline, the margins of the valve nearly parallel, but weakly swollen in the middle and at the ends, the two inflations nearly equal in the breadth, the ends broadly rounded. Raphe broadly lateral, semicomplex to complex, the central ends of the raphe slightly curved in one direction and central pores large and round, the terminal ends of the raphe shaped like question marks. Axial area linear occupying 1/3 of the breadth of the valve and narrowing towards the ends, central area roundish to somewhat variable in size and asymmetrical in outline. Striae 4 5 rows in 10 μm, strongly radiate in the middle and convergent towards the ends, longitudinal bands distinctly broad. Valves μm in length, μm in breadth. TYPE LOCALITY: Isigny in France. ECOLOGY AND DISTRIBUTION: This variety is abundant in oligotrophic, circumneutral and low electrolyte water, but not common (Krammer 2000). In Korea, the forms of Pinnularia nobilis var. regularis were already reported under the name of P. nobilis (Ehrenbeg) Ehrenberg from benthic diatoms in streams and lowland swamps (Noh 1991). It was recently reported from mountain bogs, Sohwangbyeongsanneup of Mountain Odae in Pyeong-

75 Pennales: Naviculaceae: Pinnularia 67 A B C D 10 μm Fig. 57. Pinnularia nobilis var. regularis. A D. view of valves (A, B: 600, C: 750, D: 1500, LM).

76 68 Algal Flora of Korea Freshwater Diatoms VII chang and commonly from an unnamed wetland in Daegwanryeong Pasture in Pyeongchang, May in REMARKS: Pinnularia nobilis is very common in the diatom literatures (Krammer 2000), three related taxa, P. nobilis var. regularis Krammer, P. gigas Ehrenberg (= P. dactylus Ehrenberg) and P. flexuosa Cleve, do not show any conspicuous differences, but can be differentiated on the basis of the valve outline, raphe complexity and axial area. P. flexuosa has the parallel margins of the valve and wider axial area, P. gigas has slightly swollen margins in the middle and lateral raphe, and P. nobilis var. regularis has equally swollen margins in the middle and ends of the valve. Fig. 58. Distribution of Pinnularia nobilis var. regularis. 29. Pinnularia obscura Krasske 1932: 117 (Figs. 59, 60). Krammer and Lange-Bertalot 1986: 420. pl f. 20. Krammer 2000: 50. pl. 13. f. 10. SYNONYM: Navicula obscura (Krasske) F.W. Mills 1935: Valves linear to linear-elliptical in outline with straight to weakly concave or convex margins and wedge-shaped or slightly rostrate ends. Raphe filiform to weakly lateral, the central ends of the raphe slightly bent in one direction and the central pore small, the terminal ends of the raphe shaped like question marks, but difficult to observe under light microscopy. Axial area linearly narrow and hyaline central area widening up to the valve margins. Striae rows in 10 μm, moderately radiate in the middle becoming strongly convergent towards the ends. Valves μm in length and μm in breadth. TYPE LOCALITY: Alps, In Moos eines etwas feuchten Hanges bei Kampenn, Südtirol. ECOLOGY AND DISTRIBUTION: Pinnularia obscura is widely distributed in northern alpine regions, particularly aerophytic to wet mosses and rocks (Krammer and Lange-Bertalot 1986). In an ecological threshold study for plankton and benthic diatoms, this species is the second most important indicator in cluster groups from the 235 alpine lakes of the Pyrenees, the Alps, the Tatras and four other regions (Catalan et al. 2009). It was designated as low ph indicator species in the study for

77 Pennales: Naviculaceae: Pinnularia 69 A B C D E F G H I J K 10 μm L M Fig. 59. Pinnularia obscura. A M. view of valves ( 2000, LM). estimating palaeoecological conditions from East Africa (Gasse and Tekaia 1983). In Korea, this species was found from two peatland bogs of Mountain Odae in Pyeongchang and Daeam in Inje, May in Noh (1991) first reported this species from some Korean freshwaters, but the forms of her specimens (Fig. 35f h) are different to this species. REMARKS: This species is similar to P. schoenfelderi Krammer, but striae of the central parts moderately radiate becoming more or less convergent towards the ends; it is similar to P. intermedia (Lagerstedt) Cleve, but the striae broader and more distant in the present species. Fig. 60. Distribution of Pinnularia obscura.

78 70 Algal Flora of Korea Freshwater Diatoms VII 30. Pinnularia peracuminata Krammer 2000: 157 (Figs. 61, 62). Krammer 2000: 157. pl f. 1. REPLACED SYNONYM: Pinnularia rupestris var. cuneata Krammer 1992b: 142. pl. 56. f. 1. SYNONYM: Pinnularia acuminata sensu Rabenhorst 1861: Valves linear in outline, the margins of the valve parallel and slightly swollen in the middle in larger forms, with cuneate and acutely rounded ends. Raphe lateral, the central ends of the raphe slightly bent to one side, the central pores drop-shaped, the terminal ends of the raphe shaped like question marks. Axial area linear occupying 1/4 1/3 of the valve breadth, central area rhomboid. Striae rows in 10 μm, slightly radiate in middle and weakly convergent towards the ends. Longitudinal lines present, however, sometimes indistinctly. Valves μm in length and 9 10 μm in breadth. TYPE LOCALITY: Walthers Grund in the Ober-Lössnitz near Dresden, Germany. ECOLOGY AND DISTRIBUTION: Pinnularia peracuminata was recorded in the streams of mountains and aerial diatom assemblages (Krammer 2000). In Korea, this species was first found in a peatland of Mountain Daeam in Inje, Yongneup Bog, May in REMARKS: This species is close to P. esoxiformis Fusey in morphology, but the latter has coarser striae. Fig. 61. Distribution of Pinnularia peracuminata.

79 Pennales: Naviculaceae: Pinnularia 71 A B C D E F G 10 μm H I J K L M Fig. 62. Pinnularia peracuminata. A M. view of valves ( 2000, LM).

80 72 Algal Flora of Korea Freshwater Diatoms VII 31. Pinnularia pluvianiformis Krammer 2000: 103 (Figs. 63, 64). Krammer 2000: 103. pl. 83. f. 1. SYNONYM: Pinnularia pluviana Sovereign sensu Lange- Bertalot and Metzeltin pl. 51. f. 10. Valves linear in outline, the margins of the valve triundulate, the central swelling equally wide with the other two, the ends of the valve subcapitate. Raphe straight, the central ends of the raphe slightly bent to one side, the central pores small round, the terminal ends of the raphe shaped like question marks. Axial area narrow, central area rhombic widening up to the margins of the valve. Striae rows in 10 μm, slightly radiate in middle and convergent towards the ends. Valves μm in length and 6 7 μm in breadth. TYPE LOCALITY: Julma Ölkky near Kuusamo in Finland, on rocks. ECOLOGY AND DISTRIBUTION: Until now, Pinnularia pluvianiformis is reported only from the type locality (Krammer 2000). In Korea, this species was found in a peatland, the Sumeunmulbaengdwineup Bog, in Mountain Halla of Jeju Island, August in This is the first report in Korea. Fig. 63. Distribution of Pinnularia pluvianiformis. 10 μm A B C D E F G H Fig. 64. Pinnularia pluvianiformis. A H. view of valves ( 2000, LM).

81 Pennales: Naviculaceae: Pinnularia 73 REMARKS: This species is distinguished by the outline ratio on the length and breadth of the valve and the finer striae (Krammer 2000) and resembles to a small form of Pinnularia grunowii. 32. Pinnularia pseudogibba Krammer 1992b: 129 (Figs. 65, 66). Krammer 2000: 89. pl. 48. f. 8. Valves linear in outline, the margins of the valve weakly convex or delicately undulate with broadly truncate or rounded ends. Raphe broadly lateral, the central ends of the raphe slightly bent to one side, the central pores round to drop-shaped and closely adjacent, the terminal ends of the raphe shaped like question marks. Axial area linear occupying 1/4 1/3 of the valve breadth, central area rectangular developed up to the margins of the valve. Striae rows in 10 μm, radiate in middle and weakly convergent towards the ends. A longitudinal line present, however, sometimes indistinctly. Valves μm in length and 9 10 μm in breadth. TYPE LOCALITY: Soorbach, Hertogenwald, near Eupen in Belgium. ECOLOGY AND DISTRIBUTION: Pinnularia pseudogibba is abundant in oligotrophic, cold and low mineral streams in mountainous area (Krammer 2000). In Korea, this species was found in two peatlands of Mountain Odae in Pyeongchang and Daeam in Inje, May in This is the first report in Korea. REMARKS: This species is greatly different from P. microstauron in its habitat (Krammer 2000). Fig. 65. Distribution of Pinnularia pseudogibba.

82 74 Algal Flora of Korea Freshwater Diatoms VII G C D E F 10 μm B A Fig. 66. Pinnularia pseudogibba. A G. view of valves ( 2000, LM).

83 Pennales: Naviculaceae: Pinnularia Pinnularia pulchra Østrup 1897: 263 (Figs. 67, 68). Krammer and Lange-Bertalot 1986: 414. pl f. 9, 10. Krammer 1992b: 123. pl. 49. f. 1. Krammer 2000: 119. pl. 90. f. 1. SYNONYM: Navicula pulchra (Østrup) M. Peragallo : 269. pl. 1. f. 3. Valves linear in outline, the margins of the valve weakly triundulate or biundulate in small forms. The breadth of three inflations identical or the middle inflation the most narrow. The ends of the valve broadly protracted and rostrate. Raphe slightly lateral, the outer fissures of the raphe weakly curved, the central ends of the raphe slightly bent to the one side, the central pores distinct and the terminal ends of the raphe semicircular. Axial area narrow linear and central area round to rectangular reaching the margins of the valve. Striae 9 11 rows in 10 μm on valves, slightly radiate in the middle becoming weakly convergent Fig. 67. Distribution of Pinnularia pulchra. 10 μm A B C Fig. 68. Pinnularia pulchra. A C. view of valves ( 2000, LM).

84 76 Algal Flora of Korea Freshwater Diatoms VII towards the ends. Valves μm in length and 6 11 μm in breadth. TYPE LOCALITY: East Greenland. ECOLOGY AND DISTRIBUTION: Pinnularia pulchra is found in the northern and subarctic regions as epipelic diatoms. It is most common in oligotrophic, dystrophic and low electrolyte streams and ponds, but its frequency is low (Krammer 2000). However, this species was found in the coastal wetland in Albania (Miho and Witkowski 2005). In Korea, this species was first found from two streams of the Namhan River by Noh (1991) and was collected from an unnamed wetland in Daegwanryeong Pasture, Pyeongchang, May in REMARKS: Pinnularia pulchra is closely related with P. angusta (Cleve) Krammer and P. infirma Krammer, but is distinguished from these by the size and outline of the valve, the broad protracted ends (Krammer 2000). 34. Pinnularia quadratarea (A. Schmidt) Cleve 1895: 95 (Figs. 69, 70). Hendey 1964: 232. Witkowski et al. 2000: 335. pl f. 17. BASIONYM: Navicula quadratarea A. Schmidt 1874: 90. pl. 2. f. 26. Valves linear-oblong in outline, the margins of the valve nearly parallel and the ends of the valve with broadly rounded. Raphe straight, the central ends of the raphe slightly expanded and bent in one direction, the terminal ends bent to one direction. Axial area very narrow and central area rectangular and large developed up to the margins. Striae 8 10 rows in 10 μm, parallel throughout the valve and slightly convergent towards the ends. Valves μm in length and 8 12 μm in breadth. TYPE LOCALITY: North See, Europe. No other locality given. ECOLOGY AND DISTRIBUTION: As a marine taxon, Pinnularia quadratarea is widely distributed in European coasts and entire Baltic Sea and is most commonest in the colder water (Witkowski et al. 2000). It was dominant in the phytoplankton and sea ice algal communities at the end of winter in McMurdo Sound, the Antarctic (McMinn et al. 2010). In Korea, this species was found from sand flats in intertidal area of the Nakdong River Estuary. This is the first report in Korea. REMARKS: This species is distinguished in the largely broad central area and the finer striae. Fig. 69. Distribution of Pinnularia quadratarea.

85 Pennales: Naviculaceae: Pinnularia 77 B C 10 μm A Fig. 70. Pinnularia quadratarea. A C. view of valves ( 2000, LM). 35. Pinnularia rabenhorstii var. franconica Krammer 2000: 22 (Figs. 71, 72). Krammer 2000: 22. pl. 5. f. 5. SYNONYM: Pinnularia borealis var. thuringiaca sensu Krammer 1992b. pl. 10. f. 5. Valves linear, parallel to moderately convex or concave in outline with somewhat broadly subrostrate. Raphe weakly to moderately lateral and broad, the outer fissures of the raphe curved, the terminal ends of the raphe shaped like large sickles, the central ends of the raphe slightly expanded and bent in one direction, the central pores distinctly rounded. Axial area Fig. 71. Distribution of Pinnularia rabenhorstii var. franconica.

86 78 Algal Flora of Korea Freshwater Diatoms VII E D A B C 10 μm Fig. 72. Pinnularia rabenhorstii var. franconica. A E. view of valves ( 2000, LM). narrow and central area roundly developed. Striae coarsely broad and distant, 4 5 rows in 10 μm, slightly radiate in middle and weakly convergent towards the ends. Valves μm in length and μm in breadth. TYPE LOCALITY: Rivulet near Reichenbach, Frankenwald in Germany. ECOLOGY AND DISTRIBUTION: This variety is distributed in oligotrophic water of lower mountains and as epipelic forms in some places in Bavaria in Germany (Krammer 2000). In Korea, this species was found in peatlands, the third Mujechineup Bog of Mountain Jeongjok in Ulsan and Yongneup Bog of Mountain Daeam in Inje. This is the first report in Korea. REMARKS: This variety is smaller than the nominal variety. Pinnularia rabenhorstii overlaps P. borealis in morphology, but they can be differentiated by the size. The largest specimens of P. borealis and its varieties rarely exceed the valve breadth of 12 μm.

87 Pennales: Naviculaceae: Pinnularia Pinnularia rhombarea Krammer in Metzeltin and Lange-Bertalot 1998: 185 (Figs. 73, 74). Krammer 2000: 75. pl. 53. f. 1. SYNONYM: Pinnularia microstauron (Ehrenberg) Cleve sensu morphotype 3 in Krammer 1992b: 100. pl. 33. f. 14. Valves broadly linear in outline with parallel to weakly convex margins and broadly protracted or subrostrate ends. Raphe narrowly lateral, the central ends of the raphe slightly expanded and bent in one direction and the terminal ends of the raphe shaped like question marks, difficult to see under light microscopy. Axial area narrowly linear and central area large rhombic, reaching the valve margins. Striae 9 10 rows in 10 μm on valves, radiate in the middle becoming convergent over the ends. Valves μm in length and μm in breadth. TYPE LOCALITY: Hammerfest in North Norway, on moss in a pool. ECOLOGY AND DISTRIBUTION: This taxon is most common in oligotrophic and low electrolyte waters in the northern and subarctic region rather than in the central Europe (Krammer 2000). The part of the fen that is characterized by Sphagnum species is dominated by acidobiontic and acidophilous species, some Eunotia species and Pinnularia rhombarea (Kapetanović et al. 2011). In Korea, this species was found in some peatlands, the Sohwangbyeongsanneup Bog of Mountain Odae in Pyeongchang and Yongneup Bog of Mountain Daeam in Inje, May REMARKS: This species is distinguished from the P. microstauron complex by broadly hatchet-like rostrate and flatly rounded ends. Fig. 73. Distribution of Pinnularia rhombarea.

88 80 Algal Flora of Korea Freshwater Diatoms VII 10 μm A B C D E F G H I Fig. 74. Pinnularia rhombarea. A I. view of valves (A C: 1500, D I: 2000, LM).

89 Pennales: Naviculaceae: Pinnularia Pinnularia schoenfelderi Krammer 1992b: 70 (Figs. 75, 76). Hustedt 1930: 322. f Krammer 2000: 40. pl. 10. f. 18. SYNONYM: Pinnularia microstauron var. brebissonii f. diminuta sensu Hustedt 1930: 322. f (non Navicula brebissonii var. diminuta Grunow in Van Heurck pl. 5. f. 8) Valves linear-lanceolate to linear-elliptical, gradually narrow towards the ends, with obtusely rounded. Raphe narrowly lateral, the central ends of the raphe slightly expanded and bent in one direction and the terminal ends of the raphe shaped like question marks, difficult to see under light microscopy. Axial area widening gradually from point near the base of the polar raphe fissure fork to the central area, which may reach the valve margins. Hyaline central area rectangular, reaching the margins. Striae often coarse, interrupted at the center to form a broad hyaline fascia. Striae rows in 10 μm on valves, radiate in the middle becoming convergent over the ends. Valves μm in length and 5 7 μm in breadth. TYPE LOCALITY: Drainage ditch on the path from the Teichmühle to Reichenbach, Frankenwald of Bavaria in Germany. ECOLOGY AND DISTRIBUTION: Pinnularia schoenfelderi is probably cosmopolitan in oligotrophic and low to moderate electrolyte water (Krammer 2000). In Korea, this species was found from streams in mountains (Chung and Lee 1982; Chung et al. 1985), in river (Lee et al. 1995) and in mountainous peatland bogs. REMARKS: This species can be confused with some other taxa, notably P. microstauron var. brebissonii f. diminuta sensu Hustedt, P. obscura Krasske and P. frauenbergiana Reichardt. These taxa are discriminated by the skewness and density of the striae, and other features. Fig. 75. Distribution of Pinnularia schoenfelderi.

90 82 Algal Flora of Korea Freshwater Diatoms VII A B C D E F G H I J 10 μm K L M N O P Q R S Fig. 76. Pinnularia schoenfelderi. A S. view of valves ( 2000, LM). 38. Pinnularia similiformis var. koreana Metzeltin et Krammer in Krammer 2000: 48 (Figs. 77, 78) Krammer 2000: 48. pl. 16. f. 3. Valves linear-lanceolate to rhombic-lanceolate with obtusely rounded ends. Raphe narrowly lateral, the outer fissures of the raphe weakly undulate, the central ends of the raphe slightly bent, the central pores distinct, the terminal ends of the raphe semicircular. Axial area linear in small valves and lanceolate in large valves, hyaline central area rectangular reaching the valve margins.

91 Pennales: Naviculaceae: Pinnularia 83 A B 10 μm C D F E Fig. 77. Pinnularia similiformis var. koreana. A F. view of valves ( 2000, LM). Striae rows in 10 μm, strongly radiate in the middle and becoming convergent over the ends. Valves μm in length and about 8 μm in breadth. Frustules very broad in the girdle view. TYPE LOCALITY: Korea, photographe from Metzeltin. ECOLOGY AND DISTRIBUTION: This variety is widely distributed in oligotrophic, low electrolyte

92 84 Algal Flora of Korea Freshwater Diatoms VII moorland, and particularly in aerophytic mosses in northern Europe, particularly the Alps and the northern Germany (Krammer 2000). Type locality is designated as Korea in the first description of the variety (Krammer 2000), however, the precise locality is not given. In Korea, this variety occurs in three mountain peatlands, Sinbulsanneup in Yangsan, Yongneup in Daeam Mountain in Inje and 1100 of Halla Mountain in Jeju Island. REMARKS: This variety is characterized by the skewness of sriae and the girdle view (Krammer 2000). Fig. 78. Distribution of Pinnularia similiformis var. koreana. 39. Pinnularia sinistra Krammer 1992b: 105 (Figs. 79, 80). Krammer 2000: 118. pl. 90. f. 6. SYNONYM: Pinnularia subcapitata auct. nonull. Valves linear in outline with slightly convex margins, ends broadly protracted and rostrate. Raphe filiform in smaller forms and slightly lateral in larger forms, the central ends of the raphe slightly expanded and bent in one direction, the terminal ends of the raphe shaped like question marks. Axial area narrow, linear or slightly lanceolate and central area rhombic widening towards the margins. Striae rows in 10 μm, slightly radiate in the middle and becoming convergent over the ends. Valves μm in length and 4 6 μm in breadth. TYPE LOCALITY: Rivulet on the path from the Techmühle to Reichenbach, Frankenwald in Germany. ECOLOGY AND DISTRIBUTION: Pinnularia sinistra is mostly found in oligotrophic, acidic and low electrolyte freshwaters habitat, and may be locally abundant in these area (Krammer 2000). It is the most common and dominant in the epilithic diatoms from rocks in the alpine streams and lakes of the Swiss Alps (Robinson and Kawecka 2005) and Adirondack Park in northern New York (Camburn and Charles 2000).

93 Pennales: Naviculaceae: Pinnularia 85 G H I J K A B C D E F 10 μm Fig. 79. Pinnularia sinistra. A K. view of valves ( 2000, LM). In Korea, this species was reported as P. subcapitata Gregory from periphytons of streams (Hong and Chung 1990; Noh 1991), swamps (Noh 1991), mountain peatlands (Chung and Kim 1987) and lakes (Noh 1991; Choi et al. 1994). It is frequently found from some peatlands of mountains. REMARKS: Pinnularia sinistra shows a mixture of various forms such as Patrick and Reimer pl. 55. f (Krammer 2000). Fig. 80. Distribution of Pinnularia sinistra.

94 86 Algal Flora of Korea Freshwater Diatoms VII 40. Pinnularia stomatophora var. salina Krammer 2000 (Figs. 81, 82). Krammer 2000: 126. pl. 99. f. 7. SYNONYM: Pinnularia salina (Krammer) Kulikovskiy, Lange-Bertalot et Metzeltin 2010: 365. Valves linear in outline, occasionally with slightly convex margins in larger specimen, ends not protracted, obtusely to cuneate rounded. Raphe slightly lateral, the outer fissure of the raphe slightly curved, the central ends of the raphe bent in one direction and the central pores small round, the terminal ends of the raphe shaped like very long bayonets. Axial area moderately broad and linear to linear-lanceolate, 1/4 1/3 the breadth of the valve, and central area elongate-elliptical to rhombic and broad fascia, rather narrow hyaline area reaching up to the margins and markings absent in the central area. Striae rows in 10 μm, slightly radiate in the middle and becoming convergent over the ends. Valves μm in length and μm in breadth. TYPE LOCALITY: Lake Darscho, Neusiedlersee in Austria. ECOLOGY AND DISTRIBUTION: This variety was only reported from the type locality. In Korea, this taxon was found in many peatlands, Mujechineup and Anjeokneup C near Mountain Jeongjok in Ulsan and Sohwangbyeongsanneup of Odae Mountain in Pyeongchang, but never common. This is the first report in Korea. REMARKS: This variety is characterized with the long terminal fissures of the raphe. Fig. 81. Distribution of Pinnularia stomatophora var. salina.

95 Pennales: Naviculaceae: Pinnularia 87 A B C 10 μm D E F G Fig. 82. Pinnularia stomatophora var. salina. A G. view of valves ( 2000, LM).

96 88 Algal Flora of Korea Freshwater Diatoms VII 41. Pinnularia subbrevistriata Krammer 2000: 94 (Figs. 83, 84). Hustedt 1930: 327. f Krammer 2000: 94. pl. 70. f. 7. REPLACED SYNONYM: Navicula brevistriata Grunow in VanHeurck pl. 6. fig. 5. SYNONYM: Pinnularia gibba var. parva sensu Hustedt 1930: 327. f Pinnularia gibba in Metzeltin and Lange-Bertalot pl f. 10. Valves lanceolate in outline, the margins of the valve convex with broadly rostrate ends. Raphe lateral and the outer fissures of the raphe weakly undulate, the central ends of the raphe bent in one direction, the central pores small round and close each other, the terminal ends of the raphe shaped like large question marks. Axial area, very wide and lanceolate, 1/2 2/3 of the valve breadth, connecting with the central area. Striae 8 10 rows in 10 μm, radiate in the middle and moderately convergent over the ends. Valves μm in length and μm in breadth. TYPE LOCALITY: Julma Ölkky near Kuusamo in Finland, on rocks. ECOLOGY AND DISTRIBUTION: Pinnularia subbrevistriata is widespread in oligotrophic and low electrolyte freshwater, but restricted in distribution (Krammer 2000). Additionally, it is frequent in tropical regions. In Korea, this species was reported under the name of Pinnularia gibba var. parva (Ehrenberg) Hustedt from Gyeongju (Chung and Watanabe 1984) and swamps in Haman (Chung and Noh 1987). It is rarely found from mountain peatlands of Daegwanryeong Pasture in Pyeongchang. REMARKS: Pinnularia subbrevistriata is similar to some taxa, such as P. gibba Ehrenberg sensu Krammer 2000, P. parvulissima Krammer 2000 and P. braunii var. amphicephala (Mayer) Hustedt 1930, but may be distinguished by the capitate ends or irregular markings in the central area. Further study is necessary of P. gibba var. parva sensu Hustedt (Hustedt 1930: 327. f. 603) in spite of their morphological similarity. Fig. 83. Distribution of Pinnularia subbrevistriata.

97 Pennales: Naviculaceae: Pinnularia 89 D E F B C 10 μm A Fig. 84. Pinnularia subbrevistriata. A F. view of valves ( 2000, LM). 42. Pinnularia subcapitata var. subrostrata Krammer 1992b: 108 (Figs. 85, 86). Krammer 2000: 118. pl. 90. f. 18. Valves linear to linear-lanceolate in outline with delicately subrostrate ends. Raphe filiform in small forms and narrowly lateral in large forms, the central ends of the raphe slightly bent to the one side and the central pores distinct, the terminal ends of the raphe semicircular. Axial area narrowly linear and central area round to rectangular reaching the margins. Striae rows in 10 μm on valves, slightly radiate in the middle becoming convergent towards the ends. Valves μm in length and 5 7 μm in breadth. TYPE LOCALITY: Pond near Füssen, Algäu, Bavaria, Germany, in moss. ECOLOGY AND DISTRIBUTION: This variety is distributed in northern and alpine area, found not

98 90 Algal Flora of Korea Freshwater Diatoms VII D B C E F A 10 μm G H I J K L M N O P Fig. 85. Pinnularia subcapitata var. subrostrata. A P. view of valves ( 2000, LM).

99 Pennales: Naviculaceae: Pinnularia 91 uncommonly in the oligotrophic and low electrolyte waters (Krammer 2000). In Korea, this species is found with significant frequency in the third Mujechineup of Mountain Jeongjok in Ulsan, some peatlands near the mountain and the Wangdeungjaeneup of Mountain Jiri in Sancheong. This is the first report in Korea. REMARKS: Certain size classes of P. subcapitata and its variety show a degree of convergence with P. sinistra (Krammer 2000). This variety differs from the nominal variety by subrostrate to capitate ends. Fig. 86. Distribution of Pinnularia subcapitata var. subrostrata. 43. Pinnularia subgibba var. sublinearis Krammer 1992b: 85 (Figs. 87, 88). Hustedt 1930: 327. f Krammer 2000: 85. pl. 65. f. 14. SYNONYM: (?) Pinnularia gibba var. linearis sensu Hustedt 1930: 327. f. 604 (non lectotype sensu Simonsen pl f. 1) Valves linear in outline, the margins of the valve parallel and the ends of the valve rounded. Raphe broadly lateral, the outer fissures of the raphe slightly curved, the central ends of the raphe expanded and bent in one direction, the central pores distinctly small round and close together, the terminal ends of the raphe large shaped like question marks. Axial area 1/3 of the valve breadth, widening from the ends to the center, and central area large rhombic reaching the margins to form fascia. Large markings sometimes apparent in the hyaline central area. Striae very strongly broad and distant between two striae, 8 11 rows in 10 μm, radiate in the middle and weakly convergent in the ends. Longitudinal lines absent across the striae. Valves (117) μm in length and (13.5) μm in breadth.

100 92 Algal Flora of Korea Freshwater Diatoms VII A B C F 10 μm D E Fig. 87. Pinnularia subgibba var. sublinearis. A F. view of valves ( 2000, LM).

101 Pennales: Naviculaceae: Pinnularia 93 TYPE LOCALITY: Waltham, Massachusetts, USA. ECOLOGY AND DISTRIBUTION: Pinnularia subgibba var. sublinearis is rare, and generally sporadic in boreal regions, although it is rather frequent in fossil samples (Krammer 2000). This variety is recorded in acid bogs of the English Lake District in England (Lund 1950; Round 1957) and a temporary pond from the Patagonian Andes in Argentina (Villanueva 2006). It is found in classical materials, Degernä diatomite, collected by Ehrenberg from Sweden (Mayama and Kobayasi 1990). In Korea, this species is found in two mountain peatlands, the Duncheolneup in Sancheong and an unnamed peatland of the Daegwanryeong Pasture in Pyeongchang. This is the first report in Korea. REMARKS: This variety is closely similar to P. macilenta except for the forms of valve ends. Fig. 88. Distribution of Pinnularia subgibba var. sublinearis. 44. Pinnularia subgibba var. undulata Krammer 1992b: 127 (Figs. 89, 90). Krammer 2000: 85. pl. 64. f. 4. Valves linear in outline, the margins parallel, weakly convex, rhombic or triundulate, but slightly swollen in the central and terminal portion, with broadly rounded to cuneate ends. Raphe broadly lateral, the outer fissures of the raphe slightly curved, the central ends of the raphe expanded and bent in one direction, the central pores distinctly small round and close together, the terminal ends of the raphe large shaped like question marks. Axial area broad, 1/4 1/2 of the valve breadth, and central area irregularly rhombic. Striae 9 10 rows in 10 μm, strongly radiate in the middle and becoming convergent over the ends. Valves μm in length and 8 10 μm in breadth. TYPE LOCALITY: Ditch in the forest near the Arnold-Denkmal, Erlangen, Bavaria in Germany. ECOLOGY AND DISTRIBUTION: This variety is probably widespread in the northern and arctic regions like the nominal variety. In Korea, this species is found from an unnamed peatland of the Daegwanryeong Pasture in Pyeongchang, May 2011 and some other peatlands. This is the first

102 94 Algal Flora of Korea Freshwater Diatoms VII 10 μm A B C D E F G Fig. 89. Pinnularia subgibba var. undulata. A G. view of valves ( 2000, LM).

103 Pennales: Naviculaceae: Pinnularia 95 report in Korea. REMARKS: This species is closely similar to P. gibba var. subundulata Mayer, illustrated in Hustedt (1930. p f. 601) and further study is required (Krammer 2000). Fig. 90. Distribution of Pinnularia subgibba var. undulata. 45. Pinnularia subgigas Krammer 2000: 180 (Figs. 91, 92). Krammer 2000: 180. pl f. 2. SYNONYM: Pinnularia flexuosa Cleve sensu Krammer 1992b. pl. 73. f. 1. Valves very broad and linear in outline, the margins of the valve parallel or very weakly convex with cuneately rounded ends. Raphe broadly lateral and semicomplex to complex, the central ends of the raphe laterally bent in one direction, the central pores round, the terminal ends of the raphe shaped like question marks. Axial area linear narrowing towards the ends, occupying 1/5 1/3, and central area slightly round, asymmetrically widening. Striae about 5 in 10 μm, radiate in the middle and becoming slightly convergent or parallel over the ends, and crossed by a broad longitudinal band. Valves μm in length and μm in breadth. TYPE LOCALITY: Grand Lake in Canada. ECOLOGY AND DISTRIBUTION: Pinnularia subgigas is common in some of the larger lakes of North America, and frequently occurs in the fossil records (Krammer 2000). In Korea, this species was found in the Sohwangbyeongsanneup of Mountain Odae in Pyeongchang and some peatlands near

104 96 Algal Flora of Korea Freshwater Diatoms VII A 10 μm B C D Fig. 91. Pinnularia subgigas. A D. view of valves (A: 1000, B D: 800, LM).

105 Pennales: Naviculaceae: Pinnularia 97 Mountain Jeongjok in Yangsan, May This is the first report in Korea. REMARKS: This species differs from Pinnularia gigas Ehrenberg in its complex raphe, the narrower axial area and the outline of the valve (Krammer 2000). 46. Pinnularia subinterrupta Krammer et Schroeter in Krammer 1992b: 111 (Figs. 93, 94). Hustedt 1930: 317. f Krammer 2000: 116. pl. 88. f. 44. REPLACED SYNONYM: Pinnularia interrupta var. minutissima Hustedt 1924: 566. Hustedt 1930: 317. f Valves broadly linear in outline, the margins parallel with a shoulder between the center and the end of the valve, the ends of the valve distinctly capitate. Raphe delicately filiform, the central ends of the raphe slightly bent to one side, central pores distinct, the terminal ends of the raphe indistinct in light microscopy observation. Axial area very narrow, central Fig. 92. Distribution of Pinnularia subgigas. area round and 1/2 2/3 of the valve breadth. Striae rows in 10 μm, slightly radiate in the middle and becoming convergent over the ends. Valves μm in length and 4 5 μm in breadth. 10 μm A B C D E F G H I Fig. 93. Pinnularia subinterrupta. A I. view of valves ( 2000, LM).

106 98 Algal Flora of Korea Freshwater Diatoms VII TYPE LOCALITY: Sarek Mountains, Swedish Lappland, spring in a swamp. ECOLOGY AND DISTRIBUTION: Pinnularia subinterrupta is abundant in oligotrophic freshwater with low mineral concentration (Krammer 2000). In Korea, this species is found from peatlands, Mujechineup of Mountain Jeongjok in Ulsan and the Yongneup Bog of Mountain Daeam in Inje and in May REMARKS: This species is well distinguished by the size and outline of the valve, and the form of the central area. Fig. 94. Distribution of Pinnularia subinterrupta. 47. Pinnularia transversa (A. Schmidt) Mayer 1940: 143 (Figs. 95, 96). Patrick and Reimer 1966: 630. pl. 61. f Noh 1991: 182. f. 67d. Krammer 2000: 171. pl f. 1. BASIONYM: Navicual transversa A. Schmidt in Schmidt et al pl. 43. f. 5. SYNONYM: Pinnularia major var. transversa (A. Schmidt) Cleve 1891: 24. Valves elongately linear in outline, swollen in the middle and the ends of the valve broadly rounded to swollen. Raphe strongly lateral, both lines of outer and inner fissures oppositely curved, the central ends of the raphe expanded and bent in one direction, the central pores moderately large, the terminal ends of the raphe shaped like large hooks. Axial area 1/3 1/2 of the valve breadth, tapering towards the ends. Central area a little wide than axial area and usually asymmetrical. Striae 8 9 rows in 10 μm, radiate in the middle and weakly convergent in the ends. Longitudinal lines distinct across the striae. Valves (317) μm in length and μm in breadth. TYPE LOCALITY: Monticello, northern America. ECOLOGY AND DISTRIBUTION: Pinnularia transversa is probably cosmopolitan like P. neomajor Krammer and is most common in cold, oxygen-rich and low electrolyte water (Krammer 2000).

107 Pennales: Naviculaceae: Pinnularia 99 A B C D E F 10 μm Fig. 95. Pinnularia transversa. A F. view of valves (A: 1250, B, C: 750, D: 1200, E: 800, F: 1500, LM).

108 100 Algal Flora of Korea Freshwater Diatoms VII This species was found in streams, lakes, swamps of the lowlands (Noh 1991). Recently, it rarely occurs epiphytic algal assemblages in the Seonakdong River. This is the first report in Korea. REMARKS: This species is distinguished from more elongated forms of P. neomajor by the higher ratios of length and breadth of the valve, (Krammer 2000). Fig. 96. Distribution of Pinnularia transversa. 48. Pinnularia viridiformis Krammer 1992b: 160 (Figs ). Krammer 2000: 167. pl f. 1. REPLACED SYNONYM: Pinnularia viridis var. minor Cleve 1891: 22. f. 1. SYNONYM: Pinnularia streptoraphe var. minor (Cleve) Cleve 1895: 93 pro parte. Valves broad and linear in outline, the margins of the valve parallel or slightly swollen narrowing towards the rounded ends. Raphe broadly lateral, occasionally semicomplex, the outer fissures of the raphe curved, the central ends of the raphe bent to one side, the central pores small round and close together, the terminal ends of the raphe shaped like question marks. Axial area linear, 1/5 1/4 of the valve breadth, central area asymmetrically roundish, a little wider than the axial area. Striae 7 9 in 10 μm, slightly radiate in the middle and becoming parallel to slightly convergent over the ends. Longitudinal bands distinct across the striae along the apical axis. Valves μm in length and μm in breadth. TYPE LOCALITY: Padasjoki, Tavastland län, Kemi lappmark, Finland (fossil). ECOLOGY AND DISTRIBUTION: Pinnularia viridiformis is cosmopolitan, one of the most common Pinnularia taxa in oligotrophic to mesotrophic freshwaters with low electrolyte (Krammer 2000). In

109 Pennales: Naviculaceae: Pinnularia 101 A B 10 μm C D Fig. 97. Pinnularia viridiformis (1). A D. view of valves (A C: 2000, D: 1500, LM).

110 102 Algal Flora of Korea Freshwater Diatoms VII A B 10 μm C D Fig. 98. Pinnularia viridiformis (2). A D. view of valves (A, C, D: 2000, B: 1500, LM).

111 Pennales: Naviculaceae: Pinnularia 103 a study of diatom assemblages on sediments collected from 30 clear Florida lakes, it has a wide range of tolerance from oligotrophic to eutrophic state and is most common in circumneutral conditions (Whitmore 1989). In Korea, this species was collected from two streams, a swamp and a mountain wetland in 1987 and reported as Pinnularia viridis var. minor Cleve (Noh 1991). It is rarely found from the Mujechineup of Mountain Jeongjok in Ulsan, December 2009 and the Yongneup Bog of Mountain Daeam in Inje, May REMARKS: Most of the forms designated in the past as P. viridis (Nitzsch) Ehrenberg belong to P. viridiformis. The latter is distinguished from P. viridis by outline, size, semicomplex raphe form, which has less striae in 10 μm, has broader axial area, and is much larger. Also this species is different to the much larger P. neomajor Krammer, which has always smaller longitudinal bands and larger length-to-breadth ratio (Krammer 2000). Fig. 99. Distribution of Pinnularia viridiformis. 49. Pinnularia viridiformis var. minor Krammer 1992b: 160 (Figs. 100, 101). Krammer 2000: 168. pl f. 1. Valves broad in breadth and linear in outline, the margins of the valve parallel or slightly swollen narrowing towards the rounded ends. Raphe broadly lateral, occasionally semicomplex, the outer fissures of the raphe curved, the central ends of the raphe bent to one side, the central pores small round and close together, the terminal ends of the raphe shaped like question marks. Axial area linear, 1/5 1/4 of the valve breadth, central area asymmetrically slightly rounded, a little wider than the axial area. Striae 7 9 rows in 10 μm, slightly radiate in the middle and becoming parallel to slightly convergent over the ends. Longitudinal bands distinct across the striae along the apical axis. Valves μm in length and μm in breadth. TYPE LOCALITY: Kosbach, Franken, Bavaria in Germany. ECOLOGY AND DISTRIBUTION: This variety is widely distributed in oligotrophic water like the nominal variety. In Korea, it was rarely found in the Yongneup Bog of Mountain Daeam in Inje, May This is the first report in Korea.

112 104 Algal Flora of Korea Freshwater Diatoms VII A 10 μm B C D E Fig Pinnularia viridiformis var. minor. A E. view of valves ( 2000, LM).

113 Pennales: Naviculaceae: Pinnularia 105 REMARKS: Pinnularia viridiformis is distinguished by outline, size, semicomplex raphe from P. viridis, which has less striae in 10 μm, broader axial area and wider breadth, more than 21 μm. Also this species is different to the much larger P. neomajor, which has always smaller longitudinal bands and larger length-tobreadth ratio (Krammer 2000). Fig Distribution of Pinnularia viridiformis var. minor. 50. Pinnularia viridis (Nitzsch) Ehrenberg 1843: 305 (Figs. 102, 103). Patrick and Reimer 1966: 639. pl. 64. f. 1. Krammer and Lange-Bertalot 1986: 428. pl f. 1. Krammer 2000: 175. pl f. 1. BASIONYM: Bacillaria viridis Nitzsch 1817: 97. pl. 6. f. 1. SYNONYM: Frustulia viridis (Nitzsch) Kützing 1833: 551. Navicula viridis (Nitzsch) Ehrenberg 1835: 266. Schizonema viride (Nitzsch) Kuntze 1898: 555. Valves linear in outline, the margins of the valve parallel or very weakly convex or triundulate, narrowing towards the rounded ends. Raphe broadly lateral to semicomplex, the outer fissures of the raphe undulate, sometimes three lines of the raphe visible, the central ends of the raphe laterally bent in one direction, the central pores small round, the terminal ends shaped like question marks. Axial area linear narrowing towards the ends, occupying 1/5 1/4 of the valve breadth, and central area a little wide than the axial area and commonly asymmetrical. Striae 6 7 rows in 10 μm, radiate in the middle and becoming parallel or slightly convergent over the ends, and distinctly crossed by two or three longitudinal bands. Valves μm in length and μm in breadth.

114 106 Algal Flora of Korea Freshwater Diatoms VII A B 10 μm C D Fig Pinnularia viridis. A D. view of valves ( 2000, LM).