Algal Flora of Korea

Volume 3, Number 15 Chrysophyta: Bacillariophyceae: Pennales: Raphidineae: Naviculaceae: Stauroneis, Craspedostauros, Caloneis III, Pinnularia III Freshwater Diatoms XI

2015

National Institute of Biological Resources Ministry of Environment

Algal Flora of Korea

Volume 3, Number 15 Chrysophyta: Bacillariophyceae: Pennales: Raphidineae: Naviculaceae: Stauroneis, Craspedostauros, Caloneis III, Pinnularia III Freshwater Diatoms XI

Gyeongje Joh Inje University Algal Flora of Korea Volume 3, Number 15 Chrysophyta: Bacillariophyceae: Pennales: Raphidineae: Naviculaceae: Stauroneis, Craspedostauros, Caloneis III, Pinnularia III Freshwater Diatoms XI

Published by the National Institute of Biological Resources Environmental Research Complex, Hwangyeong-ro 42, Seo-gu Incheon 22689, Republic of Korea www.nibr.go.kr

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

ISBN : 9788968111709-96470 Government Publications Registration Number 11-1480592-000819-01

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

Publisher : Kim, Sang-Bae Author : Gyeongje Joh

Published on November 30, 2015

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. Chlorococcales: 1

Preface

The biological resources include all the composition of organisms and genetic resources which possess the practical and potential values essential to human live. Biological resources will be firmed competition of the nation because they will be used as fundamental sources to make highly valued products such as new lines or varieties, new material, and drugs. As the Nagoya Protocol th was adopted in 2010 and entered into force in the 12 Conference of Parties of the Convention on

Biological Diversity (CBD) in 2014, it is expected that the competition to get biological resources will be much intensive under the rapidly changed circumstance on the access and benefic sharing of the genetic resources (ABS). Therefore, each nation is investigating and clearing information of native species within its territory in order to secure its sovereignty rights over biological resources. The National Institute of Biological Resources of the Ministry of Environment has been publish- ing the ‘Flora and Fauna of Korea’ since 2006 to manage biological resources in comprehensive ways and to enhance national competitiveness by building up the foundation for the sovereignty over biological resources. Professional research groups consisting of professors and related experts of taxonomy examined systematically a total of 12,631 species for the past eight years to publish 151 volumes in both Korean and English versions, and two volumes of World Monograph cover- ing 216 species. This year, 11 volumes of the Flora and Fauna of Korea in both Korean and English versions including 517 species of invertebrates, insects, vascular plants, algae and fungi are addi- tionally published. Flora and Fauna of Korea were the first professional records to describe all the species of the nation in a comprehensive way, and they would contribute to level up the taxonomic capacity. Furthermore, publication of flora and fauna through identification of native species and investigation of national biota would be helpful to declare sovereignty rights over our native biological resources, be used as positive proof, and be utilized to provide the basic information of biological resources for industrial application. The National Institute of Biological Resources of the Ministry of Environment will continue to accelerate the project of the publication of the ‘Flora and Fauna of Korea’. Personally I would like to express my sincere appreciation for those experts, Professor Gyeongje Joh of Inje University, who has continuously made a lot of efforts to publish an excellent version of Korean flora.

Sang-Bae, Kim President National Institute of Biological Resources

Contents

List of Taxa 3 Introduction 5 Materials and Methods 6 Taxonomic Notes 7

1. Stauroneis acuta W. Smith 8 2. Stauroneis alabamae Heiden 10 3. Stauroneis amphicephala Kützing 12 4. Stauroneis anceps Ehrenberg 14 5. Stauroneis ancepsopsis Lange-Bertalot, Cavacini, Tagliaventi et Alfinito 15 6. Stauroneis borgei Manguin 17 7. Stauroneis circumborealis Lange-Bertalot et Krammer 18 8. Stauroneis elena Kulikovskiy, Lange-Bertalot et Witkowski 20 9. Stauroneis gracilis Ehrenberg 22 10. Stauroneis gracillima Hustedt 24 11. Stauroneis heinii Lange-Bertalot et Krammer 25 12. Stauroneis jarensis Lange-Bertalot, Cavacini, Tagliaventi et Alfinito 27 13. Stauroneis kriegeri Patrick 29 14. Stauroneis kuelbsii Lange-Bertalot 31 15. Stauroneis lacusvulcani Rioual 33 16. Stauroneis nobilis Schumann 35

17. Stauroneis prominula (Grunow) Hustedt 35 18. Stauroneis pseudosmithii Van de Vijver et Lange-Bertalot 38 19. Stauroneis reichardtii Lange-Bertalot, Cavacini, Tagliaventi et Alfinito 39 20. Stauroneis respectabilis Lange-Bertalot, Cavacini, Tagliaventi et Alfinito 41 21. Stauroneis sagitta Cleve 43

22. Stauroneis siberica (Grunow) Lange-Bertalot et Krammer 44 23. Stauroneis smithii Grunow 46 24. Stauroneis sonyae Kulikovskiy, Lange-Bertalot, Witkowski et Dorofeyuk 47 25. Stauroneis staurolineata var. japonica Kobayasi et Ando 49 26. Stauroneis subaustralis Van de Vijver et Lange-Bertalot 51 27. Stauroneis subgracilis Lange-Bertalot et Krammer 53

28. Craspedostauros indubitabilis (Lange-Bertalot et Genkal) Cox 56 29. Caloneis aequatorialis Hustedt 58 30. Caloneis bryophila Manguin 60

31. Caloneis intermedia (Manguin) Moser, Lange-Bertalot et Metzeltin 61

32. Caloneis odiosa (Manguin) Moser, Lange-Bertalot et Metzeltin 62 33. Pinnularia anglica Krammer 64

34. Pinnularia brebissonii (Kützing) Rabenhorst 66 35. Pinnularia brevicostata var. sumatrana Hustedt 68 36. Pinnularia gibba var. cuneata Manguin 70 2 Algal Flora of Korea·Freshwater Diatoms XI

37. Pinnularia meridiana Metzeltin et Krammer 72

38. Pinnularia mesolepta (Ehremberg) W. Smith 74 39. Pinnularia nipponica Skvortzow 76

40. Pinnularia tirolensis (Metzeltin et Krammer) Krammer 78 Literature Cited 80 Index to Korean Names 86 Index to Korean Names as Pronounced 87 Index to Scientific Names 88 3

List of Taxa

Class Bacillariophyceae Order Pennales Suborder Raphidineae Family Naviculaceae Kützing 1944 Genus Stauroneis Ehrenberg 1843 Stauroneis acuta W. Smith 1853 Stauroneis alabamae Heiden 1903 Stauroneis amphicephala Kützing 1844 Stauroneis anceps Ehrenberg 1843 Stauroneis ancepsopsis Lange-Bertalot, Cavacini, Tagliaventi et Alfinito 2003 Stauroneis borgei Manguin 1941 Stauroneis circumborealis Lange-Bertalot et Krammer 1999 Stauroneis elena Kulikovskiy, Lange-Bertalot et Witkowski 2010 Stauroneis gracilis Ehrenberg 1843 Stauroneis gracillima Hustedt 1943 Stauroneis heinii Lange-Bertalot et Krammer 1999 Stauroneis jarensis Lange-Bertalot, Cavacini, Tagliaventi et Alfinito 2003 Stauroneis kriegeri Patrick 1945 Stauroneis kuelbsii Lange-Bertalot 2004 Stauroneis lacusvulcani Rioual 2013 Stauroneis nobilis Schumann 1867

Stauroneis prominula (Grunow) Hustedt 1959 Stauroneis pseudosmithii Van de Vijver et Lange-Bertalot 2004 Stauroneis reichardtii Lange-Bertalot, Cavacini, Tagliaventi et Alfinito 2003 Stauroneis respectabilis Lange-Bertalot, Cavacini, Tagliaventi et Alfinito 2003 Stauroneis sagitta Cleve 1881

Stauroneis siberica (Grunow) Lange-Bertalot et Krammer 1996 Stauroneis smithii Grunow 1860 Stauroneis sonyae Kulikovskiy, Lange-Bertalot, Witkowski et Dorofeyuk 2010 Stauroneis staurolineata var. japonica Kobayasi et Ando 1978 Stauroneis subaustralis Van de Vijver et Lange-Bertalot 2004 Stauroneis subgracilis Lange-Bertalot et Krammer 1999 Genus Craspedostauros E.J. Cox 1999

Craspedostauros indubitabilis (Lange-Bertalot et Genkal) Cox 1999

Genus Caloneis Cleve 1894 (3) Caloneis aequatorialis Hustedt 1921 Caloneis bryophila Manguin 1996

Caloneis intermedia (Manguin) Moser, Lange-Bertalot et Metzeltin 1998

Caloneis odiosa (Manguin) Moser, Lange-Bertalot et Metzeltin 1998

Genus Pinnularia Ehrenberg 1843 (3) Pinnularia anglica Krammer 1992

Pinnularia brebissonii (Kützing) Rabenhorst 1864 4 Algal Flora of Korea·Freshwater Diatoms XI

Pinnularia brevicostata var. sumatrana Hustedt 1934 Pinnularia gibba var. cuneata Manguin 1996 Pinnularia meridiana Metzeltin et Krammer 1998

Pinnularia mesolepta (Ehremberg) W. Smith 1853 Pinnularia nipponica Skvortzow 1936

Pinnularia tirolensis (Metzeltin et Krammer) Krammer 2000 5

Introduction

The genus Stauroneis in the history of diatom classification was already established in 1843 by C. G. Ehrenberg and its taxonomic status is characterized by stauros, a thickened hyaline fascia in the central parts of the valve. The diatoms belonging to genus Stauroneis are distinguished from the other naviculoid species by known as central nodule such as stauros, and it is a primary criterion (Andrews 1981). The stauros is characteristic and easily recognized under light microscopy. In ad- ditions, features used to identify species of Stauroneis diatoms are the outline and dimensions of the valve, the structure of the raphe and its external central endings, the number of striae and poroids occurring in a 10 segment, and the presence of pseudosepta at the ends (Van de Vijver et al. 2004). Besides of the genus Stauroneis Ehrenberg 1843, many taxa have stauros-like structure on valve, and they are Stauronella Mereschkowsky 1901, Staurophora Mereschkowsky 1903, Stauropsis Meuni- er 1910, Haslea Simonsen 1974, Proschkinia Karayeva 1978, Craspedostauros Cox 1999. Many hyaline stauros strips in the central area are derived from strong transapical ribs or their arrangements. The thickened central strips have evolved independently in several groups of raphid diatoms and they are not closely related to Stauroneis as homoplastic characters (Round et al. 1990; Cox and Wil- liams 2000). In addition, the stauros-bearing taxa show diverse structure and arrangement of chlo- roplast and raphe-sternum in cells. Diatoms of this genus, Stauroneis Ehrenberg sensu stricto, include obligate freshwater ones, cosmopolitan ones as benthic diatoms in various habitats and subaerial diatoms in moist soil and moss (Round et al. 1990; Van de Vijver et al. 2004). Approximately 140 speces and intraspecies have been currently accepted taxonomically (Guiry and Guiry 2012). Stauroneis diatoms are distributed worldwide in temperature regions (Bahls 2010), in the Antarctic and Arctic regions (Van de Vijver et al. 2004), and in the tropical area (Metzeltin and Lange-Bertalot 1998). A large numbers of Stauroneis species are endemic, being restricted to a local and regional habitat. Many species are newly reported, including 40 species, among the 63 Stauroneis described in the Arctic and Antarctic regions (Van de Vijver et al. 2005), 21 of the 52 species in the northern Rockies (Bahls 2010), and 19

Stauroenis species in tropical South America (Metzeltin and Lange-Bertalot 1998, 2007). Bahls (2012) reported the characteristic distribution of genus Stauroneis and its biogeographic disjunction. Approximately 20 taxa of Stauroneis have been previously found in a floristic study of freshwater diatoms in Korea (Lee et al. 1995; Lee 1997), and they were primarily S. anceps Ehrenberg, S. phoenicenteron (Nitzsch) Eherenberg and their varieties or forma. As many species have been separated from the two former taxa and designated to new species, and S. anceps and S. phoenicenteron sensu stricto were hardly found throughout this study. This monograph reports the species of genus Stauroneis and some other taxa residing largely in the wetland or mountain peatlands and provides detailed information on their biogeography in Korea. 6 Algal Flora of Korea·Freshwater Diatoms XI

Materials and Methods

Diatom materials were collected from various areas, including many mountain wetlands, head- waters and reservoirs of streams and rivers, lowland wetland, and other habitats. In the case of wetlands, the collection was primarily from peatlands, which are largely concentrated on the sum- mit of mountains from 600 m to 1100 m elevations, while some are at lower altitudes. The mountain peatlands of Korea are commonly covered with Japanese moor grass (Molinia japonica) and other grasses, and surrounded by konara oak (Quercus serrata) and pine trees (Pinus densiflora) at their borders. These grass plants provide the major contribution to peat deposit and formation. Small pools or water holes are scattered throughout the mountain wetlands. In these pools, the slurries or suspension in the water holes were 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. Preparations of the diatom specimens, from digestive oxidation to permanent mounting, fol- lowed the standard procedures of APHA (1995). Diatoms were observed under oil immersion by using an Axioplan microscope (Carl Zeiss, Oberkochen, Germany). For each sample, the percent- age of each taxon was evaluated by counting 350 to 400 valves. The identification and confirmation of Stauroneis species are done according to the work of Hustedt (1959), Krammer and Lange-Ber- talot (1986), Van de Vijver et al. (2004), and many other articles. The synonym and the taxonomical informations of the species are referred to website of California Academy of Science (CAS) (http:// www.calacademy.org) (Fourtanier and Kociolek 2011) and AlgaeBase (Guiry and Guiry 2012, 2015) on-line version. The check-lists authorized by Lee et al. (1995) and Lee (1997), and other documents after 1997 were the source documents used to sum up the Stauroneis flora reported in Korea. Pennales: Naviculaceae: Stauroneis 7

Taxonomic notes

Genus Stauroneis Ehrenberg 1843: 45.

Sip-ja-dol-mal-sok (십자돌말속)

Valves linear, lanceolate or elliptic-lanceolate in outline with rounded to often capitate ends. Sometimes a ridge developed at the junction of the valve face and mantle. Valve margins thickened occasionally to form pseudosepta at the terminal ends. Raphe straight and sometimes em- bedded in the longitudinal ribs, the external ends of central raphe strongly expanded and often curved to same directions, the internal ends of the central raphe simple or slightly curved, the terminal ends of the raphe curved to the secondary sides. Axial area narrow, the hyaline fascia of the central area distinctly transverse, known as stauros, occasionally one to several short striae on the valve margins. Striae uniseriate and punctate, the areolae round or elongate. Two plastids, one on each side of the girdle.

Lectotype: Stauroneis phoenicenteron (Nitzsch) Ehrenberg 1841 (1843): 311 (=Bacillaria phoenicenteron Nitzsch 1817)

Species: The 820 species and infraspecific taxa have been reported worldwide, of which approximately 140 have been currently accepted taxonomically (Guiry and Guiry 2012). In Korea, 22 taxa have been reported from freshwaters according to Lee (1997) and other literatures. Distribution: Stauroneis species occurs in both benthic and planktonic habitats in freshwaters, and they are more obligate in soft bottom as epipelic forms (Round et al. 1990). They are consid- ered to be low in species diversity, abundance or dominance, but many new taxa have been reported from some regions, South America, the Sardinia Island in Mediterranean Sea, Antarctic regions,

Rockies Mountains in USA (Metzeltin and Lange-Bertalot 1998; Lange-Bertalot et al. 2003; Van de Vijver et al. 2004; Bahl 2010).

Key reference: Lange-Bertalot et al. (2003), Van de Vijver et al. (2004), Van de Vijver (2005), Bahls (2010). 8 Algal Flora of Korea·Freshwater Diatoms XI

1. Stauroneis acuta W. Smith 1853: 59 (Figs. 1, 2).

Hustedt 1930: 259. f. 415. Van de Vijver et al. 2004: 17. pl. 94. f. 1.

Valves rhombic-lancolate in outline, the margins of the valve largely swollen or convex in middle, the ends of the valve bluntly rounded and not protracted, and pseudosepta distinctly developed in the ends. Raphe broadly lateral, the central pores of the raphe expanded, reversely deflected to a side and. Axial area wide, linear and widening towards the central area. The hyaline stauros of the central area transverse up to the margins, a little expanded towards the margins. Striae moderately radiate in the middle, strongly radiate towards the ends, 12-16 rows in 10 μm on valves, and areolae 14-16 in 10 μm of a stria.

Valves 80-166 (180) μm in length and 15-40 μm in breadth.

Type locality: Uncertian, New Zealand? Ecology and distribution: This species occurs in a variety of freshwaters, in New Zealand, in Alaska, in Iceland and in Antaectica region (Van de Vijver et Fig. 1. Distribution of Stauroneis acuta. al. 2004). In Korea, S. acuta was found in Lake Yeong- nang in Sokcho of Gangwon-do (Cho and Park 1969), in freshwaters of Gyeongju of Gyeongsangbuk-do (Chung and Watanabe 1984), in Demitarized

Zone of Gangwon-do (Cho et al. 1987), and in waterfalls of Jeju Island (Lee and Chang 1991). This species occurred recently in Hanon Crater Lowland of Jeju Island. Pennales: Naviculaceae: Stauroneis 9

A B C D

Fig. 2. Stauroneis acuta. A-D. The morphology of valves (Scales: 10 μm, A, B: ×2000, C, D: ×1500). 10 Algal Flora of Korea·Freshwater Diatoms XI

2. Stauroneis alabamae Heiden in Schmidt et al. 1903: 242 (Figs. 3, 4).

Schmidt et al. 1903. pl. 242. f. 2. Hustedt 1930: 257. f. 412.

Valves elliptical to rhombic-lancolate in outline, sharply narrowing towards the ends of the valve, and the ends not protracted and obtusely round. Raphe straight, the central ends of the raphe straight, the central pores not curved. Axial area widely linear, narrowing towards terminal and central ends, and the hyaline stauros of central area narrow rectangular, hardly expanded towards the margins. No marginal striae in the central area. Striae radiate in the middle and more radiate towards the ends, 14-17 rows in 10 μm. Valves 100-185 μm in length and 24-38 μm in breadth.

Type locality: Fossil in Montogomery of Alabama. Ecology and distribution: This species occurs rarely in freshwaters (Hustedt 1930). In Korea, it was found rarely in swamps in Haman of Gyeong- sangnam-do (Chung and Noh 1987), recently in Dong- song Reservoir in Cheolwon of Gangwon-do and in Fig. 3. Distribution of Stauroneis alaba- Hanon Crater Lowland of Jeju Island. mae. Pennales: Naviculaceae: Stauroneis 11

Fig. 4. Stauroneis alabamae. A-C. The morphology of valves (Scales: 10 μm, A: ×1000, B, C: ×1200). 12 Algal Flora of Korea·Freshwater Diatoms XI

3. Stauroneis amphicephala Kützing 1844: 105 (Figs. 5, 6).

Van de Vijver et al. 2004: 20. pl. 51. f. 1.

Synonym: Stauroneis anceps var. amphicephala (Kützing)

Van Heurck 1885: 69. pl. 4. f. 6 (1880)

Stauroneis anceps var. amphicephala (Kützing) Cleve 1894: 148.

Valves elliptic-lanceolate in outline, the ends of the valve protracted and rostrate. Raphe moderately lateral in the middle, the central ends of the raphe deflected, Axial area moderately narrow and the hyaline stauros of the central area rectangular and more expanded towards the margins. Striae radiate throughout more or less wavy in the central parts of the valve and more radiate in the ends, 19-22 rows in 10 μm, and areolae on a stria 20 puncta in 10 μm. Valves 45- 55 μm in length and 11-13 μm in breadth.

Type locality: Nordhausen in Thuringia, Germany. Ecology and distribution: This species is distributed widely in a variety of standing waters from small pools to large lakes, and in the Arctic region (Van de Fig. 5. Distribution of Stauroneis am- Vijver et al. 2004). In Korea, it occurs rarely in a pad- phicephala. dy rice field in Wando of Jeollanam-do. Remarks: This species is often confused with S. anceps Ehrenberg sensu stricto and S. amphicephaloides Metzeltin et Lange-Bertalot. S. amphicephaloides is larger in dimension and its transapical striae are more coarse than S. amphicephala (Van de Vijver et al. 2004). Pennales: Naviculaceae: Stauroneis 13

C A B

F D E

Fig. 6. Stauroneis amphicephala. A-F. The morphology of valves (Scale: 10 μm, ×2000). 14 Algal Flora of Korea·Freshwater Diatoms XI

4. Stauroneis anceps Ehrenberg 1843: 306 (18) sensu stricto (Figs. 7, 8).

Lange-Bertalot et al. 2003: 132. pl. 35. p. 1. Van de Vi- jver et al. 2004. pl. 43. p. 1. Bahl 2011a in Diatoms of the United States.

Valves lanceolate to linear-lanceolate in outline, the ends of the valve moderately protracted and rostrate. Raphe lateral from the middle, the central ends of the raphe slightly expanded and deflected. Axial area moderately narrow, linear-lanceolate, and the hyaline stauros of the central area rectangular and slightly expanded towards the margins. Striae radiate throughout more or less wavy in the central parts of the valve and more radiate in the ends, 22-24 rows in 10 μm. Valves 40-70 μm in length and 10-13 μm in breadth.

Type locality: Cayenne in French Guiana. Ecology and distribution: Stauroneis anceps is one of the most frequent and widespread species in the freshwaters, but its abundance or dominance is relatively low. In Korean freshwaters, this species was reported 22 times before 1997 (Lee 1997). However, S. anceps sensu stricto is found less frequently, occurring Fig. 7. Distribution of Stauroneis anceps.

G F D E B C A

Fig. 8. Stauroneis anceps. A-G. The morphology of valves (Scale: 10 μm, ×2000). Pennales: Naviculaceae: Stauroneis 15 rarely in lowland wetlands in Haman of Gyeongsangnam-do. Remarks: This species is the most popular and common in this genus worldwide, and was described in a variety of waters without the type specimens. Reichardt (1995) designated the lectotype for this species from a specimen in Ehrenberg’s type material collected from Cayenne, French Guyana and many new species has been separated S. anceps sensu lato.

5. Stauroneis ancepsopsis Lange-Bertalot, Cavacini, Tagliaventi et Alfinito

2003: 138 (Figs. 9, 10).

Lange-Bertalot et al. 2003: 132. pl. 38. f. 15.

Valves elliptic-lanceolate to elliptical in outline, the ends of the valve abruptly protracted, comparatively long and rostrate. Raphe almost straight or weakly lateral, the central ends of the raphe straight, the central pores of the raphe not curved. Axial area narrow, linear-lanceolate, the hyaline stauros of the central area narrowly rectangular and not expanded towards the margins. Striae slightly radiate in the central parts of the valve and strongly radiate towards the ends,

19?20 (not 22-24) rows in 10 μm, and areolae on a stria ca. 18 (not 24) puncta in 10 μm. Valves 52-65 μm in length and 13.5-14.5 μm in breadth.

Type locality: Pauli Murdegu in Insula Sardinia, Italy. Ecology and distribution: This species was newly recorded from Sardinia Island of the Mediterranean

Sea (Lange-Bertalot et al. 2003). It occurs rarely in a paddy rice field in Wando of Jeollanam-do. Remarks : This species is differentiated from Stau- Fig. 9. Distribution of Stauroneis anceps- roneis anceps Ehrenberg sensu stricto a neighboring opsis. species, Stauroneis accedens Lange-Bertalot et al., has narrower breadth in valves than S. ancepsopsis, and S. amphicephala Kützing has more linear-lanceolate valve in outline and more expanded central area towards the margin (Lange-Bertalot et al. 2003). 16 Algal Flora of Korea·Freshwater Diatoms XI

A B C

D E

Fig. 10. Stauroneis ancepsopsis. A-E. The morphology of valves (Scale: 10 μm, ×2000). Pennales: Naviculaceae: Stauroneis 17

6. Stauroneis borgei Manguin in Allorge and Manguin 1941: 179 (Figs. 11, 12).

Hustedt 1959: 811. f. 1157h.

Basionym: Stauroneis borgei Manguin in Allorge and Manguin 1941: 179. f. 69.

Synonym: Stauroneis smithii var. borgei (Manguin) Hustedt 1959: 811. f. 1157h.

Valves elliptical to rhombic-lanceolate with convex margins, the ends of the valve not protracted, and pseudosepta developed near the ends. Raphe filiform and straight, and the central ends of the raphe distinctly expanded. Axial area narrow and linear, and the hyaline stauros of the central area narrow rectangular, extending to the margins of the valve. Tran- sapical striae parallel or slightly radiate throughout the valve, fine and not resolved in light microscopy. Valves about 20 μm in length, about 7.5 μm in breadth.

Type locality: Basque in the western Pyrenees mountains of Etang d’Yrieu, France. Ecology and distribution: Stauroneis borgei is distributed in the Canadian Arctic Archipelago (Bouch- Fig. 11. Distribution of Stauroneis borgei. ard et al. 2004). This species was rarely found only in a lake, Dongsong Reservoir in Cheolwon of Gang- won-do.

Fig. 12. Stauroneis borgei. The morphology of a valve (Scale: 10 μm, ×2000). 18 Algal Flora of Korea·Freshwater Diatoms XI

7. Stauroneis circumborealis Lange-Bertalot et Krammer in Lange-Bertalot

and Genkal 1999: 90 (Figs. 13, 14).

Lange-Bertalot and Genkal 1999: 90. pl. 34. Van de Vi- jver et al. 2004: 31. pl. 10. f. 1.

Valves broadly elliptic-lanceolate in outline, the ends of the valve weakly protracted and obtusely rounded. Raphe distinctly lateral, the central ends of the raphe strongly curved and the terminal pores of the raphe drop-like expanded, the central pores of the raphe expanded and curved to the side. Axial area broad, slightly narrowing towards the central area. The hyaline stauros of the central area broadly rectangular, and expanded towards the margins with irregularly shortened striae in the margins. Striae radiate throughout, 15-16 rows in 10 μm on valves, puncta in a stria 14-16 in 10 μm. Valves 70-130 μm in length and 23-27 μm in breadth.

Type locality: Mestnyi Island near Yugorskij Pen- insula in the northwestern Siberia. Ecology and distribution: This species was reported firstly from the sediments of the Russian Arctic (Lange-Bertalot and Genkal 1999), from submerged Fig. 13. Distribution of Stauroneis cir- mosses of the Zackenberg area of Greenland (Van de cumborealis. Vijver et al. 2004) and from small lakes and ponds of the northern Rockies (Bahl 2011b). In Korea, it was found in the bottoms of a lowland wetland, Jilnalneup, of Haman Gyeongsangnam-do, in peatlands of Daegwanryeong Pasture of Pyeongchang, Gangwon-do and Jangdo Island of Sinan, Jeol- lanam-do. Remarks: Morphological similarity exists in some large Stauroneis species, but Stauroneis sonya differs by a large size, narrower ends of the valve and narrow axial area (Bahl, 2011f). Pennales: Naviculaceae: Stauroneis 19

A C D

Fig. 14. Stauroneis circumborealis. A-E. The morphology of valves (Scale: 10 μm, ×1500). 20 Algal Flora of Korea·Freshwater Diatoms XI

8. Stauroneis elena Kulikovskiy, Lange-Bertalot et Witkowski in

Kulikovskiy et al. 2010: 59 (Figs. 15, 16).

Kulikovskiy et al. 2010: 59. pl. 92. f. 1.

Valves elliptic-lancolate to broadly lanceolate in outline, the ends of the valve protracted and broadly rostrate. Raphe distinctly lateral with reverse-lateral ends, the central pores of the raphe expanded and bent to a side. Axial area rather wide linear or linear-lanceolate, slightly narrowing near the central ends. The hyaline stauros of the central area broadly transverse, expanded towards the margins and short striae absent in the margins of the central area. Striae moderately radiate in the middle, strongly radiate towards the ends, 15-17 rows in 10 μm on valves, and areolae 14-17 in a stria in 10 μm. Valves 103-134 μm in length and 23-27.5 μm in breadth.

Type locality: Sphagnum bog Nur in northern area of Hentai highlands, Mongolia. Ecology and distribution: This species occurs in the type locality, Sphagnum peatland, Nur, in the northern mountain region of Mongolia (Kulikovs- kiy et al. 2010). S. elena occurred frequently in some Fig. 15. Distribution of Stauroneis elena. mountain peatlands, Wangdeunjae of Jiri Mountain in Sancheong of Gyeongsangnam-do, Sohwangbyeong- san of Odae Mountain and peatlands of Daegwanryeong Pasture in Pyeongchnag of Gangwon-do. Remarks: The central area of this species is less expanded towards the margins and characteristic in the elliptic-lanceolate forms in outline (Kulikovskiy et al. 2010). Pennales: Naviculaceae: Stauroneis 21 m μ 10

B E D

Fig. 16. Stauroneis elena. A-E. The morphology of valves (Scales: 10 μm, ×1500). 22 Algal Flora of Korea·Freshwater Diatoms XI

9. Stauroneis gracilis Ehrenberg 1843: 386 (Figs. 17, 18).

Van de Vijver et al. 2004: 38. pl. 17. f. 1. Bahl 2011c in Diatoms of the United States.

Synonym: Stauroneis anceps var. gracilis (Ehrenberg) Brun 1880: 89. pl. 9. f. 2.

Stauroneis anceps var. gracilis (Ehrenberg) Cleve 1894: 147. Non synonym: Stauroneis anceps f. gracilis Rabenhorst 1864: 247.

Stauroneis anceps f. gracilis (Ehrenberg) Hustedt 1930: 256. f. 406.

Valves lancolate to somewhat linear-lanceolate in outline, and the ends of the valve slightly protracted and broadly subrostrate. Raphe distinctly lateral, the central ends of the raphe deflected, the central pores of the raphe expanded and curved to the side. Axial area relatively broad, linear to linear-lanceolate, narrow towards the central area. The hyaline stauros of the central area rectangular, slightly expanded towards the margins. Striae moderately radiate in the middle, more radiate towards the ends, 16-19 rows in Fig. 17. Distribution of Stauroneis graci- 10 μm on valves, and areolae 18-21 puncta in 10 μm. lis. Valves 75-114 μm in length and 15-19 μm in breadth.

Type locality: In Germany. Ecology and distribution: This species occurs in oligotrophic and mesotrophic waters, from small and shallow pools to large and deep lakes, and in wet soil and even dry mosses (Van de Vi- jver et al. 2004). It is widely distributed in both the Arctic and Antarctic regions (Van de Vijver et al. 2004). In Korea, it was found rarely in peatlands of Daegwanryeong Pasture of Pyeongchang of Gangwon-do, in lowlands, Jilnalneup of Haman and Daebong Reservoir in Changryeong of Gyeo- ngsangnam-do.

Remarks: In a bibliotheca of genus Stauroneis written by Van de Vijver et al. (2004), S. gracilis and S. subgracilis are closely related in the valve morphology, and it is hard to discriminate two species each other. The dimensions or breadth of the valve and the density of striae are used to differentiate the species, and the valve of S. gracilis is wider than 14 μm in breadth and striae are less than 19 rows in 10 μm. In addition, S. supergracilis Van de Vijver et Lange-Bertalot has narrower valves relative to the length and lower areola density, S. phoenicenteron has straight proximal raphe ends and lacks protracted ends (Bahl 2011d). Pennales: Naviculaceae: Stauroneis 23

A B C D

Fig. 18. Stauroneis gracilis. A-E. The morphology of valves (Scale: 10 μm, ×2000). 24 Algal Flora of Korea·Freshwater Diatoms XI

10. Stauroneis gracillima Hustedt 1943: 154 (Figs. 19, 20).

Krammer and Lange-Bertalot 1986: 248. pl. 90. f. 28. Simonsen 1987: 311. pl. 469: 7.

Synonym: Nupela gracillima (Hustedt) Lange-Bertalot 1993: 158.

Valves linear in outline, with parallel, slightly convex or concave margins, the ends of the raphe protracted and distinctly capitate. Raphe filiform, the central ends of the raphe straight, the central pores distinct. Axial area narrow linear and the hyaline stauros of the central area rectangular up to the margins. Striae and their puncta very fine and not resolved under light microscopy. Valves 12-21 μm in length and 3-5 μm in breadth.

Type locality: Graubünden near Davos, Switzer- land. Ecology and distribution: Stauroneis gracillima is known as moss attached diatoms from the central Eu- rope (Krammer and Lange-Bertalot 1986), and showed significant abundance in fossil diatom assemblages Fig. 19. Distribution of Stauroneis gracil- of a long sediment core from Lake Plešné, Czech Re- lima. public (Štefková 2008). In addition, it dominates the diatom assemblages on sediment in Lake Big Moose, which is ranged from pH 4.6 to 5.0 in strong acidic environments (Charles 1984). In Korea, this species occurs only in a peat wetland, the 1100 wetland, which name indicates the local altitude of the peatland, in Halla Mountain of Jeju Island.

Remarks: This species was transferred to Nupela as N. gracillima (Hustedt) Lange-Bertalot

(Lange-Bertalot 1993), and this genus occur mainly in oligotrophic waters (Wojtal 2009).

F A B C D E

Fig. 20. Stauroneis gracillima. A-F. The morphology of valves (Scale: 10 μm, ×2000). Pennales: Naviculaceae: Stauroneis 25

11. ‌Stauroneis heinii Lange-Bertalot et Krammer in Lange-Bertalot and

Genkal 1999: 91 (Figs. 21, 22).

Lange-Bertalot and Genkal 1999: 91. pl. 27. f. 1. Van de Vijver et al. 2004: 40. pl. 1. f. 1.

Valves lancolate in outline, the ends of the valve slightly protracted and broadly subrostarte. Raphe distinctly lateral, the central ends of the raphe deflected, the central pores of the raphe expanded and curved. Axial area linear or linear-lanceolate, narrowing towards terminal ends, and the hyaline stauros of the central area rectangular and weakly expanded towards the margins. No marginal striae in the central area. Striae moderately radiate throughout the valve, 15-16 (18) rows in 10 μm on valves, areolae 15-

18 puncta in 10 μm. Valves (110)125-167 μm in length and 23-27 μm in breadth.

Type locality: Mestnyi Island near Yugorskij Pen- insula in the northwestern Siberia. Ecology and distribution: This species was reported newly from the Siberian Arctica (Lange-Bertalot and Genkal 1999), and have been found in small and circumneutral pools with low conductivity in Arc- Fig. 21. Distribution of Stauroneis heinii. tic and Antarctic regions (Kulikovskiy et al. 2010). In Korea, it was found rarely in some peatlands, Muje- chi Peatland of Jeongjok Mountain in Ulsan, Daegwanryeong Pasture and Sohwangbyeongsan of Odae Mountian of Gangwon-do in Pyeongchang. Remarks: This species is similar to some large forms of Stauroneis taxa, but they differ in the size and dimensions, axial area and proximal raphe endings (Van de Vijver et al. 2004; Bahl 2011d). S. gracilis and S. supergracilis have narrower valves, relative to their length, and S. sonyae has elliptic-lanceolate or rhombic-lanceolate valves and a very wide stauros, often with numerous short striae. S. circumborealis Lange-Bertalot and Krammer has wider axial and central areas and more curved raphe endings in the center, and S. phoenicenteron Ehrenberg and S. rex Bahl have straight, rather than curved, proximal raphe ends. 26 Algal Flora of Korea·Freshwater Diatoms XI

A C

Fig. 22. Stauroneis heinii. A-E. The morphology of valves (Scales: 10 μm, A, B, D: ×1500, C, E: ×1200). Pennales: Naviculaceae: Stauroneis 27

12. ‌Stauroneis jarensis Lange-Bertalot, Cavacini, Tagliaventi et Alfinito

2003: 138 (Figs. 23, 24).

Lange-Bertalot et al. 2003: 138. pl. 38. f. 1. Van de Vi- jver et al. 2004: 44. pl. 31. f. 1.

Valves lanceolate to linear-lanceolate in outline, the ends of the valve protracted and rostrate. Raphe almost straight or weakly lateral, the central ends of the raphe slightly curved, the central pores of the raphe expanded. Axial area narrow to moderately broad, linear or linear-lanceolate, slightly narrow towards the central area. The hyaline stauros of the central area rectangular and expanded towards the margins. Stri- ae radiate throughout the valve, 20-24 rows in 10 μm on valves, and areolae 21-26 puncta in 10 μm. Valves 25-55 μm in length and 8.5-11 μm in breadth.

Type locality: Pauli Murtas in Insula Sardinia, Ita- ly. Ecology and distribution: This species is known from several pools of Sardinia Island in the Medterra- nean Sea (Lange-Bertalot et al. 2003) and from rarely in the Arctic regions (Van de Vijver et al. 2004). In Ko- rea, the species was found in Yongneup peatland of Fig. 23. Distribution of Stauroneis jaren- Daeam Mountain, in Inje Gangwon-do. sis. Remarks: Stauroneis jarensis is differentiated from related of S. anceps Ehrenberg, and has large similarity to adjacent taxa. This species differs in the central endings of the raphe and stria number with small forms of S. subgracilis Lange-Bertalot et Krammer, and in the valve outlines with S. anceps Eh- renberg (Lange-Bertalot et al. 2003; Van de Vijver et al. 2004). 28 Algal Flora of Korea·Freshwater Diatoms XI

D C B A

G F E

Fig. 24. Stauroneis jarensis. A-G. The morphology of valves (Scale: 10 μm, ×2000). Pennales: Naviculaceae: Stauroneis 29

13. Stauroneis kriegeri Patrick 1945: 175 (Figs. 25, 26).

Patrick and Reimer 1966: 362. pl. 30. f. 5. Krammer and Lange-Bertalot 1986: 248. pl. 90. f. 23. Van de Vijver et al. 2004: 45. pl. 59. f. 1.

Synonym: Stauroneis pygmaea Krieger 1929: 272. pl. 2. f. 28.

Valves linear-lanceolate in outline, with parallel to slightly convex margins, the ends of the valve broadly protracted and capitate. Raphe filiform, and straight or weakly lateral, the central ends of the raphe indistinct. Axial area narrowly linear. The hyaline stauros of the central area narrow rectangular reaching the margins. Striae weakly radiate in the middle, parallel to convergent towards the ends. Striae very fine and not resolved under light microscopy, slightly radiate and 26-30 rows in 10 μm on valves. Valves 17-24 μm in length and 4-6 μm in breadth.

Type locality: Lake Diebelsee in Hochmoor, Ger- many. Ecology and distribution: This species is com- Fig. 25. Distribution of Stauroneis krieg- monly distributed in the freshwater of lowlands, eri. headwater and mountains (Krammer and Lange-Ber- talot 1986), occurs in the Arctic and Antarctic regions

(Van de Vijver et al. 2005), and are abundant in bogs of Lake County, Illinois (Cochran-Stafira and Andersen 1984). In Korea, the species was found in two peatlands, Daeseongneup near Jeongjok Mountain in Yangsan of Gyeongsangnam-do and Jangdo Island in Sinan of Jeollanam-do, and in moss plants collected in Biseul Mountain in Dalseong of Daegu city, Misiryeong Mountain in Inje of Gangwon-do and Halla Mountain in Jeju. 30 Algal Flora of Korea·Freshwater Diatoms XI

A B C D E F G H

I J K L M N

Fig. 26. Stauroneis kriegeri. A-O. The morphology of valves (Scale: 10 μm, ×2000). Pennales: Naviculaceae: Stauroneis 31

14. ‌Stauroneis kuelbsii Lange-Bertalot in Van de Vijver et al. 2004: 45 (Figs. 27, 28).

Van de Vijver et al. 2004: 45. pl. 40. f. 1.

Valves lanceolate to linear-lanceolate in outline, the ends of the valve shortly protracted, subrostrate or subcapitate. Raphe moderately lateral in the middle, the central ends of the raphe deflected, Axial area moderately broad and the hyaline stauros of the central area narrow rectangular and slightly or not expanded towards the margins. Striae radiate in the central parts of the valve and more radiate in the ends, 15-18 rows in 10 μm, and areolae on a stria 18- 20 puncta in 10 μm. Valves 50-70 μm in length and 15-18 μm in breadth.

Type locality: In Franconia of Germany. Ecology and distribution: This species was newly reported from freshwaters in Germany (Van de Vijver et al. 2004). In Korea, it occurs rarely in a lowland wetland, Jilnalneup, in Haman of Gyeongsangnam-do. Remarks: This species can be often confused with Stauroneis gracilis Ehrenberg and S. subgracilis Lange-Bertalot et Krammer. But they differ by the Fig. 27. Distribution of Stauroneis kuelb- shape of the central area, the expansion of the central sii. area towards the margins and strial number (Van de Vijver et al. 2004). 32 Algal Flora of Korea·Freshwater Diatoms XI

A B C

Fig. 28. Stauroneis kuelbsii. A-C. The morphology of valves (Scale: 10 μm, ×2000). Pennales: Naviculaceae: Stauroneis 33

15. ‌Stauroneis lacusvulcani Rioual 2013: 49 (Figs. 29, 30).

Rioual et al. 2013: 49. f. 2.

Valves typically long lanceolate in outline, sharply tapering to the the ends, the ends of the valve long protracted and capitate. Raphe slightly lateral in the middle parts of the raphe, filiform at the terminal and central parts of the raphe, the central pores of the raphe not curved. Axial area narrow and more or less linear, and the hyaline stauros of the central area narrowly rectangular, expanded up to the margins. Striae parallel in the middle parts of the valve, radiate in the terminal parts of the valve, and more radiate toqrds the ends, very fine, 21-28 rows in 10 μm. Valves 62-

83 (90) μm in length and 9.2-12 (12.5) μm in breadth.

Type locality: Inner Mongolia, Great Xing’An Mountain, Sifangshan Tianchi. Ecology and distribution: Stauronei lacusvulcani occurred in the type locality in China (Rioual et al. 2013). In Korea, it was found in Dongsong Reservoir in Cheolwon of Gangwon-do, and is newly reported. Remarks: This species is closely related with S. Fig. 29. Distribution of Stauroneis la- gracilior Reichardt in the valve morphology, but dif- cusvulcani. fers by the valve dimension, the long protracted and strongly capitate ends, and the parallel striation in the middle part (Rioual et al. 2013). 34 Algal Flora of Korea·Freshwater Diatoms XI

A B C

Fig. 30. Stauroneis lacusvulcani. A-C. The morphology of valves (Scale: 10 μm, ×2000). Pennales: Naviculaceae: Stauroneis 35

16. Stauroneis nobilis Schumann 1867: 59 (Figs. 31, 32).

Krammer and Lange-Bertalot 1986: 242. pl. 87. f. 1.

Valves rhombic-lanceolate in outline, the ends of the valve long protracted and capitate. Raphe straight, the central ends of the raphe straight, the central pores not curved. Axial area widely linear, narrowing towards terminal and central ends, and the hyaline stauros of central area narrow rectangular, narrow towards the margins. The hyaline stauros of the central area rectangular and narrow towards the margins. Striae radiate in the central area and more radiate to- wars the ends, 14-17 rows in central area, and 19-21 rows in 10 μm in the terminal ends. Valves 100-185 μm in length, 23-38 μm in breadth.

Type locality: In Prussia of Germany. Ecology and distribution: Stauroneis nobilis is cosmopolitan in the freshwaters. This species occurs frequently in Korean freshwaters, the lowland wetlands in Haman of Gyeongsangnam-do (Chung and Noh,

1987), Gwang River in Uljin of Gyeongsangbuk-do (Lee et al. 1994), recently rare in Dongsong Reservoir Fig. 31. Distribution of Stauroneis nobil- in Cheolwon of Gangwon-do, and in Hanon Crater is. Lowland in Jeju Island.

17. ‌Stauroneis prominula (Grunow) Hustedt 1959: 805 (Figs. 33, 34).

Patrick and Reimer 1966: 366. pl. 30. f. 14. Krammer and Lange-Bertalot 1986: 247. pl. 90. f. 16.

Basionym: Pleurostauron (Pleurostaurum) prominulum Grunow in Cleve and Möller 1879: 261.

Synonym: Stauroneis parvula var. prominula (Grunow in Cleve and Möller) Grunow in Cleve 1894: 149. Stauroneis ignorata Hustedt 1939: 620. f. 48.

Valves linear with weakly three undulating margins, the ends of the valve narrow protracted and rostrate, and pseudoespta distinctly developed at the terminal parts of the valve. Raphe straight or the outer fissure of the raphe slightly lateral and filiform, the central ends of the raphe straight, the central pores of the raphe distinct. Axial area very narrow linear and the hyaline stauros of the central area transverse up to the margins. Striae parallel or weakly radiate, very fine and not resolved under light microscopy, 20-24 rows in 10 μm on valves. Valves 18-42 μm in length and 3-8 36 Algal Flora of Korea·Freshwater Diatoms XI

B C

Figs. 32. Stauroneis nobilis. A-C. The morphology of valves (Scales: 10 μm, A: ×1500, B, C: ×2000). Pennales: Naviculaceae: Stauroneis 37

μm in breadth.

Type locality: Tana of Finmark county in Gulf of Bothnia, Norway. Ecology and distribution: This species is cosmopolitan in stagnant and flowing waters, but not concentrated and rather scattered distribution (Krammer and Lange-Bertalot 1986). These diatoms prefer me- dium and high electrolytic conditions and even in the slightly brackish waters (Patrick and Reimer 1966). In Korea, the species was found in two peatlands, Jang- do Island in Sinan of Jeollanam-do and Oegogyeneup of Jiri Mountain in Sancheong of Gyeongsangnam-do. Remarks: This species is easily identified by the septa near the terminal parts of the valve and the sub- sequently characteristic outline.

Fig. 33. Distribution of Stauroneis prominula.

A B C

Fig. 34. Stauroneis prominula. A-C. The morphology of valves (Scale: 10 μm, ×2000). 38 Algal Flora of Korea·Freshwater Diatoms XI

18. ‌Stauroneis pseudosmithii Van de Vijver et Lange-Bertalot in

Van de Vijver et al. 2004: 58 (Figs. 35, 36).

Van de Vijver et al. 2004: 58. pl. 103. f. 1.

Valves narrow linear-lanceolate with triundulate margins, the ends of the valve narrow protracted and rostrate, and pseudosepta developed near the ends. Raphe filiform and the central ends of the raphe expanded. Axial area narrow and the stauros in the central area rectangular, extending to the margins of the valve. Striae slightly radiate to even parallel in the middle and more radiate towards, 20-21 in 10 μm. Valves 25-40 μm in length, 5-7 μm in breadth.

Type locality: Dovers Moraine in Heard Island, Subantarctica. Ecology and distribution: Stauroneis pseudosmithii occurs in sub-Antarctic regions (Van de Vijver et al. 2004). This species is very rare in a small pool of Baekyang Mountain in Busan and the peatland of Jangdo Island in Sinan of Jeollanam-do, and recently occurring in mosses of Daebong Reservoir in Chang- nyeong of Gyeongsangnam-do. Fig. 35. Distribution of Stauroneis pseudosmithii.

C D B E A

Fig. 36. Stauroneis pseudosmithii. A-D. The morphology of valves. D, E. The girdle (Scale: 10 μm, ×2000). Pennales: Naviculaceae: Stauroneis 39

19. ‌Stauroneis reichardtii Lange-Bertalot, Cavacini, Tagliaventi et Alfinito

2003: 142 (Figs. 37, 38).

Lange-Bertalot et al. 2003: 142. pl. 36. f. 1. Van de Vi- jver et al. 2004: 60. pl. 54. f. 6.

Valves linear to slightly linear-lanceolate with almost parallel margins, the ends of the valve abruptly protracted and distinctly subcapitate to capitate. Raphe slightly lateral in the middle, the central ends of the raphe straight, the central pores of the raphe indistinct. Axial area rather narrowly linear, narrow towards the central areas and terminal ends. The hyaline stauros of the central area rectangularly transverse, hardly expanded towards the margins. Striae weakly radiate in the middle and more radiate in the ends, 21-23 rows in 10 μm, and areolae unresolved in the light microscopy. Valves 22-48 μm in length and 6.2-11.5 μm in breadth.

Type locality: Pozza S’Arenarzu near Oristano of Sardinia Island, Italy. Ecology and distribution: This species occurs in tropical South America and in Sardinia of the Mediter- ranean Sea (Van de Vijver et al. 2004). It was found in Fig. 37. Distribution of Stauroneis reich- some mountain peatlands, Wangdeungjaeneup of Jiri ardtii. Mountain in Sancheong of Gyeongsangnam-do, Sand- eulneup of Jaeyak Mountain in Miryang, Yongneup of Daeam Mountain of Gangwon-do, particularly common in an unnamed peatland of Daegwanryeo- ng Pasture in Pyeongchang. Remarks: This species is similar closely to S. acidoclinatopsis Van de Vijver et Lange-Bertalot, The former has more lanceolate outline of the valve, the bigger dimensions and rather than the differ- ence in strial number (Van de Vijve et al. 2004). 40 Algal Flora of Korea·Freshwater Diatoms XI

A B C D

E F G H

Fig. 38. Stauroneis reichardtii. A-H. The morphology of valves (Scale: 10 μm, ×2000). Pennales: Naviculaceae: Stauroneis 41

20. ‌Stauroneis respectabilis Lange-Bertalot, Cavacini, Tagliaventi et Alfinito

2003: 144 (Figs. 39, 40).

Lange-Bertalot et al. 2003: 144. pl. 31. f. 3.

Valves absolutely lanceolate in outline, the ends of the valve not protracted in large froms and slightly protracted in small forms. Raphe distinctly lateral in the middle, the central ends of the raphe not curved and straight, the central pores of the raphe distinct. Axial area rather broadly linear. The hyaline stauros of the central area rectangularly transverse, more or less expanded towards the margins, and a few short striae occasionally occurring in the margins. Striae radiate in the middle and more radiate in the ends, 14- 16 rows in 10 μm, and puncta 14-16 in 10 μm. Valves 85-125 μm in length and 20-24 μm in breadth.

Type locality: Pauli Murdegu of Insula Sardinia, Italy. Ecology and distribution: This species was reported newly from Sardinia, the second largest island in the Mediterraean Sea, but has usually not been distinguished from Stauroneis phoenicenteron (Nitzsch)

Ehrenberg sensu lato (Lange-Bertalot et al. 2003). This Fig. 39. Distribution of Stauroneis respect- may occur more widespread over the freshwaters. In abilis. Korea, it was found rarely in a lowland wetland, Jinal- neup, in Haman of Gyeongsangnam-do, in peatlands of Sohwangbyeongsanneup of Odae Mountain and Daegwanryeong Pasture in Pyeongchang of Gangwon-do.

Remarks: This species is similar to some large forms of Stauroneis phoenicenteron (Nitzsch) Ehren- berg sensu stricto, but differ in the lacking ridge in the valve margins and central endings of the raphe (Lange-Bertalot et al. 2003). 42 Algal Flora of Korea·Freshwater Diatoms XI

Fig. 40. Stauroneis respectabilis. A-E. The morphology of valves (Scale: 10 μm, ×1500). Pennales: Naviculaceae: Stauroneis 43

21. Stauroneis sagitta Cleve 1881: 15 (Figs. 41, 42).

Hustedt 1959: 810. f. 1158. Van de Vijver et al. 2004: 63. pl. 101. f. 1.

Synonym: Stauroneis smithii var. sagitta (Cleve) Hustedt 1959: 811. f. 1158.

Valves elliptic-lanceolate to rhombic-lanceolate with convex margins, the margins of the valve weakly triundulate, the ends of the valve narrow protracted and rostrate, and pseudosepta developed near the ends. Raphe filiform and the central ends of the raphe expanded. Axial area narrow and the hyaline stauros in the central area long rectangular, extending to the margins of the valve. Striae parallel to slightly radiate throughout the valve, 19-21 rows in 10 μm, puncta on the striae unresolved in light microscopy. Valves 34- 49 μm in length, 8-11 μm in breadth.

Type locality: Finmark County in Gulf of Bothnia, Norway. Ecology and distribution: Stauroneis sagitta is found in the Arctic region and in slightly brackish wa- Stauroneis sagitta ters as type locality (Van de Vijver et al. 2004). This Fig. 41. Distribution of .

A B C

Fig. 42. Stauroneis sagitta. A-D. The morphology of valves (Scale: 10 μm, ×2000). 44 Algal Flora of Korea·Freshwater Diatoms XI species occurs only in a lake, Dongsong Reservoir in Cheolwon of Gangwon-do.

22. ‌Stauroneis siberica (Grunow) Lange-Bertalot et Krammer in

Lange-Bertalot and Metzeltin 1996: 104 (Figs. 43, 44).

Lange-Bertalot and Metzeltin 1996: 104. pl. 35. f. 1. Van de Vijver et al. 2004: 65. pl. 45. f. 1. Bahls 2011e in Diatoms of the United States.

Basionym: Stauroneis anceps var. siberica Grunow in Cleve and Grunow 1880: 48. pl. 3. f. 65.

Valves elliptic-lanceolate in outline, the ends of the valve protracted and rostrate to subcapitate. Raphe filiform and straight, the central ends of the raphe straight and the central pores indistinct. Axial area very narrow, linear and slightly narrow towards the central area. The hyaline stauros of the central area rectangular, narrow toward the margins and areolae bordering the stauros coarse. Transapical striae parallel or slightly convergent in the middle parts, weakly radiate towards the ends, and very fine, 24-32 rows in 10 μm. Valves 50-60 μm in length, 9-11 μm in breadth.

Type locality: Jenissey in Russia. Ecology and distribution: Stauroneis siberica oc- Fig. 43. Distribution of Stauroneis siberi- curs in oligodystrophic waters (Lange-Bertalot and ca. Metzeltin 1996) and slightly alkaline and low electrolyte lakes and pools in Arctic region (Van de Vijver et al. 2004). This species was reported in the Kwang River in Uljin Gyeongsangbuk-do (Lee et al. 1994), and was rare in a lowland wetland in Haman and in a pool of Odo Mountain in Hapcheon of Gyeongsangnam-do in this study. Remarks: Almost parallel arrangement of fine striae in middle parts is characteristic in this species. Pennales: Naviculaceae: Stauroneis 45

A B C

D E F

Fig. 44. Stauroneis siberica. A-F. The morphology of valves (Scale: 10 μm, ×2000). 46 Algal Flora of Korea·Freshwater Diatoms XI

23. Stauroneis smithii Grunow 1860: 564 (Figs. 45, 46).

Hustedt 1959: 810. f. 1157a-c.

Valves lanceolate with triundulate and convex margins, the ends of the valve narrow protracted and rostrate, and pseudosepta developed near the ends. Raphe filiform and straight, and the central ends of the raphe expanded. Axial area narrow and the hyaline stauros of the central area rectangular, extending to the margins of the valve. Striae parallel to slightly radiate throughout the valve, 25-30 rows in 10 μm, puncta on the striae unresolved in light microscopy. Valves 18-28 μm in length, 5.8-7.7 μm in breadth.

Type locality: In Austria. Ecology and distribution: Stauroneis smithii is widely distributed in freshwaters. This species occurred in Donghwa Stream in Daegu city (Hong and Chung 1990), and recently in mosses collected in the peatland of Jangdo Island in Sinan of Jeollanam-do.

Fig. 45. Distribution of Stauroneis smithii.

Figs. 46. Stauroneis smithii. The morphology of a valve (Scale: 10 μm, ×2000). Pennales: Naviculaceae: Stauroneis 47

24. ‌Stauroneis sonyae Kulikovskiy, Lange-Bertalot, Witkowski et

Dorofeyuk in Kulikovskiy et al. 2010: 62 (Figs. 47, 48).

Kulikovskiy et al. 2010: 62. pl. 93. f. 1. Bahl 2011f in Diatoms of the United States.

Valves lancolate to elliptic-lanceolate in outline, the ends of the valve not protracted and obtusely rounded. Raphe distinctly lateral, the central ends of the raphe hooked, the central pores of the raphe expanded and curved to the side. Axial area broad, slightly narrowing towards the central area. The hyaline stauros of the central area broadly rectangular, and expanded towards the margins with irregularly shortened striae in the margins. Striae radiate throughout, 14-16 rows in 10 μm on valves, areolae in a stria 15-18 puncta in

10 μm. Valves 120-170 (not 70-133) μm in length and

27-36 (not 18-25) μm in breadth.

Type locality: Sphagnum bog Nur in northern area of Hentai highlands, Mongolia. Ecology and distribution: This species was recorded newly in a Sphagnum bog, Nur, in Hentai highlands of Mongolia, and occurred in springs, lakes, rivers and other peat bogs in Mongolia (Kulikovskiy Fig. 47. Distribution of Stauroneis sonyae. et al. 2010). In Korea, it was found in some mountain peatlands, the peatland of Jangdo Island in Sinan of Jeollanam-do, unnamed peatlands of Daegwanryeong Pasture in Pyeongchang of Gangwon-do and a lowland wetland, Jilnalneup in Haman of Gyeongsangnam-do. Remarks: Morphological similarity exists in some large Stauroneis species. Stauroneis heinii differs by a narrower central area, less widening towards the margins and denser spaced striae in 10 μm, S. circumborealis Lange-Bertalot et Krammer by a broader axial area and a wide raphe, and S. medioasiatica Metzeltin et al. by its larger and broader valves and narrower ends (Kulikovskiy et al. 2010; Bahl 2011b). 48 Algal Flora of Korea·Freshwater Diatoms XI

A C D F

Fig. 48. Stauroneis sonyae. A-F. The morphology of valves (Scales: 10 μm, A, C, D, F: ×1000, B, E: ×1500). Pennales: Naviculaceae: Stauroneis 49

25. ‌Stauroneis staurolineata var. japonica Kobayasi et Ando 1978: 15 (Figs. 49, 50).

Kobayasi and Ando 1978: 15. pl. 2. f. 18.

Valves lanceolate in outline, the ends of the valve not protracted and acute. Raphe straight, the central ends of the raphe straight, the terminal ends of the raphe also straight. Axial area very narrow and the hyaline stauros of the central area rectangular extending up to the valve margins. Striae parallel throughout the valve, 15-16 rows in 10 μm, striae crossed regular- ly by many longitudinal lines. Valves 116-127 μm in length and 18-20 μm in breadth.

Type locality: Senjoga-ika pond in Saitama Prefec- ture, Japan. Ecology and distribution: This variety occurs in the type locality and a Sphagnum peatland near Lake

Biwa, Yamakado Moor (Kihara et al. 2009). In Korea, it occurs only in the 1100 wetland in Jeju Island. Remarks: This species was published invalidly in

1971 as Stauroneis stodderi var. japonica H. Kobayasi (Ando et al. 1971). Fig. 49. Distribution of Stauroneis staurolineata var. japonica. 50 Algal Flora of Korea·Freshwater Diatoms XI

A B C D

Fig. 50. Stauroneis staurolineata var. japonica. A-D. The morphology of valves (Scales: 10 μm, A-C ×2000, D: ×1500). Pennales: Naviculaceae: Stauroneis 51

26. ‌Stauroneis subaustralis Van de Vijver et Lange-Bertalot in

Van de Vijver et al. 2004: 69 (Figs. 51, 52).

Van de Vijver et al. 2004: 69. pl. 4. f. 1.

Valves typically lancolate in outline, the ends of the valve slightly protracted and bluntly rounded. Raphe broadly lateral, the central pores of the raphe expanded and curved or deflected to a side. Axial area very wide, linear and over 1/3 the bread of the valve. The hyaline stauros of the central area broadly transverse, a little expandedup tos the margins and sometimes short striae in the margins of the central area. Striae moderately radiate in the middle, strongly radiate towards the ends, 16 rows in 10 μm on valves, and areolae 16-18 in 10 μm of a stria. Valves 85-165 μm in length and 16.5-24 μm in breadth.

Type locality: Vallée de la Hébé in Ile de la Posses- sion, Crozet Archipelago of Subantarctica. Ecology and distribution: This species occurs on wet soil in drying mud pools of Antarctic region (Van de Vijver et al. 2004; Zidarova et al. 2014). In Korea, S. subaustralis was found only in mosses of wet area in a forest, Pyeongchang of Gangwon-do, and is newly re- Fig. 51. Distribution of Stauroneis sub- ported. australis. Remarks: This species is characterized by the wide axial area of the valve, and is compared with S. phoenicenteron (Nitzsch) Ehrenberg by narrower breadth of the valve, deflected central endings of the raphe (Van de Vijver et al. 2004). 52 Algal Flora of Korea·Freshwater Diatoms XI

C F B

D A

Fig. 52. Stauroneis subaustralis. A-F. The morphology of valves (Scales: 10 μm, A, D: ×1500, B, C, E, F: ×1200). Pennales: Naviculaceae: Stauroneis 53

27. ‌Stauroneis subgracilis Lange-Bertalot et Krammer in Lange-Bertalot and

Genkal 1999: 96 (Figs. 53, 54).

Lange-Bertalot and Genkal 1999: 96. pl. 29. f. 1. Van de Vijver et al. 2004: 71. pl. 27. f. 1.

Valves lanceolate to slightly elliptic-lanceolate in outline, the ends of the valve slightly protracted and subrostrate. Raphe lateral, the central ends of the raphe expanded, and the central pores slightly deflected. Axial area narrow and slightly narrow towards the central area and the terminal ends, the hyaline stauros of the central area rectangular and expanded towards the valve margins. Striae moderately radiate in the middle, more radiate towards the ends, 19-22 rows in 10 μm on valves, and areolae in a stria 18-20 puncta in 10 μm. Valves 46-76 μm in length and 12- 16 μm in breadth.

Type locality: Yugorskij Peninsula of northwestern Siberia. Ecology and distribution: This is known to be bipolar species in the Arctic and Antarctic regions, more abundant in the Arctica, and prefers freshwaters with a low to moderate electrolyte (Van de Vijver et al. Fig. 53. Distribution of Stauroneis sub- 2004). In Korea, it was found in mountain peatlands, gracilis. Danjoneup in Yangsan of Gyeongsangnam-do and Sumeunmulbaengdwineup in Jeju Island. Remarks: This species is most similar in morphology of S. pergracilis Van de Vijver et Lange-Ber- talot, but is distinguished by the lanceolate form of the valve rather than the linear to linear-lanceolate form (Van de Vijver et al. 2004). 54 Algal Flora of Korea·Freshwater Diatoms XI

A B C

D E F

Fig. 54. Stauroneis subgracilis. A-F. The morphology of valves (Scale: 10 μm, ×2000). Pennales: Naviculaceae: Craspedostauros 55

Genus Craspedostauros E.J. Cox 1999: 134.

Keu-ra-seu-pe-do-seu-ta-u-ro-seu-sok (크라스페도스타우로스속)

Valves linear or narrowly lanceolate in outline, often the margins of the valve constricted with the narrow stauros in the middle, the ends of the valve rounded. The face of the valve deeply convex, particularly towards the ends, the mantle of the valve wide. In girdle view, the cells generally constricted in the middle with many narrow girdle bands or copulae. Sometimes a clear bound- ary layer between the valve face and mantle. Occasionally the stauros biarcuate in the margins of the valve with a broader hyaline area. The raphe usually straight in a distinct raphe sternum, the ends of the raphe curved strongly and unilaterally, and the terminal ends of the internal fissures of the raphe fused into an additional silica knob with helictoglossae. The axial area narrow and linear, the hyaline and thickened stauros in central area narrowly and transversely developed up to the margins. The striae punctate, very delicate and invisible in light microscopy, but in the view of electron microscopy, the areolae on the valve cribriform with four to many small pores and similar pores along the girdle bands. In live cells, two plate-like chloroplasts arranged along apical axis, connected with a central pyrenoid, appressed against one side of the girdle and extending towards both valves. Holotype: Craspedostauros neoconstrictus E.J. Cox 1999.

Species: Seven species have been recorded in the database at present and are accepted as valid taxonomically (Guiry and Guiry 2015). Distribution: They are distributed in the brackish waters and are primarily benthic forms on sediments waters (Cox 1999; Lange and Tiffany 2002), but not frequent with sporadical fashions rather than cosmopolitan or widespread ones. Species of the genus Craspedostauros never occur with high abundance. But they adhere well to the hard substratum to establish biofilms or are fouling microorganisms in the marine environments (Higgins et al. 2002; Holland et al. 2004), and are particularly resistant to antifouling surfaces that inhibit the colonization of microorganisms (Chio- vitti et al. 2003). Craspedostauros australis E.J. Lox Cox, C. britannicus E.J. Lox Cox and C. decipiens

(Hustedt) E.J. Lox Cox are major foulers among them (Higgins et al. 2002; de Brouwer et al. 2006).

The species were found in Europe, Russia, South Africa, South America and Australia (Cox 1999; Lange-Bertalot and Genkal 1999; Rivera et al. 2011). They also occurred in the palaeolimnological records of Antarctica region, and sometimes with dominance (Verleyen et al. 2004; Hodgson et al. 2006).

Key reference: Cox (1999). Remarks: Diatoms belonging to this genus were previously placed in the genus Stauroneis Eh- renberg, but were separated from the previous group to be designated as genus Stauronella by

Mereschkowsky (1901). Although C. Mereschkowsky erected the genus Stauronella from Stauroneis constricta Ehrenberg 1843, he applied a misidentified taxon lacking stauros or other characters to the type specimen for the genus (Cox 1999). The name of genus Stauronella cannot be used for no- menclatural reasons. Genus Craspedostauros has some characteristic or different features from genus Stauroneis, the narrow stauros, constriction in the middle of frustule and cribriform areolae (Cox 1999). The genus Craspedostauros is more closely related to the family Mastogloiaceae than the family Stauroneidaceae (Cox and William 2000). 56 Algal Flora of Korea·Freshwater Diatoms XI

28. ‌Craspedostauros indubitabilis (Lange-Bertalot et Genkal) Cox 1999: 146 (Figs. 55, 56).

Lange-Bertalot and Genkal 1999: 100. pl. 36. f. 1. Rive- ra et al. 2011: 124. f. 1.

Basionym: Stauronella indubitabilis Lange-Bertalot et Genkal 1999: 100. pl. 36. f. 1.

Valves linear in outline, but linear-elliptical in small forms, the ends of the valve rounded to stumpi- ly cuneate, and not protracted. Raphe filiform and straight, the central ends of the raphe punctum-like, and the terminal ends of the raphe strongly curved in one direction. Axial area narrowly linear. The hyaline stauros of the central area thickened and narrow rectangular reaching the margins. Striae very fine and not resolved under light microscopy, parallel in the middle, more or less convergent towards the ends, and 25-27 rows in 10 μm. Valves 25-60 μm in length and 6-7 μm in breadth.

Type locality: Matveev Island near Yugorskij Pen- insula, northwestern Siberia of Russia. Ecology and distribution: Craspedostauros indu- Fig. 55. Distribution of Craspedostauros bitabilis is distributed probably in brackish waters in indubitabilis. the northern Hemisphere (Lange-Bertalot and Genkal 1999). This species occurred in the type locality of in northwestern Siberia in Russia (Lange-Bertalot and Genkal 1999), in the Red Sea near Hurghada in

Egypt (Gerasimyuk and Kovtun 2014), in the estuaries of the Belek River and the Balaklava Bay of the Crimean Peninsula in Ukraine (Nevrova 2013, 2014). This species was also found in a marine coast of Caldera in northern Chile (Rivera et al. 2011). It was reported as Stauronella indubitabilis Lange-Bertalot et Genkal in Russia, Egypt and Ukraine. In Korea, this species was found in the es- tuary of the Osip Stream in Yeongdeok of Gyeongsangbuk-do, and is newly reported. Pennales: Naviculaceae: Craspedostauros 57

A B E C D

Fig. 56. Craspedostauros indubitabilis. A-F. The morphology of valves (A-E. Scale: 10 μm, ×2000. F. The internal view of the valve. SEM, Scale: 5 μm). 58 Algal Flora of Korea·Freshwater Diatoms XI

Genus Caloneis Cleve 1894: 46 (3)

O-i-dol-mal-sok (오이돌말속)

29. Caloneis aequatorialis Hustedt 1921: 148 (Figs. 57, 58).

Simonsen 1987: 54. pl. 68. f. 14. Gotoh and Kubota 2010: 19. f. 96.

Valves lanceolate in outline, the margins of the valve slightly convex, the ends of the valve not protracted and rounded. Raphe straight or weakly curved, the central fissures of the raphe curved to one side and the terminal fissures of the raphe distinct. Axial area lanceolate and central area distinctly rectangular, forming hyaline fascia up to the margins of the valve. Striae 19-20 rows in 10 μm, slightly radiate in the middle and slightly convergent towards the ends. Valves 25-54 μm in length and 7-8.5 μm in breadth.

Type locality: East Africa. Ecology and distribution: Caloneis aequatorialis was found in the Jukskei River in South Africa (Taylor et al. 2005), in Lake Kinrin in Japan (Gotoh and Kubo- ta 2010) and in crater Lake Kyaninga in Uganda (Coc- quyt et al. 2010). In Korea, this species occurred as subaerial diatoms in mosses of Gageo Island in Sinan of Jeollanam-do, and is newly reported. Fig. 57. Distribution of Caloneis aequato- Remarks: This species is similar to C. molaris rialis. (Grunow) Krammer (Krammer and Lange-Bertalot 1986: 394. pl. 174. f. 16) in valve outline, strial number and arrangement and other characters, and it is actually hard to distinguish one from the other. However, C. aequatorialis has a distinct lanceolate outline and the convex margins of the valve in comparison with the linear-lancelate form of C. molaris. Pennales: Naviculaceae: Caloneis 59

A B C D E

F G H

Fig. 58. Caloneis aequatorialis. A-I. The morphology of valves (Scale: 10 μm, ×2000). 60 Algal Flora of Korea·Freshwater Diatoms XI

30. ‌Caloneis bryophila Manguin ex Kociolek and Reviers 1996: 206 (Figs. 59, 60).

Kociolek and Reviers 1996: 206. Moser et al. 1998: 25. pl. 1. f. 14.

Valves linear in outline, sometimes the margins of the valve slightly concave in the middle, the ends of the valve cuneately rounded to rounded. Raphe curved or slightly lateral, the central fissures of the raphe curved to one side and the terminal fissures of the raphe distinct. Axial area very wide and linear, and central area distinctly rectangular, forming hyaline fascia up to the margins of the valve. Striae 14- 15 rows in 10 μm, convergent in the middle and more radiate towards the ends. Valves 19-36 μm in length and 5.3-6 μm in breadth.

Type locality: New Caledonia, the east coast of Australia. Ecology and distribution: Caloneis bryophila seems to be reported only from the type locality. In Korea, this species occurred rarely as subaerial diatoms in mosses of Daebong Reservoir in Changnyeo- ng of Gyeongsangnam-do, and is newly reported. Fig. 59. Distribution of Caloneis bryophi- Remarks: This species is characterized by strial ar- la. rangement, the convergent striae in the middle parts and the strongly radial ones in the terminal parts of the valve.

Fig. 60. Caloneis bryophila. A, B. The morphology of valves (Scale: 10 μm, ×2000). Pennales: Naviculaceae: Caloneis 61

31. ‌Caloneis intermedia (Manguin ex Kociolek and Reviers) Moser,

Lange-Bertalot et Metzeltin 1998: 26 (Figs. 61, 62).

Kociolek and Reviers 1996: 207. Moser et al. 1998: 26. pl. 1. f. 14.

Basionym: Caloneis clevei var. intermedia Manguin ex Kociolek and Reviers 1996: 207.

Valves lanceolate in outline, and the ends of the valve protracted and broadly capitate. Raphe straight or weakly curved, the central fissures of the raphe curved to one side and the terminal fissures of the raphe distinct. Axial area lanceolate, and central area distinctly rectangular, widening up to the margins of the valve. Striae 22-26 rows in 10 μm, parallel to weakly radiate in the middle and parallel to weakly convergent in the ends of the valve. Valves 13-32 μm in length and 4.5-6 μm in breadth.

Type locality: Nouvelle-Calédonie in New Caledo- nia. Ecology and distribution: Caloneis intermedia was found in type locality of New Caledonia (Kociolek and Reviers 1996; Moser et al. 1998) and in Ratnagiri Fig. 61. Distribution of Caloneis interme- District of India (Dhumal and Sabale 2014). In Korea, dia. this species occurred rarely as subaerial or benthic diatoms in three localities, a lowland wetland, Jilnalne- up, in Haman, and Daebong Reservoir in Changnyeong of Gyeongsangnam-do, and Jangdo Island

B A C D E F G H I J K

Fig. 62. Caloneis intermedia. A. The girdle view. B-K. The morphology of valves (Scale: 10 μm, ×2000). 62 Algal Flora of Korea·Freshwater Diatoms XI

Bog Peatland in Sinan of Jeollanam-do. This is newly reported in Korea. Remarks: The dimension of the valves in the type locality was 28-31 μm in length and 5.5-6 μm in breadth (Kociolek and Reviers 1996), but in the local area, was wider in length of the valves, 13- 26 μm, and 4.5-5 μm in breadth.

32. ‌Caloneis odiosa (Manguin ex Kociolek and Reviers) Moser,

Lange-Bertalot et Metzeltin 1998: 26 (Figs. 63, 64).

Manguin 1962: 24. pl. 3. f. 7. Kociolek and Reviers 1996: 206. Moser et al. 1998: 26. pl. 1. f. 20.

Basionym: Pinnularia odiosa Manguin ex Kociolek and Reviers 1996: 206.

Valves more or less linear in outline, slightly convex in the middle, but in the large forms the margin of the valve inflated in the middle, and the ends of the valve obtuse to rounded. Raphe straight or weakly curved, the central fissures of the raphe slightly curved to one side and the terminal fissures of the raphe distinct. Axial area broadly linear-lanceolate, and central area distinctly rectangular, forming hyaline fascia up to the margins of the valve. Striae 16-20 rows in 10 μm, parallel to slightly radiate in the middle and slightly radial in the ends of the valve. Valves (29)32-34 (52) μm in length and 4.5-5.5 (8) μm in breadth.

Type locality: New Caledonia of the eastern Aus- tralia. Ecology and distribution: Caloneis odiosa was Fig. 63. Distribution of Caloneis odiosa. reported from the type locality, New Caledonia (Ko- ciolek and Reviers 1996; Moser et al. 1998) and lakes in Japan (Gotoh and Kubota 2010; Hirota et al. 2013). In Korea, this species occurred commonly as subaerial diatoms in mosses of Daebong Reservoir in Changnyeong, and in bottom of a lowland, Jilnalneup, in Haman of Gyeongsangnam-do. This is newly reported in Korea. Remarks: This species is linear in valve outline of small forms and is overlapped with C. molaris (Grunow) Krammer and C. aequatorialis Hustedt, especially in small forms. However, C. odiosa of large forms are characteristic by the inflation of the middle parts in the margins of the valve. The dimensions of the valves in the type locality accounted for only 32-34 μm in length and 4.5-5.5 μm in breadth to show narrow range, but 29-52 μm and 5.5-8 μm respectively in the local area of Ko- rea. Pennales: Naviculaceae: Caloneis 63

A B C D E

F G H

I J

Fig. 64. Caloneis odiosa. A-J. The morphology of valves (Scale: 10 μm, ×2000). 64 Algal Flora of Korea·Freshwater Diatoms XI

Genus Pinnularia Ehrenberg 1843: 45, nom. et typ. Cons. (3)

Bit-sal-dol-mal-sok (빗살돌말속)

33. Pinnularia anglica Krammer 1992: 109 (Figs. 65, 66).

Krammer 2000: 106. pl. 80. f. 7.

Valves linear in outline, the margins of the valve straight, the ends of the valve distinctly subcapitate. Raphe filiform as the outer fissures of the raphe lin- early straight, the central pores distinct and laterally bent, the terminal fissures of the raphe shaped like question marks. Axial area narrowly linear, the central area rhomboid expanding from the axial area, incompletely or asymmetrically developed as some marginal striae absent and irregular. Striae radiate in the middle, convergent at the ends, and 9-11 rows in 10 μm, No longitudinal lines on the margins of the valve across the striae. Valves 30-70 μm in length and 10-13 μm in breadth.

Type locality: Haverfordwest in England. Ecology and distribution: This species is common in moorland waters with low electrolyte and in oligotrophic waters of Europe (Krammer 2000). In Korea, Pinnularia anglica occurred abundantly in two mountain peatlands, the 1100 and Sumeunmulbangdwine- Fig. 65. Distribution of Pinnularia angli- up Bog on Halla Mountan of Jeju Island, and rarely ca. in other peatlands, Sinbulsanneup Bog, in Yangsan of Gyeongsangnam-do. This is newly reported to Korea. Remarks: The specimens collected in Korean freshwaters are closely related to P. anglica mor- photype 1 Krammer rather than P. angliciformis Van de Vijver et Beyens, with the primary reference of the valve breadth. Actually it is hard to differentiate P. anglica Krammer from P. angliciformis Van de Vijver et Beyens. Pennales: Naviculaceae: Pinnularia 65

B C D E

F G

Fig. 66. Pinnularia anglica. A-G. The morphology of valves (Scale: 10 μm, ×2000). 66 Algal Flora of Korea·Freshwater Diatoms XI

34. Pinnularia brebissonii (Kützing) Rabenhorst 1864: 222 (Figs. 67, 68).

Hustedt 1930: 321. f. 584. Krammer 2000: 69. pl. 45. f. 1.

Basionym: Navicula brebissonii Kützing 1844: 93. pl. 3. f. 49. Synonym: Pinnularia stauroneiformis W. Smith 1853: 57. pl. 19. f. 178.

Pinnularia microstauron var. brebissonii (Kützing) Mayer 1913: 186.

Pinnularia microstauron var. brebissonii (Kützing) Hustedt 1930: 321. f. 584.

Valves linear-lanceolate to linear elliptical in outline, the ends of the valve broadly rounded to weakly wedge-shaped. Raphe filiform to weakly lateral, central pores large round, the terminal ends of the raphe shaped indistinctly like question marks. Axial area narrow, but to be gradually wide towards the cental parts, uniting with the central area. Central area more or less broad fascia up to the margins, rarely short striae at the fascia margins, the central area uniting with the axial area forming rhombic shape. Striae slightly radiate in the middle, convergent towards the Fig. 67. Distribution of Pinnularia brebi- - μ - μ ends, and 10 12 rows in 10 m. Valves 28 60 m in ssonii. length and 9-12 μm in breadth.

Type locality: In süssem Wasser Falaise: De Brébisson in France. Ecology and distribution: Pinnularia brebissonii or P. microstauron var. brebissonii has been found in the oligotrophic and low electrolyte habitats (Hustedt 1930; Patrick and Reimer 1966), a mire in

North Norway (Foged 1978), streams of acid drainage area in Appalachians of the eastern USA(Kelly

1988), Shara Mountains in Macedonia (Levkov et al. 2005), Spahgnum bogs in Russia (Kulikovs- kiy 2009). On the contrary, this species prefers high electrolyte waters or brackish coastal waters (Krammer and Lange-Bertalot 1986; Krammer 2000), and was reported from the sand of Puck Bay coastal shallows in Poland (Witkowski 1991). In Korea, it occurred in streams of Mountain Gye- bangsan in Gangwon-do (Chung and Lee 1982), streams of Mountain Juwangsan and near Gyeo- ngju in Gyeongsangbuk-do (Chung and Watanabe 1984; Chung et al. 1985), and some lowland wetlands in Haman of Gyeongsangnam-do (Chung and Noh 1987). Pennales: Naviculaceae: Pinnularia 67

F E D C

Fig. 68. Pinnularia brebissonii. A-F. The morphology of valves (Scale: 10 μm, ×2000). 68 Algal Flora of Korea·Freshwater Diatoms XI

35. ‌Pinnularia brevicostata var. sumatrana Hustedt in Schmidt et al.

1934: 389 (Figs. 69, 70).

Schmidt et al. 1934. pl. 389. f. 7. Simonsen 1987: 156. pl. 251: 1. Krammer 2000: 146. pl. 125. f. 1.

Valves linear in outline, the margins of the valve parallel to slightly swollen in the middle, the ends of the valve broadly rounded. Raphe broadly lateral as outer fissures curved, the central ends of the raphe bent in one direction, central pores drop-shaped, the terminal ends of the raphe sickle-shaped. Axial area very wide, narrowing towards the ends, 3/4 the breadth of the valve, central area not developed independently. Striae parallel to slightly radiate in the middle, parallel to slightly convergent at the ends, and 7.5-9 rows in 10 μm. Longitudinal lines along the apical axis across the striae absent. Valves 77-130 μm in length and 17-19 μm in breadth.

Type locality: Moorland in Toba-Heide of Sumatra Island. Ecology and distribution: This variety was found in type locality of Sumatra (Simonsen 1987), in ponds of Japan (Negoro 1981; Murakami et al. 1988), in South Fig. 69. Distribution of Pinnularia brevi- Africa (Cholnoky 1962). In Korea, this species was costata var. sumatrana. found in a lowland wetland, Jilnalneup, in Haman of Gyeongsangnam-do, and is newly reported. Remarks: This variety is characterized by the swollen margins in the middle parts of the valve. Specimens collected from local area are 105-130 μm in length and 17-19 μm in breadth, and strial number is 7.5-8.5 in 10 μm. Pennales: Naviculaceae: Pinnularia 69

B C

Fig. 70. Pinnularia brevicostata var. sumatrana. A-E. The morphology of valves (Scale: 10 μm, A-E: ×1500). 70 Algal Flora of Korea·Freshwater Diatoms XI

36. ‌Pinnularia gibba var. cuneata Manguin ex Kociolek and Reviers

1996: 205 (Figs. 71, 72).

Manguin 1962: 23. pl. 4. f. 4. Kociolek and Reviers 1996: 205.

Valves lanceolate in outline, gradually tapering towards the rounded ends, the ends of the valve not protracted. Raphe lateral and narrow, the outer fissures of the raphe slightly curved, 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 question marks. Axial area widening from the ends towards the center to form a large rhombic hyaline area and a fascia up to the margins, and axial and central area not clearly differentiated. Usually four large markings distinct in the hyaline central area. Striae short in the margins of the valve, radiate in the middle, convergent towards the ends, and 9-10 rows in 10 μm. Longitudinal lines across the striae absent. Valves collected from the local area 57-76 μm in length and 9.5-10.5 μm in breadth.

Type: New Caledonia. Fig. 71. Distribution of Pinnularia gibba Ecology and distribution: This variety is report- var. cuneata. ed only in the type locality, New Caledonia. In Korea, this taxon occurred in the bottom of a lowland, Jilnal- neup, in Haman of Chungcheongnam-do. Pennales: Naviculaceae: Pinnularia 71

A B C D

E F G

Fig. 72. Pinnularia gibba var. cuneata. A-G. The morphology of valves (Scale: 10 μm, ×2000). 72 Algal Flora of Korea·Freshwater Diatoms XI

37. ‌Pinnularia meridiana Metzeltin et Krammer in Metzeltin and

Lange-Bertalot 1998 (Figs. 73, 74).

Metzeltin and Lange-Bertalot 1998: 180. pl. 181. f. 1.

Valves broadly linear in outline, the margins of the valve straight to slightly convex, and the ends of the valve very broadly protracted and rounded. Raphe slightly lateral, the central ends of the raphe bent, the central pores distinct drop-shaped, the terminal ends of the raphe indistinct in light microscopy. Axial area broadly lanceolate to rhombic, greatly wide towards the central area, not well defined from central area, and hyaline central area rhombic, but not up to the margins. Striae parallel or slightly radiate in the middle, slightly convergent towards the ends, and 9-11 rows in 10 μm. No longitudinal band crossing with striae. Valves 37-50 μm in length and 12-13 μm in breadth.

Type locality: Porto Alegre, Rio Grande do Sul in Brazil. Ecology and distribution: This species was found in streams of Porto Alegre and in the Amazon River of

Brazil (Metzeltin and Lange-Bertalot 1998), and in oth- Fig. 73. Distribution of Pinnularia meridi- er regions of Brazil (Souza and Senna 2009; Bartozek ana. et al. 2013). In Korea, this taxon occurred only from a lowland wetland, Jilnalneup, in Haman of Gyeong- sangnam-do, and is newly reported. Pennales: Naviculaceae: Pinnularia 73

A B C

D E

Fig. 74. Pinnularia meridiana. A-F. The morphology of valves (Scale: 10 μm, ×2000). 74 Algal Flora of Korea·Freshwater Diatoms XI

38. Pinnularia mesolepta (Ehremberg) W. Smith 1853: 58 (Figs. 75, 76).

Hustedt 1930: 319. f. 575a. Patrick and Reimer 1966: 600. pl. 55. f. 17.

Basionym: Navicula mesolepta Ehrenberg 1843: 419 (131). pl. 4/2. f. 4.

Valves linear in outline, the margin of the valve triundulate, the central hump narrow than the other two, the ends of the valve broadly rostrate to capitate. Raphe nearly straight, the central ends of the raphe bent in one direction and the central pores tear- shaped, the terminal ends of the raphe shaped like small question marks. Axial area linear to lanceolate, wide towards the central area, and central area rhom- boidal forming a fascia up to the margins. Striae radiate in the middle, convergent towards the ends, and 10-14 rows in 10 μm on valves. Valves 30-65 μm in length and 9-12 μm in breadth.

Type locality: Okak in Labrador, Canada. Ecology and distribution: This species is cosmopolitan worldwide in freshwaters (Hustedt 1930), Fig. 75. Distribution of Pinnularia meso- prefers the low electrolyte waters (Patrick and Reiemr lepta. 1966), and was dominant in Holocene lacustrine diatom records from Greenland (Cremer et al. 2001). In

Korea, this taxon occurs in lowland swamps (Chung and Noh 1987; Noh 1991), streams (Chung and

Watanabe 1984; Hong and Chung 1990; Lee and Chang 1991; Noh 1991), and reservoirs and lakes (Noh 1991; Choi et al. 1994). This was recently found in a mountain peatland, Yongneup, in Inje and a wet surface of a forest near Seonghwagsa Temple in Pyeongchang of Gangwon-do. Remarks: This taxon is similar to Pinnualria biceps Gregory and P. grunowii Krammer, but the former has more or less straight margins of the valve and the latter has narrower breadth of the valve, less 9 μm. Pennales: Naviculaceae: Pinnularia 75

A B C D E F

Fig. 76. Pinnularia mesolepta. A-F. The morphology of valves (Scale: 10 μm, ×2000). 76 Algal Flora of Korea·Freshwater Diatoms XI

39. Pinnularia nipponica Skvortzow 1936a: 45 (Figs. 77, 78).

Skvortzow 1936a: 45. pl. 7. f. 12. Kawashima and Mayama 2000: 77. f. 8E.

Valves broadly linear in outline, the margins of the valve straight, weakly convex or concave, and the ends of the valve narrowing, not protracted and broadly rounded. Raphe slightly lateral as the outer fissures of the raphe curved, the central ends of the raphe bent, the central pores drop-shaped, the terminal ends of the raphe shaped like question marks. Axial area linear to lanceolate, widening towards the central area, and hyaline central area rectangular reaching up to the margins, but somewhat irregular and asym- metrical along the apical axis. Striae slightly radiate in the middle, convergent towards the ends, and 8-9 rows in 10 μm. No longitudinal band crossing with striae. Valves 71-81.5 μm in length and 13.5-15 μm in breadth.

Type locality: Lake Kizaki in Shinano Province, Ja- pan. Ecology and distribution: This species occurred Fig. 77. Distribution of Pinnularia nip- only in some lakes in Japan, Lake Kizaki of Nagano ponica. Prefecture (Skvortzow 1936a), Lake Biwa of Shiga Pre- fecture (Skvortzow 1936b), and Lake Akan of Hokkai- do (Kawashima and Mayama 2000). In Korea, this taxon was found in Dongsong Reservoir in Che- olwon of Gangwon-do, and is newly reported. Remarks: Pinnularia nipponica is characterized by the robust striae on valves and the irregular forms of the central area. Pennales: Naviculaceae: Pinnularia 77

E C D

B A

Fig. 78. Pinnularia nipponica. A-E. The morphology of valves (Scale: 10 μm, ×2000). 78 Algal Flora of Korea·Freshwater Diatoms XI

40. ‌Pinnularia tirolensis (Metzeltin et Krammer) Krammer 2000: 88 (Figs. 79, 80).

Lange-Bertalot and Metzeltin 1996: 100. pl. 86. f. 7. Krammer 2000: 88. pl. 85. f. 2.

Basionym: Pinnularia subgibba var. tirolensis Metzeltin et Krammer in Lange-Bertalot and Metzeltin 1996: 100. pl. 86. f. 7.

Valves linear in outline, the margins of the valve straight to slightly convex, and the ends of the valve narrowing, not protracted and broadly rounded. Ra- phe slightly lateral, the central ends of the raphe bent, the central pores drop-shaped, the terminal ends of the raphe shaped like question marks. Axial area broadly linear to lanceolate, 1/4-1/2 the breadth of the valve, and hyaline central area rectangular up to the margins. Striae radiate in the middle, convergent towards the ends, and 9-10 rows in 10 μm. No longitudinal band crossing with striae. Valves 41-66 (82)

μm in length and 6.4-9.4 (10) μm in breadth.

Type locality: Lake Mittersee near Lermoos in Ti- rol, Austria. Ecology and distribution: Pinnularia tirolensis was found in oligotrophic and low electrolyte lakes such as Lake Mittersee in Austria and some lakes in the Alps Fig. 79. Distribution of Pinnularia tirolen- in Europe (Krammer 2000). This species occurred in sis. acidic conditions, in Shara Mountains in Mecedonia (Levkov et al. 2005), in sphagnum bogs in European

Russia (Kulikovskiy 2009) and in montain wetlands in Manchuria of China (Zhang et al. 2011). In addition, they were found in some fossil diatoms (Lotter and Hofmann 2003; Denys 2009). In Ko- rea, this taxon occurred from an unnamed peatland of Daegwanryeong Pasture and Sohwangbyeo- ngsanneup Peatland of Odae Mountain in Pyeongchang of Gangwon-do. This is newly reported to Korea. Remarks: The local specimens collected in Korea are larger than those in the type locality, 59-82 μm in length of the valves and 9.5-10 μm in breadth. Pennales: Naviculaceae: Pinnularia 79

A B C D E

Fig. 80. Pinnularia tirolensis. A-G. The morphology of valves (Scale: 10 μm, ×2000). 80 Algal Flora of Korea·Freshwater Diatoms XI

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Index to Korean Names

ㅂ O

빗살돌말속 64 오이돌말속 58

ㅅ ㅋ

십자돌말속 7 크라스페도스타우로스속 55 Index to Scientific Names 87

Index to Korean Name as Pronounced

B O

Bit-sal-dol-mal-sok 64 O-i-dol-mal-sok 58

K S

Keu-ra-seu-pe-do-seu-ta-u-ro-seu-sok 55 Sip-ja-dol-mal-sok 7 88 Algal Flora of Korea·Freshwater Diatoms XI

Index to Scientific Names

B Pinnularia biceps 74 Pinnularia anglica 64, 65 Bacillaria phoenicenteron 7 Pinnularia angliciformis 64 Pinnularia brebissonii 66, 67 Pinnularia brevicostata var. sumatrana 68, 69 C Pinnularia gibba var. cuneata 70, 71 Pinnularia grunowii 74 Caloneis 58 Pinnularia meridiana 72, 73 Caloneis aequatorialis 58, 59, 62 Pinnularia mesolepta 74, 75 Caloneis bryophila 60 Pinnularia microstauron var. brebissonii 66 Caloneis clevei var. intermedia 61 Pinnularia nipponica 76, 77 Caloneis intermedia 61 Pinnularia odiosa 62 Caloneis molaris 58, 62 Pinnularia stauroneiformis 66 Caloneis odiosa 62, 63 Pinnularia subgibba var. tirolensis 78 Craspedostauros 5, 55 Pinnularia tirolensis 78, 79 Craspedostauros australis 55 Pinus densiflora 6

Craspedostauros britannicus 55 Pleurostauron (Pleurostaurum) prominulum 35 Craspedostauros decipiens 55 Proschkinia 5 Craspedostauros indubitabilis 56, 57 Craspedostauros neoconstrictus 55 Q

H Quercus serrata 6

Haslea 5 S

M Stauroneis 5, 6, 7, 18, 22, 25, 47, 55 Stauroneis accedens 15 Molinia japonica 6 Stauroneis acidoclinatopsis 39 Stauroneis acuta 8, 9, 15 Stauroneis alabamae 10, 11 N Stauroneis amphicephala 12, 13, 15 Stauroneis anceps 5, 14, 15, 27 Navicula brebissonii 66 Stauroneis anceps f. gracilis 22 Navicula mesolepta 74 Stauroneis anceps var. amphicephala 12 Nupela 24 Stauroneis anceps var. gracilis 22 Nupela gracillima 24 Stauroneis anceps var. siberica 44 Stauroneis ancepsopsis 15, 16 Stauroneis borgei 17 P Stauroneis circumborealis 18, 19, 25, 47 Stauroneis constricta 55 Pinnualria 64 Stauroneis elena 20, 21 Index to Scientific Names 89

Stauroneis gracilis 22, 23, 25, 31 Stauroneis sagitta 43 Stauroneis gracillima 24 Stauroneis siberica 44, 45 Stauroneis heinii 25, 47 Stauroneis smithii 46 Stauroneis ignorata 35 Stauroneis smithii var. borgei 17 Stauroneis jarensis 27, 28 Stauroneis smithii var. sagitta 43 Stauroneis kriegeri 29, 30 Stauroneis sonyae 25, 47, 48 Stauroneis kuelbsii 31, 32 Stauroneis staurolineata var. japonica 49, 50 Stauroneis lacusvulcani 33, 34 Stauroneis stodderi var. japonica 49 Stauroneis medioasiatica 47 Stauroneis subaustralis 51, 52 Stauroneis nobilis 35, 36 Stauroneis subgracilis 25, 27, 31, 53, 54 Stauroneis parvula var. prominula 35 Stauroneis supergracilis 22 Stauroneis pergracilis 53 Stauronella 5, 55 Stauroneis phoenicenteron 5, 7, 22, 25, 41, 51 Stauronella indubitabilis 56 Stauroneis prominula 35, 37 Sphagnum 20, 47, 66 Stauroneis pseudosmithii 38 Staurophora 5 Stauroneis pygmaea 29 Stauropsis 5 Stauroneis reichardtii 39, 40 Stauroneis amphicephaloides 12 Stauroneis respectabilis 41, 42 Stauroneis gracilior 33 Stauroneis rex 25