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Diatoms of Bering Island, Kamchatka, Russia

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DOI: 10.1127/1438-9134/2014/004

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C Nova Hedwigia, Beiheft 143, p. 63–102
Stuttgart, July 2014

Diatoms of Bering Island, Kamchatka, Russia

Marina Potapova
The Academy of Natural Sciences of Drexel University, 1900 Benjamin Franklin Parkway, Philadel-
phia, PA 19103, USA; Potapova@ansp.org

With 401 figures and 1 table

Abstract: The goal of this study was an inventory of diatoms of Bering Island, the largest of the Commander
Islands located east of Kamchatka Peninsula, Russia. Diatom samples were collected in summer 2008
from 85 sites, mostly from ponds, small lakes, rivers, and streams. A total of 313 species and infraspecific
diatom taxa were found. The diatom flora of the island was mostly comprised of species characteristic for
northern and alpine regions of Eurasia and North America, but several species were so far found only in
Asia, or North America, or in both Eastern Asia and North America, were also observed. Five new species,
Eunotia ninae Potapova, sp. nov., E. prilezhaevii Potapova, sp. nov., Diadesmis mochalovae Potapova, sp.
nov., Psammothidium strelnikovae Potapova, sp. nov., and Pinnularia beringensis Potapova, sp. nov., are
described and a new combination Placoneis cuneata (M. Moller ex Foged) Potapova is established. This
study provides information for future analysis of biogeograhical patterns of diatoms and may serve as a
baseline for assessing the changes in diatom assemblages caused by climate change and human impacts.

Key words: Bering Island, Commander Islands, diatoms, Kamchatka, new species, tundra ponds.

Introduction
Bering Island is the largest in the group of the Commander Islands located 175 kilometers east of
the Kamchatka Peninsula, Russia (Fig. 1). These islands belong to the Aleutian volcanic island
arc, which separates Bering Sea from the Pacific Ocean. Bering Island is 95 kilometers long
and 15 kilometers wide, with an area of 1,667 square kilometers. The climate of the island is
maritime subarctic with mild winters and cold foggy summers. The average annual temperature
is + 2.1oC and mean annual precipitation 500 mm (Kursanova & Savchenko 1966). The northern
part of the islands consists of the plateaus and coastal plain, while the southern part is covered
by low mountains. The Island is treeless, covered by tundra vegetation. Shallow lakes and nival
ponds are a common landscape feature. Small ponds usually have peaty bottoms and abundant
aquatic plants growth. The common plant species in lakes and ponds are Hippuris vulgaris
L., Sparganium hyperboreum Beurling ex Laest., Batrachium trichophyllum (Chaix) Van der
Bosche, Callitriche palustris L., and Eleocharis palustris (L.) Roem. & Schult. (Mochalova &
Yakubov 2004).
The oceanic island chains are considered natural laboratories that allow studying various
aspects of biogeography. Comparisons of the diatom floras on island chains may shed light
on the mechanisms that generate diatom diversity, such as dispersal, extinction and speciation
and thus further the understanding of microbial biogeography. One of the first observations on
diatom biogeography, for instance, was that of considerable similarity between diatom floras of
Siberia and Western North America expressed by Ehrenberg (1949). Still, no formal analysis has
yet been possible to quantitatively assess this similarity or reveal other biogeographical patterns
in the area as there is no sufficient and well documented floristic data available. While diatoms of
the American Northwest are being currently systematically studied at modern level (e.g., Bahls
2005, 2010, 2011, 2012), and diatoms of at least some parts of Alaska have been documented by
LM photographs (Foged 1971, 1981), there are few modern well-documented floristic studies on
diatoms of Alaska, North-Eastern Asia (Kamchatka and Chukotka Peninsulas), and island chains
between Asia and North America.
© 2014 J. Cramer in der Gebr. Borntraeger Verlagsbuchhandlung, Stuttgart, Germany www.borntraeger-cramer.de
DOI 10.1127/1438-9134/2014/004 1438-9134/2014/004 $ 10.00
64 M. Potapova

Fig. 1. Map showing location of Bering Island and sampling sites.

Diatoms of Bering Island, Kamchatka, Russia 65

The diatoms of the Commander Islands have not been studied with the exception of one
species, Stephanodiscus niagarae Ehrenberg, reported from the plankton of Lake Sarannoe,
which is the largest lake on the Bering Island (Lepskaya 2000, Genkal & Lepskaya 2009).
On Aleutian Islands, the diatoms were surveyed on Adak Island (Judson 1946, Hein 1990)
and on Amchitka Island (Skuja 1969). Only one of these publications (Hein 1990) contains
photographic documentation of observed diatom species. Numerous studies dealt with recent
freshwater diatoms of the Kamchatka Peninsula, with majority dedicated to communities in
thermal springs (Petersen 1946, Golovenkina 1981) or lake plankton (e.g., Genkal & Lupikina
1998, Genkal et al. 2004, Kulikovskyi & Shkurina 2009, Lepskaya 2004, 2007, Lepskaya &
Lupikina 2007, Lupikina 2005, Shkurina et al. 2005). Nevertheless, the diversity of freshwater
diatoms in Kamchatka is still insufficiently known and new species are discovered in the course
of various research projects (e.g. Hoff et al. 2011). The goal of this study was to describe diatom
flora of Bering Island and document species occurrence by microphotographs.

Materials and methods

Diatom samples used in this study were collected July 8–23, 2008 from 85 sampling sites (Fig. 1) on Bering
Island. Fifty-six samples were collected from lakes and ponds and twenty-nine from rivers and streams.
Conductivity and pH were measured in 59 sites with ExStikII conductivity/pH meter. Conductivity ranged
from 4 to 415 S/cm, with the average value of 62 S/cm in lakes and ponds and 177 S/cm in streams and
rivers. pH varied from 5.2 to 9.6 with the average value of 6.1 in lakes and ponds and 8.0 in streams and
rivers. Two sites were located near the seashore and were obviously brackish, but no conductivity and pH
measurements were taken at these sites. Diatom samples were collected by squeezing mosses and aquatic
plants, scraping rocks, or siphoning the upper layer of silty sediments.
The samples were treated by 30 % hydrogen peroxide and potassium dichromate, then rinsed several
times with distilled water. The permanent diatom slides were prepared with Naphrax mounting medium.
The slides were examined with a Zeiss AxioImager A1 light microscope equipped with an AxioScope MRm
digital camera. For SEM examination diatom slurries were dried on aluminum stubs, sputter-coated with
Pt-Pd and observed with Zeiss Supra 50 scanning electron microscope under 10 kV accelerating voltage.

Results
A total of 313 species and infraspecific diatom taxa were found in studied samples (Table 1).
The most common diatoms in slightly acidic and soft-water tundra ponds were Aulacoseira
nivalis, Stauroforma exiguiformis, and various species of Frustulia, Eunotia, and Pinnularia. In
rivers that mostly had slightly alkaline water, dominant species were Planothidium lanceolatum,
P. haynaldii, Hannaea arcus, and Diatoma mesodon. In a few localities that were obviously
influenced by the influx of marine waters, Melosira moniliformis, Masogloia elliptica,
Ctenophora pulchella, and Pseudostaurosira subsalina were most abundant.

Class Coscinodiscophyceae
The genus Aulacoseira was represented by six species. Among those, A. nivalis (Figs 2,
9–11) was especially abundant in shallow tundra ponds. A. nivaloides (Figs 3, 4, 13, 14), A.
crassipunctata (Figs 5, 6) and A. italica were also frequently found in ponds. A. subarctica (Fig.
12) was abundant in the large lake Sarannoe. Two species of Melosira were abundant in some
rivers (M. varians, Fig. 15) and in brackish water localities (M. moniliformis, Figs 16, 17).
In the plankton of the lake Sarannoe two species of Stephanodiscus were abundant. The
small-sized species (Figs 18–28) is tentatively identified here as S. cf. binatus. This diatom
belongs to the group of relatively small-sized Stephanodiscus species with lacunate/scutate
valves, one marginal rimoportula, and a ring of marginal spines located at each interfascicle.
Similarly to S. binatus and S. minutulus and unlike S. nipigonensis Kling & Håkansson, it has
a single fultoportula located slightly off valve center (Fig. 27). Unlike S. minutulus that mostly
66 M. Potapova

Figs 2–17. Aulacoseira and Melosira species. Figs 2 (LM), 9–11 (SEM). Aulacoseira nivalis. Figs 3, 4
(LM), 13, 14 (SEM). A. nivaloides. Figs 5, 6 (LM). A. crassipunctata. Figs 7, 8 (LM). A. alpigena. Fig. 12
(SEM). A. subarctica. Fig. 15 (SEM) Melosira varians. Figs 16 (LM), 17 (SEM). M. moniliformis. Scale
bar for LM images = 10 μm, SEM images = 2 μm.

has been reported to have three-strutted fultoportulae, this diatom has all fultoportulae with two
satellite pores (Figs 27, 28). Håkansson & Kling (1990) noted that the most striking feature of S.
binatus is the long external tube of the rimoportula that is paired with a spine. The same feature
was observed in the specimens from the lake Sarannoe (Fig. 24). The reason that I was not
entirely convinced that this diatom is conspecific with S. binatus, was an unusual structure of its
external valve surface. It appeared to be often covered by an additional silica layer that created
Diatoms of Bering Island, Kamchatka, Russia 67

Figs 18–31. Stephanodiscus species. Figs 18–23 (LM) and 24–28 (SEM). S. cf. binatus. Figs 29–31.
S. cf. niagarae. Scale bar for LM images = 10 μm, SEM images = 2 μm in Figs 24–27, 0.5 μm in Fig.
28, 10 μm in Figs 29–31.
68 M. Potapova

a stellar pattern either in the valve central part (Fig. 24) or towards the periphery (Fig. 26). The
lacunes forming this stellar pattern gave an appearance of uniseriate striae in the SEM images
of the external valve surfaces, while the internal views of the valves always showed that striae
become bi- and multiseriate in their mid-course (Fig. 27).
The second, large-celled species of Stephanodiscus (Figs 29–31) has been described in detail
by Genkal & Lepskaya (2009). This diatom, found in the lake Sarannoe, has all characteristics
of S. niagarae, but lacks any central fultoportulae and has marginal spines that range in shape
from valve to valve from bifurcate (Fig. 30) to simple conical (Fig. 31). Genkal & Lepskaya
(2009) concluded that it belongs to S. niagarae because some populations of S. niagarae have
been reported in the past to lack central fultoportulae and there is evidence that spines may
vary in shape within a species. On the other hand, the bifurcate spines on some valves of the
Stephanodiscus from lake Sarannoe are so unusual, that I cannot be entirely sure that this is not
a separate species, so it is listed here as S. cf. niagarae.

Table 1. List of diatom taxa found on Bering Island in 2008.

Taxon Figures
Achnanthidium exiguum (Grunow) Czarnecki
Achnanthidium kranzii (Lange-Bertalot) Round & Bukhtiyarova 348, 349
Achnanthidium minutissimum (Kützing) Czarnecki
Actinella punctata F.W. Lewis
Adlafia bryophila (Petersen) Lange-Bertalot 215
Adlafia minuscula (Grunow) Lange-Bertalot 231
Amphora pediculus (Kützing) Grunow in Schmidt et al.
Aneumastus tuscula (Ehrenberg) Mann & Stickle
Asterionella ralfsii W. Smith 52, 53
Aulacoseira alpigena (Grunow) Krammer 7, 8
Aulacoseira ambigua (Grunow) Simonsen
Aulacoseira crassipunctata Krammer 5, 6
Aulacoseira italica (Ehrenberg) Simonsen
Aulacoseira nivalis (Smith) English & Potapova 2, 9–11
Aulacoseira nivaloides (Camburn) English & Potapova 3, 4, 13, 14
Aulacoseira subarctica (O. Müller) Haworth 12
Boreozonacola hustedtii Lange-Bertalot et al. 243
Boreozonacola natchikae (Petersen) Lange-Bertalot et al. 242
Brachysira brebissonii Ross
Brachysira microcephala (Grunow) P. Compère
Brachysira neoacuta Lange-Bertalot
Caloneis bacillum (Grunow) Cleve 262
Caloneis silicula (Ehrenberg) Cleve 260
Caloneis sp. 1 266
Caloneis sp. 2 267
Caloneis undulata (Gregory) Krammer 261
Cavinula cocconeiformis (Gregory ex Greville) Mann & Stickle 211, 212
Cavinula jaernefeltii (Hustedt) Mann & Stickle 210
Diatoms of Bering Island, Kamchatka, Russia 69

Taxon Figures
Cavinula pseudoscutiformis (Hustedt) Mann & Stickle 213
Cavinula variostriata (Krasske) Mann 214
Chamaepinnularia cf. begeri (Krasske) Lange-Bertalot 191, 192
Chamaepinnularia cf. krookiformis (Krammer) Lange-Bertalot & Krammer 187–190, 201, 202
Chamaepinnularia circumborealis Lange-Bertalot 182, 183
Chamaepinnularia krookiformis (Krammer) Lange-Bertalot & Krammer 184–186, 200, 203
Chamaepinnularia mediocris (Krasske) Lange-Bertalot 193, 197
Chamaepinnularia soehrensis (Krasske) Lange-Bertalot & Krammer 196
Chamaepinnularia soehrensis var. capitata (Krasske) Lange-Bertalot &
194, 195, 199
Krammer
Chamaepinnularia sp. 1 206
Chamaepinnularia vyvermanii Lange-Bertalot 198, 204, 205
Cocconeis placentula Ehrenberg
Cosmioneis pusilla (W. Smith) Mann & Stickle
Ctenophora pulchella (Ralfs ex Kuetzing) Williams & Round 54
Cyclotella antiqua W. Smith
Cyclotella meneghiniana Kützing
Cymbella amplificata Krammer 311
Cymbella mexicana var. kamtschatica (Grunow) Krammer 310
Cymbella neocistula Krammer 312
Cymbella proxima Reimer 313
Cymbella sp. 1 316
Cymbellonitzschia diluviana Hustedt
Cymbopleura crassipunctata var. nonornata Krammer 309
Cymbopleura cuspidata (Kutzing) Krammer
Cymbopleura peranglica Krammer 317
Decussata placenta (Ehrenberg) Lange-Bertalot
Diadesmis contenta (Grunow ex Van Heurck) Mann 226
Diadesmis laevissima (Cleve) Mann 227
Diadesmis mochalovae sp. nov. 219–224, 228–230
Diadesmis perpusilla (Grunow) Mann 225
Diatoma anceps (Ehrenberg) Kirchner 44
Diatoma hiemalis (Lyngbye) Heiberg 42
Diatoma mesodon (Ehrenberg) Kützing 43
Diatoma moniliformis Kützing 45
Diatoma tenuis Agardh
Diatomella balfouriana Greville 268, 269
Didymosphenia geminata (Lyngbye) M. Schmidt
Diploneis elliptica (Kützing) Cleve 251
Diploneis ovalis (Hilse) Cleve
Diploneis pseudovalis Hustedt 252
70 M. Potapova

Taxon Figures
Discostella pseudostelligera (Hustedt) Houk & Klee
Encyonema cf. gauemannii (Meister) Krammer 325
Encyonema hebridicum Grunow ex Cleve 322
Encyonema kamtschaticum Krammer 321
Encyonema minutum Hilse
Encyonema neogracile Krammer
Encyonema perpusillum (A. Cleve) Mann 323
Encyonema reichardtii (Krammer) Mann 340
Encyonema rostratum Krammer 324
Encyonema silesiacum (Bleisch) Mann
Encyonema temperei Krammer 315
Encyonopsis behrei (Foged) Krammer & Metzeltin
Encyonopsis descripta (Hustedt) Krammer 335
Encyonopsis sp.1 326
Eolimna minima (Grunow) Lange-Bertalot
Epithemia adnata (Kützing) Brébisson
Epithemia sorex Kützing
Epithemia turgida (Ehrenberg) Kützing
Eunotia bidens Ehrenberg 136
Eunotia bilunaris (Ehrenberg) Mills 129
Eunotia cristagallii Cleve 119
Eunotia diadema Ehrenberg 137
Eunotia elegans Ostrup
Eunotia fallax A. Cleve 101, 102
Eunotia iatriaensis Foged 126
Eunotia incisa Gregory 115
Eunotia lapponica Grunow ex A. Cleve 130
Eunotia meisteroides Lange-Bertalot 122
Eunotia microcephala Krasske ex Hustedt 123, 124
Eunotia minor (Kützing) Grunow 104–106
Eunotia minuitula Grunow 127, 128
Eunotia mucophila (Lange-Bertalot & Nörpel-Schempp) Lange-Bertalot 107
Eunotia neocompacta Mayama 108
99–100, 109–113,
Eunotia ninae sp. nov.
146–148
Eunotia paludosa Grunow 114
Eunotia parallela Ehrenberg
Eunotia praerupta Ehrenberg 145
Eunotia prilezhaevii sp. nov. 131–135, 149–151
Eunotia rhomboidea Hustedt 116, 117
Eunotia subarcuatoides Alles et al. 125
Diatoms of Bering Island, Kamchatka, Russia 71

Taxon Figures
Eunotia ursamajoris Lange-Bertalot & Nörpel-Schempp 118
Eunotia paratridentula Lange-Bertalot & Kulikovskiy 120, 121
Fragilaria acidoclinata Lange-Bertalot & Hofmann 41
Fragilaria atomus Hustedt 82
Fragilaria bidens Heiberg 48
Fragilaria cassubica Witkowski & Lange-Bertalot 77, 78, 83
Fragilaria cf. vaucheriae (Kützing) Petersen 49
Fragilaria gracilis Østrup 50,51
Fragilaria neoproducta Lange-Bertalot 63
Fragilaria sp. 1 57
Fragilaria vaucheriae (Kützing) Petersen 46, 47
Fragilariforma bicapitata (A. Mayer) Williams & Round 58, 84, 85
Fragilariforma constricta (Ehrenberg) Williams & Round 64–66
Fragilariforma lata (A. Cleve-Euler)) Williams & Round 62
Frustulia crassinervia (Brebisson) Lange-Bertalot & Krammer 241
Frustulia inculta Siver et al. 250
Frustulia krammeri Lange-Bertalot & Metzeltin
Frustulia lange-bertalotii Metzeltin 248
Frustulia saxonica Rabenhorst 249
Frustulia vulgaris (Thwaites) De Toni
Geissleria acceptata (Hustedt) Lange-Bertalot & Metzeltin
Gomphoneis eriense (Grunow) Skvortzow & Meyer 320
Gomphonema acuminatum var. pusillum Grunow 328
Gomphonema cf. mexicanum Grunow 347
Gomphonema cf. olivaceum var. minutissimum Hustedt 337, 346
Gomphonema cf. pumilum var. rigidum Reichardt & Lange-Bertalot 342
Gomphonema gracile Ehrenberg 330
Gomphonema interpositum Reichardt 336
Gomphonema khentiiense Lange-Bertalot et al. 329
Gomphonema micropus Kutzing 333
Gomphonema olivaceoides Hustedt 327
Gomphonema olivaceum (Hornemann) Ehrenberg 332
Gomphonema pala Reichardt 319
Gomphonema sp. 1 341
Gomphonema sp. 2 343, 345
Gomphosphenia sp. 1 341
Gomphosphenia tackei (Hustedt) Lange-Bertalot 344
Halamphora cf. holsatica (Hustedt) Levkov 314
Hannaea arcus (Ehrenberg) Patrick 60
Hannaea arcus f. recta (Cleve) Foged 59
Hantzschia amphioxys (Ehrenberg) Grunow
72 M. Potapova

Taxon Figures
Hippodonta capitata (Ehrenberg) Lange-Bertalotet al.
Hippodonta subcostulata (Hustedt) Lange-Bertalot et al. 308
Hygropetra balfouriana (Grunow ex Cleve) Krammer & Lange-Bertalot 180, 181
Karayevia clevei (Grunow in Cleve & Grunow) Round & Bukhtiyarova
Karayevia laterostrata (Hustedt) Round & Bukhtiyarova
Karayevia suchlandtii (Hustedt) Bukhtiyarova
Kobayasiella parasubtilissima (Kobayasi & Nagumo) Lange-Bertalot 255
Kobayasiella subtilissima (Cleve) Lange-Bertalot 256
Mastogloia elliptica (Agardh) Cleve 253, 254
Mayamaea permitis (Hustedt) Bruder & Medlin
Melosira moniliformis (O. F. Muller) Agardh 16, 17
Melosira varians Agardh 15
Meridion circulare (Greville) Agardh 61
Meridion circulare var. constrictum (Ralfs) Van Heurck
Microcostatus cf. naumannii (Hustedt) Lange-Bertalot
Microcostatus cf. maceria (Schimanski) Lange-Bertalot 232
Navicula aurora Sovereign 282, 283
Navicula cf. antonii Lange-Bertalot 292
Navicula cf. arctotenelloides Lange-Bertalot & Metzeltin 300,301
Navicula cf. exilis Kützing 302
Navicula cf. hoffmanniae Pantocsek 296
Navicula cf. microcari Lange-Bertalot 304
Navicula cf. subalpina Reichardt (as 42:2 in ID 20 299
Navicula cf. veneta Kützing 293–295
Navicula cryptotenella Lange-Bertalot ex Krammer & Lange-Bertalot 297
Navicula cryptotenelloides Lange-Bertalot 303
Navicula detenta Hustedt 305
Navicula difficillima Hustedt 233
Navicula digitulus Hustedt 306
Navicula gregaria Donkin 291
Navicula leptostriata Jørgensen 1948 290
Navicula peregrina (Ehrenberg) Kützing 280
Navicula radiosa Kützing 281
Navicula salinarum Grunow 289
Navicula schmassmannii Hustedt
Navicula streckerae Lange-Bertalot & Witkowski 298
Navicula vaneei Lange-Bertalot 286–288
Navicula vulpina Kützing 285
Neidium alpinum Hustedt 263, 264
Neidium amphigomphus (Ehrenberg) Pfitzer 278
Neidium ampliatum (Ehrenberg) Krammer 259
Diatoms of Bering Island, Kamchatka, Russia 73

Taxon Figures
Neidium bisulcatum (Lagerstedt) Cleve 265
Neidium cf. alpinum Hustedt 272
Neidium sacoense Reimer 258
Neidium sp. 1 257
Neidium sp. 2 277
Nitzschia acidoclinata Lange-Bertalot 389–392
Nitzschia angustata (W. Smith) Grunow 379
Nitzschia angusteforaminata Lange-Bertalot 385, 386
Nitzschia bryophila (Hustedt) Hustedt 400, 401
Nitzschia cf. homburgiensis Lange-Bertalot 393
Nitzschia cf. paleacea Grunow 397
Nitzschia cf. solita Hustedt 398
Nitzschia cf. sublinearis Hustedt 380, 381
Nitzschia dissipata var. media (Hantzsch) Grunow 399
Nitzschia fossilis Grunow 384
Nitzschia inconspicua Grunow 394, 395
Nitzschia microcephala Grunow 396
Nitzschia sp.1 387, 388
Nitzschia tubicola Grunow 382, 383
Nupela gracillima (Hustedt) Lange-Bertalot 216, 235, 236
Nupela impexiformis (Lange-Bertalot) Lange-Bertalot 218, 237
Nupela silvahercynia (Lange-Bertalot) Lange-Bertalot 217, 238–240
Oxyneis binalis var. elliptica (Flower) Kingston 97
Parlibellus protracta (Grunow) Witkowski et al.
Pinnularia nobilis (Ehrenberg) Ehrenberg 152
Pinnularia subanglica Krammer 166
Pinnularia viridis (Nitzsch) Ehrenberg 153
Pinnularia acrosphaeria W. Smith 163
Pinnularia anglica Krammer 158
Pinnularia beringensis sp. nov. 167–170, 178, 179
Pinnularia burkii Patrick 175
Pinnularia cf. kuetzingii Krammer 177
Pinnularia cf. rhomboelliptica Krammer 157
Pinnularia curticostata Krammer & Lange-Bertalot 176
Pinnularia divergentissima var. subrostrata A. Cleve 140–142
Pinnularia hemiptera var. bielawskii (Heribaud & M. Peragallo) Cleve-
156
Euler
Pinnularia inconstans Mayer 160
Pinnularia nodosa (Ehrenberg) W. Smith 161
Pinnularia obscura Krasske 172
Pinnularia obscuriformis Krammer 143
74 M. Potapova

Taxon Figures
Pinnularia peracuminata Krammer 165
Pinnularia persudetica Krammer 164
Pinnularia pluvianiformis Krammer 144
Pinnularia rhombarea Krammer 159
Pinnularia sp. 1 173
Pinnularia sp. 2 174
Pinnularia subborealis Hustedt sensu Metzeltin et al. 2009 171
Pinnularia subcapitata var. subrostrata Krammer 139
Pinnularia subgibba Krammer 138
Pinnularia transversa (Schmidt) Mayer 154
Pinnularia viridiformis Krammer 162
Placoneis clementis (Grunow) Cox
Placoneis cuneata (M. Moller ex Foged) comb. nov. 331
Planothidium calcar (Cleve) Round et Bukhtiyarova
Planothidium daui (Foged) Lange-Bertalot
Planothidium frequentissimum (Lange-Bertalot) Lange-Bertalot
Planothidium haynaldii (Schaarschmidt) Lange-Bertalot
Planothidium joursacense (Héribaud) Lange-Bertalot 366, 367
Planothidium lanceolatum (Brébisson ex Kützing) Lange-Bertalot
Planothidium oestrupii (Cleve-Euler) Round & Bukhtiyarova
Planothidium peragallii (Brun & Héribaud) Round & Bukhtiyarova
Platessa lutheri (Hustedt) Potapova
Psammothidium acidoclinatum (Lange-Bertalot) Lange-Bertalot 362, 363
Psammothidium altaicum Bukhtiyarova 358, 359
Psammothidium bioretii (Germain) Bukhtiyarova & Round 354, 355
Psammothidium chlidanos (Hohn & Hellerman) Lange-Bertalot 356, 357
Psammothidium daonense (Lange-Bertalot) Lange-Bertalot
Psammothidium helveticum (Hustedt) Bukhtiyarova et Round 338, 339
Psammothidium levanderi (Hustedt) Bukhtiyarova & Round 364, 365
Psammothidium scoticum (Flower & Jones) Bukhtiyarova & Round 360, 361
Psammothidium strelnikovae sp. nov. 370–378
Psammothidium subatomoides (Hustedt) Bukhtiyarova & Round
Pseudofallacia losevae (Lange-Bertalot et al.) Liu et al. 273
Pseudostaurosira brevistriata (Grunow) Williams & Round 72
Pseudostaurosira microstriata var. spinosa Flower 74, 86, 87
Pseudostaurosira parasitica var. subconstricta (Grunow) Morales
Pseudostaurosira pseudoconstruens (Marciniak) Williams & Round 75
Pseudostaurosira robusta (Fusey) Williams & Round 73
Pseudostaurosira subsalina (Hustedt) Morales 88–91, 92–94
Rexlowea navicularis Kociolek & Thomas 284
Rhoicosphenia abbreviata (C. Agardh) Lange-Bertalot
Diatoms of Bering Island, Kamchatka, Russia 75

Taxon Figures
Rhopalodia acuminata Krammer
Rhopalodia gibba (Ehrenberg) Otto Müller
Rossithidium nodosum (Cleve) Aboal 352, 353
Rossithidium pusillum (Grunow) Round & Bukhtiyarova 350, 351
Sellaphora bacillum (Ehrenberg) Mann sensu lato 244
Sellaphora cf. blackfordensis Mann & Droop 245
Sellaphora joubaudii (Germain) Aboal 307
Sellaphora laevissima (Kützing) Mann “very coarse” 247
Sellaphora pseudopupula (Krasske) Lange-Bertalot 246
Sellaphora seminulum (Grunow) Mann
Stauroforma exiguiformis (Lange-Bertalot) Flower et al. 79–81
Stauroneis cf. intricans Van de Vijver & Lange–Bertalot 276
Stauroneis cf. minutula Hustedt 271
Stauroneis gracilis Ehrenberg 279
Stauroneis kriegeri Patrick 275
Stauroneis producta Grunow 274
Stauroneis thermicola (Petersen) Lund 270
Staurophora sp. 1 318
Staurosira construens Ehrenberg
Staurosira construens var. binodis (Ehrenberg) Hamilton 96
Staurosira construens var. exigua (W. Smith) Kobayasi 98
Staurosira sp. 1
Staurosira venter (Ehrenberg) Grunow 70, 95
Staurosirella lapponica (Grunow in Van Heurck) Williams & Round 55, 56
Staurosirella pinnata (Ehrenberg) Williams & Round 67, 68, 71
Stephanodiscus cf. binatus Hakansson & Kling 18–23, 24–28
Stephanodiscus cf. niagarae Ehrenberg 29–31
Surirella amphioxys W. Smith
Surirella brebissonii Krammer & Lange-Bertalot
Surirella cf. linearis W. Smith
Surirella pinnata W. Smith
Synedra mazamaensis Sovereign
Tabellaria fenestrata (Lyngbye) Kützing
Tabellaria flocculosa (Roth) Kützing
Tabularia fasciculata (Agardh) Williams & Round 39, 40
Tryblionella debilis Arnott ex O’Meara
Tryblionella salinarum (Grunow) Pelletan
Ulnaria acus (Kützing) Aboal 38
Ulnaria ulna (Nitzsch) Compère
Ulnaria ulna cf. var. spatulifera (Grunow) Aboal 32, 33, 34–36
Ulnaria ulna var. spatulifera (Grunow) Aboal sensu Patrick & Reimer
37
1966
76 M. Potapova

Class Fragilariophyceae
An unusual species of Ulnaria with swollen valve apices identified here as U. cf. ulna var.
spatulifera was found in two lakes (Figs 32–36). This diatom is remarkably similar to a
specimen illustrated by Metzeltin et al. (2009, l. 17, fig. 11) as (?) Synedra ulna var. spatulifera
Grunow. The type material of S. ulna var. spatulifera has never been investigated, so its identity
is questionable. S. ulna var. spatulifera as illustrated by Patrick & Reimer (1966, pl.7, fig. 8)
has a different shape of valve apices and is relatively common, at least in North America. It
was also found on Bering Island (Fig. 37). The valve outline of U. cf. ulna var. spatulifera is
almost identical to that of Synedra ulna var. claviceps Hustedt described from Sumatra, which
apparently does not have a central area (Hustedt 1937, Simonsen 1987).

Figs 32–40. Ulnaria and Tabularia species. Figs 32, 33 (SEM), 34–36 (LM). Ulnaria cf. ulna var.
spatulifera. Fig. 37 (LM). U. ulna var. spatulifera sensu Patrick & Reimer (1966). Fig. 38 (LM) U. acus.
Figs 39, 40 (LM). Tabularia fasciculata. Scale bars for LM images = 10 μm, SEM images = 2 μm.
Diatoms of Bering Island, Kamchatka, Russia 77

Figs 41–76. Araphid diatoms, LM. Fig. 41. Fragilaria acidoclinata. Fig. 42. Diatoma hyemalis. Fig.
43. D. mesodon. Fig. 44. D. anceps. Fig. 45. D. moniliformis. Figs 46, 47. Fragilaria vaucheriae. Fig.
48. F. bidens. Fig. 49. F. cf. vaucheriae. Figs 50, 51. F. gracilis. Figs 52, 53. Asterionella ralfsii. Fig.
54. Ctenophora pulchella. Figs. 55, 56. Staurosirella lapponica. Fig. 57. Fragilaria sp. 1. Fig. 58.
Fragilariforma bicapitata. Fig. 59. Hannaea arcus var. recta. Fig. 60. H. arcus. Fig. 61. Meridion circulare.
Fig. 62. Fragilariforma lata. Fig. 63. Fragilaria neoproducta. Figs 64–66. Fragilariforma constricta. Figs
67, 68, 71. Staurosirella pinnata sensu lato. Fig. 70. Staurosira venter. Fig. 72. Pseudostaurosira
brevistriata. Fig. 73. P. robusta. Fig. 74. P. microstriata var. spinosa. Fig. 75. P. pseudoconstruens. Fig.
76. Staurosira sp. 1. Scale bar = 10 μm.
78 M. Potapova

Tabularia fasciculata (Figs 39, 40) and Ctenophora pulchella (Fig. 54) were common in
brackish localities. Both diatoms had capitate ends. This shape of the valves was observed
previously in some population of both taxa, but it is not obvious at this time whether it has
taxonomic significance. Two other taxa of araphid diatoms common in brackish waters were
Fragilaria cassubica (Fig 77, 78, 83) and Pseudostaurosira cf. subsalina (Figs 88–94). The
type material of P. subsalina was recently studied by Cejudo-Figueras et al. (2011), who also
described two species similar to P. subsalina. The diatom identified here as P. cf. subsalina
morphologically somewhat differs from the type population. The striae density is 14–16, mostly
around 15 in 10 m, which is on average higher than striae density in the type material, 13–14
in 10 m. The valve width is also only slightly overlaps: it is 3.6–4.1 m in the Bering Island
populations and 4.0–5.3 m in the type population. In the valve shape and the shape of the axial
area, P. cf. subsalina from the Bering Island is more similar to P. americana Morales, than to
the type of P. subsalina, but the valves of P. americana are considerably wider (4.5–5.0 m,
and the striae density is higher (16–18 in 10 m). It is possible that P. cf. subsalina from the
Bering Island is a separate biological species, but a more detailed study of this species comples
is necessary before new species are established based on subtle morphological differences.
Among other araphid diatoms, representatives of the genera Diatoma (Figs 42–45), Hannaea
(Figs 59, 60), and Meridion (Fig. 61) were often found in rivers and streams. In small acidic ponds,
Stauroforma exiguiformis (Figs 79–81) was often a dominant species. Fragilaria acidoclinata
(Fig. 41) and representatives of Fragilariforma (Figs 58, 62, 64–66) were likewise common in
low-pH waterbodies. In a few ponds, Asterionella ralfsii (Figs 52, 53) and Oxyneis binalis var.
elliptica (Fig. 97) were abundant. In larger lakes with higher water pH various cosmopolitan
species of Fragilara, Staurosira, Pseudostaurosira, Staurosirella and Synedra mazamaensis, a
species previously reported only from Western North America, were common.

Class Bacillariophyceae
Subclass Eunotiophycidae
Representatives of the genus Eunotia were especially common in small ponds with low water
pH. The most common species were E. minor (Figs 104–106) and E. rhomboidea (Figs 116,
117). Two new Eunotia species were found.

Eunotia ninae Potapova sp. nov. Figs 99, 100, 109–113, 146–148
DESCRIPTION: Valves straight to weakly concave on the ventral margin, moderately convex on the
dorsal margin, 2.2–3.6 m wide, 14–38 m long. Apices rounded, subrostrate, delimited by a
change in the slope of the dorsal margin. Helictoglossae set in thick internal costae extending to
the dorsal margin. Rimoportula at one end, at the center of the apex, parallel to the apical axis of
the valve. Raphe with distal ends only slightly curved onto the valve surface, without terminal
fissures. Striae 18–23 in 10 μm. Areolae round, 43–46 in 10 μm along the striae.
HOLOTYPE: Circled specimen on slide ANSP GC 36341 (Fig. 112).
ISOTYPES: Circled specimens on slides ANSP GC 36342, CANA 93664.
TYPE LOCALITY: Unnamed tundra pond on Bering Island, Kamchatka, Russia, 55.03106 N,
166.13646 W. Epipelon. Collected by M. Potapova, July 9, 2008.
ETYMOLOGY: This species is dedicated to Prof. Nina Ivanovna Strelnikova.
SIMILAR SPECIES: Eunotia ninae is similar in size and shape to a widely distributed species E.
paludosa, but is easily distinguishable from the latter by the conspicuous costae on the internal
valve surface extending from the helictoglossae to the dorsal margin. Other Eunotia species with
similar costae, such as E. pienitzii and E. lewisii found in North America, have larger cell sizes.
Diatoms of Bering Island, Kamchatka, Russia 79

Figs 77–87. Araphid diatoms. Figs 77, 78 (LM), 83 (SEM). Fragilaria cassubica. Figs 79, 80 (LM),
81 (SEM). Stauroforma exiguiformis. Fig. 82. Fragilaria atomus, SEM. Figs 84, 85. Fragilariforma
bicapitata, SEM. Figs 86, 87. Pseudostaurosira microstriata var. spinosa, SEM. Scale bar for LM
images = 10 μm, SEM images = 2 μm.
80 M. Potapova

Figs 88–98. Araphid diatoms. Figs 88–91 (LM), Figs 92–94 (SEM). Pseudostaurosira subsalina. Fig.
95, SEM. Staurosira venter. Fig. 96, SEM. Staurosira construens var. binodis. Fig. 97, SEM. Oxyneis
binalis var. elliptica. Fig. 98, SEM. Staurosira construens f. exigua.
Diatoms of Bering Island, Kamchatka, Russia 81

Figs 99–128. Eunotia species, LM. Figs 99–100, 109–113. E. ninae, sp. nov., type material. Fig 112.
Holotype specimen. Figs 101, 102. E. fallax. Figs 104–106. E. minor. Fig. 107. E. mucophila. Fig. 108.
E. neocompacta. Fig. 114. E. paludosa. Fig. 115. E. incisa. Figs 116, 117. E. rhomboidea. Fig. 118. E.
ursamajoris. Fig. 119. E. cristagallii. Figs 120, 121. E. paratridentula. Fig. 122. E. meisterioides. Figs
123, 124. E. microcephala. Fig. 125. E. subarcuatoides. Fig. 126. E. iatriaensis. Figs 127, 128. E.
minutula. Scale bar = 10 μm.
82 M. Potapova

Figs 129–145. Eunotia and Pinnularia species, LM. Fig. 129. Eunotia bilunaris. Fig. 130. E. lapponica.
Figs 131–135. E. prilezhaevii, type material. Fig. 134. Holotype specimen. Fig. 136. E. bidens. Fig.
137. E. diadema. Fig. 138. Pinnularia subgibba. Fig. 139. P. subcpitata var. subrostrata. Figs 140–142.
P. divergentissima var. subrostrata. Fig. 143. P. obscuriformis. Fig. 144. P. pluvianiformis. Fig. 145.
Eunotia praerupta. Scale bar = 10 μm.
Diatoms of Bering Island, Kamchatka, Russia 83

Figs 146–151. Eunotia species, SEM. Figs 146–148. E. ninae, sp. nov., type material. Figs 149–151.
E. prilezhaevii, sp. nov., type material. Scale bars = 2 μm.
84 M. Potapova

Eunotia prilezhaevii Potapova sp. nov. Figs 131–135, 149–151

DESCRIPTION: Valves with strongly convex dorsal margin and slightly concave ventral margin, 6.4–
9.6 m wide, 16–40 m long. Apices broad, rounded, slightly to moderately protracted. Terminal
raphe ends noly slightly curved into valve face, terminal fissures very short. Helictoglossae close
to apices. Rimoportulae absent. Striae radiate, 8–12 in 10 m in the middle part of the valve, up
to 16 m at apices. Areolae round, 28–32 in 10 μm along the striae.
HOLOTYPE: Circled specimen on slide ANSP GC 36343 (Fig. 134).
ISOTYPES: Circled specimens on slides ANSP GC 36344, CANA 93665.
TYPE LOCALITY: Unnamed tundra pond on Bering Island, Kamchatka, Russia, 55.027 N, 166.13383
E. Epipelon. Collected by M. Potapova, July 9, 2008.
ETYMOLOGY: This species is dedicated to Ivan Prilezhaev, a colleague who participated in the
2008 expedition to Bering Island and whose help in the field is greatly appreciated.
SIMILAR SPECIES: Eunotia prilezhaevii shares many characters with E. faba Ehrenberg, but differs
from the latter species by protracted valve apices and absence of rimoportula. This species was
also reported as Eunotia spec. cf. faba by Lange-Bertalot et al. (2011, pl. 165, figs 25, 26) from
the material collected in Adak Island.

Subclass Bacillariophycidae
Various, mostly cosmopolitan species of Pinnularia were especially abundant in tundra ponds.
P. curtocostata previously reported from the northern part of Eurasia (REF) was common in
brackish water bodies. A new species of Pinnularia is described here.

Pinnularia beringensis Potapova sp. nov. Figs 167–170, 178, 179

DESCRIPTION: Valves linear-lanceolate to lanceolate with rounded subrostrate apices, 6.1–6.8 m
wide, 25–38 m long. Raphe slightly lateral with slightly deflected proximal ends and question
mark-shaped terminal fissures. Axial area narrow. Central area a broad fascia. Striae slightly
radiate in the middle, becoming strongly convergent at apices, 14–15 in 10 m in the middle, up
to 18 in 10 m at apices. Longitudinal bands absent.
HOLOTYPE: Circled specimen on slide ANSP GC 36345 (Fig. 169).
ISOTYPES: Circled specimens on slides ANSP GC 36346, CANA 63666.
TYPE LOCALITY: Unnamed tundra pond on Bering Island, Kamchatka, Russia, 55.027 N, 166.13383
E. Epipelon. Collected by M. Potapova, July 9, 2008.
ETYMOLOGY: This species is named after its type locality, Bering Island.
SIMILAR SPECIES: Pinnularia beringensis is similar to P. jungii Krammer described from
Spitsbergen (Krammer 2000) and P. certa Krammer & Metzeltin from Guyana (Metzeltin &
Lange-Bertalot 1998). All three species have similar valve shape and very wide central area. P.
beringensis, differs, however, by smaller valve size and higher striae density. The valve width
in P. jungii is 8–20 m, in P. certa it is 8.4–9.4 m, while striae density is 11–13 and 10–
11 in 10 m, respectively. A diatom very similar to P. beringensis, but with slightly narrower
(5.5 m) valve was illustrated by Camburn and Charles (2000, pl. 23, fig. 26) under the name
Pinnularia braunii var. amphicephala f. subconica Venkataraman. The identity of the latter
taxon is not well understood as no photographs of the type material exist. According to the
protologue (Venkataraman 1939), the valves of P. braunii var. amphicephala f. subconica are
slightly narrower (5.5–6 m) and longer (40–43 m) than those of P. beringensis. The original
illustration shows that central area is not as long in the apical direction as in P. beringensis, but
it the accuracy of the drawing is questionable: for instance the proximal raphe ends were shown
as deflected to the same side as the opening of the curvature of the terminal fissures, which is
unlikely in Pinnularia.
Diatoms of Bering Island, Kamchatka, Russia 85

Figs 152–165. Pinnularia species, LM. Fig. 152. P. nobilis. Fig. 153. P. viridis. Figs 154. 155. P.
transversa. Figs 156. P. hemiptera var. bielawskii. 157. P. cf. rhomboelliptica. Fig. 158. P. anglica. Fig.
159. P. rhombarea. Fig. 160. P. inconstans. Fig. 161. P. nodosa. Fig. 162. P. viridiformis. Fig. 163. P.
acrosphaeria. Fig. 164. P. persudetica. Fig. 165. P. peracuminata. Scale bar = 10 μm.
86 M. Potapova

Figs 166–181. Pinnularia and Hydropetra species. Figs 166–177, 180. LM. Figs 178, 179, 181. SEM.
Fig. 166. P. subanglica. Figs 167–170, 178, 179. P. beringensis, sp. nov. Fig. 169. Holotype specimen.
Fig. 171. P. subborealis sensu Metzeltin et al. 2009. Fig. 172. P. obscura. Fig. 173. P. sp. 1. Fig. 174. P.
sp. 2. Fig. 175. P. burkii. Fig. 176. P. curticostata. Fig. 177. P. cf. kuetzingii. Figs 180, 181. Hydropetra
balfouriana.
Diatoms of Bering Island, Kamchatka, Russia 87

Figs 182–209. Chamaepinnularia and Eolimna species. Figs 182–196, 204–208. LM. Figs 197–203,
209. SEM. Figs 182, 183. Chamaepinnularia circumborealis. Figs 184–186, 200, 203. C. krookiformis.
Figs 187–190, 201, 202. C. cf. krookiformis, LM. Figs 191, 192. C. cf. begeri. Figs 193, 197. C.
mediocris. Figs 194, 195, 199. C. soehrensis var. capitata. Fug. 196. C. soehrensis. Figs 198, 204,
205. C. vyvermanii. Fig. 206. C. sp. 1. Figs 207–209. Eolimna minima.
88 M. Potapova

Figs 210–230. Raphid diatoms. Fig. 210. Cavinula jaernefeltii. Fig. 211, 212. C. cocconeiformis. Fig.
213. C. pseudoscutiformis. Fig. 214. C. variostriata. Fig. 215. Adlafia bryophila. Fig. 216. Nupela
gracillima. Fig. 217. N. sylvahercynia. Fig. 218. N. impexiformis. Figs 219–224 (LM), 228–230 (SEM).
Diadesmis mochalovae, sp. nov. Fig. 219. Holotype specimen. Fig. 225. D. perpusilla. Fig. 226. D.
contenta. Fig. 227. D. laevissima. Fig. 231. Adlafia minuscula. Fig. 232. Microcostatus cf. maceria. Fig.
233. Navicula difficillima. Fig. 234. Microcostatus cf. naumannii.
Diatoms of Bering Island, Kamchatka, Russia 89

Small-sized species of Chamaepinnularia, such as C. mediocris (Fig. 193) and C. soehrensis

var. capitata (Figs 194, 195, 199) were often very abundant in tundra ponds. C. krookiformis
(Figs 184–186, 200, 203) was found in larger lakes. A diatom that was smaller, but otherwise
very similar to C. krookiformis (Figs 188–190, 201, 202), was found in a few samples. According
to the original description of C. krookiformis (Krammer 1992), the valve width varies from 5
to 11 m. C. cf. krookiformis from Bering Island, however had valves only 2.5–3.2 m wide.
Such a drastic size difference could probably justify the establishment of a new species, but
considerable variation in valve size and shape has been previously reported for P. krookiformis
(e.g., pl. 45, figs 7–10, pl. 47, fig. 16 in Lange-Bertalot & Genkal 1999).
Among a few species of Diademis encountered on Bering Island, one was identified as new.

Diadesmis mochalovae Potapova sp. nov. Figs 219–224, 228–230

DESCRIPTION: Valves elliptic-lanceolate with wide rounded subrostrate to capitate apices, 3.9–4.8
m wide, 10–18 m long. Striae 16–18 in 10 m, moderately radiate, consist of 2–3 transapically
elongated areolae clearly visible in LM. Axial area lanceolate, central area round or elliptic.
Raphe filiform. Two bean-shaped depressions on external valve surface at proximal raphe ends
are often visible in LM.
HOLOTYPE: Circled specimen on slide ANSP GC 36347 (Fig. 219).
ISOTYPES: Circled specimens on slides ANSP GC 36348, CANA 63667.
TYPE LOCALITY: Unnamed tundra pond on Bering Island, Kamchatka, Russia, 55.03106 N,
166.13646 W. Epipelon. Collected by M. Potapova, July 9, 2008.
ETYMOLOGY: This species is dedicated to Olga Mochalova, a colleague who participated in the
2008 expedition to Bering Island and whose help in the field is greatly appreciated.
Among various naviculoid diatoms, representatives of Frustulia were most common on
Bering Island. In tundra ponds F. crassinervia (Fig. 241), F. saxonica (Fig. 249) were especially
abundant, while in more alkaline lakes and ponds of southern part of the island, F. langebertalotii
was frequently observed.
Two species of Boreozonacola previously reported from Asia and Western North America
were found in Bering Island (Figs 242, 243). Boreozonacola natchikae is considered as here
as conspecific with B. olympica (Sovereign) Lange-Bertalot et al. The latter specific epithet
is commonly used, but the name Navicula pseudosilicula var. olympica was published later
(Sovereign 1963) than N. natchikae (Petersen 1946).
A number of Neidium, Caloneis, and Navicula species are reported here as not identified
to species level. They often did not form large enough populations to assess morphological
variation and to identify them with certainty.
Among gomphocybelloid diatoms, three species of Encyonema, E. hebridicum (Fig. 322),
E. perspusillum (Fig. 323), and E. rostratum (Fig. 324) were abundant in tundra ponds with
relatively low pH. Representatives of the genus Gomphonema were mostly found in larger lakes
with alkaline water. They included several unidentified taxa. Gomphonema sp. 1 (Fig. 334) was
also reported by Foged (1981, pl. 64, fig. 21) from Alaska and by Hein (1990, pl. 23, fig. 7) from
Adak Island as G. brasiliense, but is another, probably not yet established taxon. Gomphonema
cf. olivaceum var. minutissimum (Figs 337, 346) is quite similar to, but has slightly narrower
valves and more parallel striae in the valve center that G. olivaceum var. minutissimum Hustedt.
It is, however, considerably different from G. olivaceoides (Fig. 327), although these two taxa
are often considered synonymous (e.g., Krammer & Lange-Bertalot 1986). A very small diatom
identified here as Gomphosphenia sp. 1 (Fig. 341) is also known from Western North America,
but a detailed comparison of populations from various locales is necessary to accurately delineate
this taxon. The diatom known as Navicula elginensis var. cuneata (M. Moller ex Foged) Lange-
Bertalot in Krammer & Lange-Bertalot and originally described as Navicula dicephala var.
dicephala f. cuneata M. Moller ex Foged has been found in several localities on Bering Island
90 M. Potapova

Figs 235–240. Nupela species. SEM. Figs 235, 236. N. gracillima. Fig. 237. N. impexiformis. Figs
238–240. N. sylvahercynia. Scale bars = 2 μm.
Diatoms of Bering Island, Kamchatka, Russia 91

Figs 241–256. Raphid diatoms, LM. Fig. 241. Frustulia crassinervia. Fig. 242. Boreozonacola
natchikae. Fig. 243. B. hustedtii. Fig. 244. Sellaphora bacillum. Fig. 245. S. cf. blackfordensis. Fig. 246.
S. pseudopupula. Fig. 247. S. laevissima. Fig. 248. Frustulia lange-bertalotii. Fig. 249. F. saxonica. Fig.
250. F. inculta. Fig. 251. Diploneis elliptica. Fig. 252. D. pseudovalis. Figs 253, 254. Mastogloia elliptica.
Fig. 255. Kobayasiella parasubtilissima. Fig. 256. K. subtilissima. Scale bar = 10 μm.
92 M. Potapova

Figs 257–276. Raphid diatoms, LM (Figs 257–161, 263–276), SEM (Fig. 262). Fig. 257. Neidium sp.
1. Fig. 258. N. sacoense. Fig. 259. N. ampliatum. Fig. 260. Caloneis silicula. Fig. 261. C. undulata.
Fig. 262. C. bacillum. Figs 263, 264. Neidium alpinum. Fig. 265. N. bisulcatum. Fig. 266. Caloneis sp.
1. Fig. 267. C. sp. 2. Figs 268, 269. Diatomella balfouriana. Fig. 270. Stauroneis termicola. Fig. 271.
S. cf. minutula. Fig. 272. Neidium cf. alpinum. Fig. 273. Pseudofallacia losevae. Fig. 274. Stauroneis
producta. Fig. 275. S. kriegeri. Fig. 276. S. cf. intricans. Scale bar for LM images = 10 μm, SEM image
= 5 μm.
Diatoms of Bering Island, Kamchatka, Russia 93

Figs 277–285. Raphid diatoms, LM. Fig. 277. Neidium sp. 2. Fig. 278. N. amphigomphus. Fig. 279.
Stauroneis gracilis. Fig. 280. Navicula peregrina. Fig. 281. N. radiosa. Figs 282, 283. N. aurora. Fig.
284. Rexlowea navicularis. Fig. 285. Navicula vulpina. Scale bar = 10 μm.
94 M. Potapova

Figs 286–308. Raphid diatoms, LM. Figs 286–288. Navicula vaneei. Fig. 289. N. salinarum. Fig. 290.
N. leptostriata. Fig. 291. N. gregaria. Fig. 292. N. cf. antonii. Figs 293–295. N. cf. veneta. Fig. 296.
N. cf. hoffmanniae. Fig. 297. N. cryptotenella. Fig. 298. N. streckerae. Fig. 299. N. cf. subalpina. Figs
300, 301. N. cf. arctotenelloides. Fig. 302. N. cf. exilis. Fig. 303. N. cryptotenelloides. Fig. 304. N.
cf. microcari. Fig. 305. N. detenta. Fig. 306. N. digitulus. Fig. 307. Sellaphora joubaudii. Fig. 308.
Hippodonta subcostulata. Scale bar = 10 μm.
Diatoms of Bering Island, Kamchatka, Russia 95

Figs 309–321. Raphid diatoms, LM. Fig. 309. Cymbopleura crassipunctata var. nonornata. Fig. 310.
Cymbella mexicana var. kamtschatica. Fig. 311. C. amplificata. Fig. 312. C. neocistula. Fig. 313. C.
proxima. Fig. 314. Halamphora cf. holsatica. Fig. 315. Encyonema temperei. Fig. 316. Cymbella sp. 1.
Fig. 317. Cymbopleura peranglica. Fig. 318. Staurophora sp. Fig. 319. Gomphonema pala. Fig. 320.
Gomphoneis eriense. Fig. 321. Encyonema kamtschaticum.
96 M. Potapova

Figs 322–347. Raphid diatoms, LM. Fig. 322. Encyonema hebridicum. Fig. 323. E. perpusillum. Fig.
324. E. rostratum. Fig. 325. E. cf. gauemannii. Fig. 326. Encyonopsis sp. 1. Fig. 327. Gomphonema
olivaceoides. Fig. 328. G. acuminatum var. pusillum. Fig. 329. G. khentiiense. Fig. 330. G. gracile.
Fig. 331. Placoneis cuneata comb. nov. Fig. 332. G. olivaceum. Fig. 333. G. micropus. Fig. 334.
Gomphonema sp. 1. Fig. 335. Encyonopsis descripta. Fig. 336. Gomphonema interpositum. Figs
337, 346. G. cf. olivaceum var. minutissimum. Figs 338, 339. Psammothidium helveticum. Fig. 340.
Encyonema reichardtii. Fig. 341. Gomphosphenia sp. 1. Fig. 342. Gomphonema cf. rigidum var.
pumilum. Figs 343, 345. Gomphonema sp. 2. Fig. 344. Gomphosphenia tackei. Fig. 347. Gomphonema
cf. mexicanum. Scale bar = 10 μm.
Diatoms of Bering Island, Kamchatka, Russia 97

Figs 348–378. Monoraphid diatoms, LM. Figs 348–375. LM. Figs 376–378. SEM. Figs 348, 349.
Achnanthidium kranzii. Figs. 350, 351. Rossithidium pusillum. Figs 352, 353. Rossithidium nodosum.
Figs 354, 355. Psammothidium bioretii. Figs 356, 357. P. chlidanos. Figs 358, 359. P. altaicum. Figs
360, 361. P. scoticum. Figs 362, 363. P. acidoclinatum. Figs 364, 365. P. levanderi. Figs 366, 367.
Planothidium joursacence. Figs 368–378. Psammothidium strelnikovae, sp. nov. Figs 372, 373.
Holotype specimen. Fig. 376. External view of raphe valve. Fig. 377. Internal view of raphe valve. Fig.
378. Internal view of rapheless valve. Scale bar for LM images = 10 μm, SEM images = 2 μm.
98 M. Potapova

Figs 379–401. Nitzschia species. Fig. 379. N. angustata. Figs 380, 381. N. cf. sublinearis. Figs 382,
383. N. tubicola. Fig. 384. N. fossilis. Figs 385, 386. N. angustiforaminata. Figs 387, 388. N. sp. 1. Figs
389–392. N. acidoclinata. Fig. 393. N. cf. homburgiensis. Figs 394, 395. N. inconspicua. Fig. 396. N.
microcephala. Fig. 397. N. cf. paleacea. Fig. 398. N. cf. solita. Fig. 399. N. dissipata var. media. Figs
400, 401. N. bryophila. Scale bar for LM images = 10 μm, SEM image (Fig. 389) = 2 μm.
Diatoms of Bering Island, Kamchatka, Russia 99

(Fig. 331). This species is transferred here to the genus Placoneis and elevated to the species
rank.

Placoneis cuneata (M. Moller ex Foged) Potapova, comb. et stat. nov.

BASIONYM: Navicula dicephala f. cuneata M. Moller ex Foged 1977, Bibliotheca Phycologica
34, p. 78; pl. 29, fig. 6.
REMARKS: Navicula dicephala f. cuneata was originally described from Ireland by Foged (1977)
and later transferred to N. elginensis var. cuneata by Kobayasi (Kobayasi & Ando 1975).
This species (Fig. 331) is indeed similar to Placoenis elginensis (Gregory) Cox, but is easily
distinguished from the latter by the lanceolate valve with cuneate apices.
Monoraphid diatoms were most abundant in flowing waters. Common species Planothidium
lanceolatum and P. frequentissimum and cold-water P. haynaldii were extremely abundant in
rivers with alkaline water. Achnanthidium kranzii (Figs. 348, 349), Rossithidium nodosum (Figs
352, 353), R. pusillum (Figs 350, 351) and various species of Psammothidium were frequently
observed in both lentic and lotic habitats. A new species of Psammothidium was found in several
creeks.

Psammothidium strelnikovae Potapova sp. nov. Figs 368–378

DESCRIPTION: Valves elliptic with cuneate apices, 4.6–5.6 m wide, 8–14 m long. Raphe valve
with narrow linear axial area and large transversely rectangular central area almost reaching
valve margins. Raphe filiform, straight with straight external distal and proximal ends. Terminal
raphe fissures absent. External proximal raphe ends in a groove. Internal proximal raphe ends
turned to opposite sides, internal distal endings with small helictoglossae. Rapheless valve with
rhomboid axial area of variable size, from small to wide often with irregular edge. Central area
is not differentiated from axial area. Striae in both valves uniseriate, slightly radiate, 32–36 in 10
m on raphe valve, around 30 in 10 m on rapheless valve. Areolae on raphe valve isodiametric,
indistinguishable in LM, approximately 40 in 10 m, 6–8 areolae in longest striae. Areolae on
rapheless valve transapically elongated, visible in LM, approximately 30–32 in 10 m, 4–5
areolae in longest striae. One row of mantle areolae on both valves.
HOLOTYPE: Circled specimen on slide ANSP GC 36349 (Figs 372, 373).
ISOTYPES: Circled specimens on slides ANSP GC 36350, CANA 63668.
TYPE LOCALITY: Unnamed creek on Bering Island, Kamchatka, Russia, 55.03019 N, 166.13867
W. Epipelon. Collected by M. Potapova, July 9, 2008.
ETYMOLOGY: This species is dedicated to Prof. Nina Ivanovna Strelnikova.
SIMILAR SPECIES: Psammothidium strelnikovae is most similar morphologically to Achnanthes
subsalsa Petersen, which has, however wider (5–10 m) valves and much lower density of striae
(20–28 in 10 m on raphe valve, 18–26 in 10 m on rapheless valve).
Canal-raphe diatoms rarely formed large populations in Bering Island. In tundra ponds
Nitzschia acidoclinata (Figs 389–391) and N. bryophila (Figs 400, 401) were frequently
observed. N. tubicola (Figs 382, 383), N. cf. sublinearis (Figs 380, 381) and Surirella brebissonii
were occasionally abundant in lakes and rivers of the southern part of the Island.

Discussion
This study is the first survey of Bering Island diatoms comprising all types of aquatic habitats
found on the island. From 313 taxa reported here, 48 are listed under provisional names. Most
of these diatoms did not fit entirely descriptions of published species and did not form large
enough populations so that their identity could not be studied in detail. It is important, however,
100 M. Potapova

that images of these diatoms are now available for comparison and as an information source for
future biogeographical studies.
Although most diatoms identified to species level are either cosmopolitan or known to have
a wide distribution range in Northern Hemisphere, many are especially characteristic for arctic
and alpine regions. These include, for example, Aulacoseira nivalis, Diatoma mesodon and D.
hiemalis, Hannaea arcus, Staurosirella lapponica, all three encountered species of Nupela:
N. impexiformis, N. gracillima, and N. silvahercynia, Chamaepinnularia circumborealis and
C. krookiformis, Frustulia lange-bertalotii, Boreozonacola hustedtii, Achnanthidium kranzii,
Rossithidium pusillum and R. nodosum.
A number of species from Bering Island are known to have limited geographic distributions.
So far reported only from North America and Eastern Asia were Cymbella amplificata, C.
mexicana var. kamtschatica (Krammer 2002), Gomphoneis eriense (Kociolek & Stoermer
1988, Skvortzow & Meyer 1928), and Gomphosphenia sp.1. Neidium sacoense and Frustulia
inculta were previously found only in Eastern North America (Siver et al. 2005, 2009). Navicula
aurora was only observed in Western North America (Lange-Bertalot 2001). Several species
found on Bering Island have been known so far only from Asia. These were: Gomphonema
khentiinense reported from Mongolia (Kulikovskiy et al. 2010, Metzeltin et al. 2009), Encyonema
kamtschaticum known from Kamchatka (Krammer 1997), and Cymbopleura crassipunctata var.
nonornata listed by Krammer (2003) and reported by Hein (1990) as Cymbella acutisucula
Cleve).
A comparison with diatom flora of Adak Island (Hein 1990) is possible because all diatoms
encountered in the latter study were illustrated by LM images. From 321 taxa found in Adak
Island, 127 (40 %) were also found in Bering Island, although different names were often attached
to these diatoms. The most abundant species on Adak Island were Achnanthidium minutissimum,
Rossithidium pusillum, Brachysira brebissonii (reprted as Anomoneis brachysira) and Diatoma
moniliformis (reported as D. tenue). On Bering Island A. mintissimum and D. moniliformis
were rather uncommon, while R. pusillum was far from being an abundant species. There are,
therefore, considerable differences in diatom communities between these islands, although at
this point it is impossible to determine what proportion of this variation is due to environmental
differences or historical processes.

Acknowledgements
This study was supported by a grant from the Trust for Mutual Understanding. The use of the Centralized
Research Facilities, Drexel University, for the SEM is gratefully acknowledged. I thank Olga Mochalova,
Eugene Potapov, Larisa Zelenskaya, Ivan Prilezhaev, and Mikhail Romanov for the logistic support and
invaluable help in the field.

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