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CZECH POLAR REPORTS 5 (2): 112-133, 2015
———
Received November 4, 2015, accepted December 21, 2015.
*Corresponding author: Petra Vinšová <vinsova@gmail.com>
Acknowledgements: This research was made during the Polar Ecology Course (2014) supported by
project nr. CZ.1.07/2/2.2.00/28.0190, and funded by Czech Ministry of Education (MSMT) via
grant LM2010009. PV was also funded by Fond Mobility UK (2014). Part of this research was
financially supported by the Fund for Scientific Research—Flanders (FWO-Flanders, Belgium,
funding of EP as a PhD student). This study was also supported as a long-term research
development project RVO no. 67985939.
112
Diatoms in cryoconite holes and adjacent proglacial freshwater
sediments, Nordenskiöld glacier (Spitsbergen, High Arctic)
Petra Vinšová1*, Eveline Pinseel2,3,4, Tyler J. Kohler1, Bart Van de
Vijver3,4, Jakub D. Žárský 1, Jan Kavan5, Kateřina Kopalová1,5,6
1Charles University in Prague, Faculty of Science, Department of Ecology, Viničná 7,
CZ-128 44 Prague 2, Czech Republic
2Ghent University, Faculty of Science, Department of Biology, Protistology & Aquatic
Ecology (PAE), Krijgslaan 281-S8, BE-9000 Ghent, Belgium
3Botanic Garden Meise, Department Bryophyta & Thallophyta, Nieuwelaan 38, BE-
1860 Meise, Belgium
4University of Antwerp, Faculty of Science, Department of Biology, Ecosystem
Management Research Group (ECOBE), Universiteitsplein 1, BE-2610 Wilrijk, Belgium
5University of South Bohemia, Faculty of Science, Centre for Polar Ecology (CPE),
Branišovská 31, CZ-370 05 České Budějovice, Czech Republic
6Academy of Science of the Czech Republic, Institute of Botany, Section of Plant
Ecology, Dukelská 135, 379 82 Třeboň, Czech Republic
Abstract
Cryoconite holes are small, extreme habitats, widespread in the ablation zones of gla-
ciers worldwide. They can provide a suitable environment for microorganisms including
bacteria, cyanobacteria, algae, fungi, and invertebrates. Diatoms have been previously
recovered from cryoconite holes of Greenland and of Svalbard, and recent findings from
Antarctica suggest that cryoconite holes may harbor a unique diatom flora distinct from
other aquatic habitats nearby. In the present study, we characterize the diatom communi-
ties of Nordenskiöld glacier cryoconite holes in Billefjorden (Svalbard, Spitsbergen), and
multivariate approaches were used to compare them with three freshwater localities in
the immediate vicinity to investigate possible sources of the species pool. We found
cryoconite holes to have similar or greater average genus-richness than adjacent lake/
ponds habitats, even though lower numbers of valves were recovered. Overall, cryo-
conite hole diatom communities differed significantly from those observed in lakes,
suggesting that other sources actively contribute to these communities than nearby lakes
alone. This further suggests that (i) diatoms present in cryoconite might not exclusively
originate from aquatic habitats, but also from (semi-) terrestrial ones; and (ii) that a
much wider area than the immediate surroundings should be considered as a possible
source for cryoconite diatom flora.
DOI: 10.5817/CPR2015-2-11
P. VINŠOVÁ et al.
113
Key words: ecology, limnology, lakes, cryosphere, Svalbard, polar region
Introduction
Glaciers and ice sheets cover almost
10% of the Earth’s landmass (Clark 2009),
and as a result, their suitability for life is of
great importance. Cryoconite holes repre-
sent microhabitats formed by dust trans-
ported into the ablation zones of the gla-
cial surface, which leaves behind small
water-filled depressions during thawing
periods. These supraglacial habitats are
found worldwide and can persist for days
to decades (Hodson et al. 2008). Since the
1930’s, cryoconite holes have been recog-
nized as a micro-environmnent harboring
life (Steinbock 1936), and modern investi-
gations have shown that they support a
range of micro-organisms including ar-
chaea, heterotrophic bacteria, fungi, micro-
algae, filamentous cyanobacteria, nema-
todes, rotifers and tardigrades (e.g. Wharton
et al. 1981 and 1985, Mueller et al. 2001,
Edwards et al. 2013a). In contrast to the
surrounding glacier surface, cryoconite
holes supply liquid water to organisms,
can shelter microorganisms from UV light,
and are presumably also less susceptible to
temperature extremes. Biovectors and wind
are thought to serve as dispersal agents for
microbial propagules to the glacier sur-
face. Both local and long range aeolian
input (Šabacká et al. 2012, Budgeon et al.
2012), as well as debris from avalanches
and erosion, can supply the surface with
sediment (Hodson et al. 2008) and organic
carbon (Stibal et al. 2008).
Antarctic cryoconite holes may be
isolated from the atmosphere for multiple
melt seasons by a thick ice lid (Fountain et
al. 2004), allowing the development of
unique habitats that harbor diverse com-
munities (Mueller et al. 2001). In contrast,
only a thin ice lid may develop during
summer months in the Arctic, resulting in
hydrologically connected environments that
are frequently flushed with meltwater, pro-
moting a more homogenous resident com-
munity (Edwards et al. 2011). Never-
theless, biological activity of inhabitants
(e.g. granule formation, darkening proc-
esses and photosynthetic activity) remains
surprisingly high for such an extreme
habitat (Vonnahme 2014), and as known
from the Antarctic, cryoconites can serve
as refuges for aquatic and terrestrial micro-
organisms (Foreman et al. 2007, Stanish et
al. 2013). Cryoconite communities may
also (re-)seed downstream microbial com-
munities residing in proglacial lakes and
streams with cells and propagules (Yallop
et Anesio 2010, Stanish et al. 2013) that
can stand at the very onset of microbial
colonization (Stibal et al. 2006).
Most studies of glacial microbial com-
munities have focused on bacteria or cyano-
bacteria (Mueller et al. 2001, Christner et
al. 2003, Cameron et al. 2012, Edwards et
al. 2013b), which are the most abundant
primary producers in these habitats (Muel-
ler et al. 2001, Porazinska et al. 2004, Stibal
et al. 2006, Stibal et Tranter 2007). Our
recent knowledge of other cryoconite hole
phototrophs such as diatoms (Bacillario-
phyceae) is, however, rather poor. It is in
spite of the fact that diatoms are one of the
most successful groups of unicellular algae
worldwide, inhabiting a wide range of
aquatic and terrestrial environments in-
cluding polar regions (Jones 1996, Van de
Vijver et Beyens 1999, Sabbe et al. 2003,
Van de Vijver et al. 2005, Antoniades et al.
2008, 2009). Furthermore, the species-
specific characteristics of their outer silica
cell-wall, as well as individual responses
to the physico-chemical environment make
diatoms excellent bio-indicators in applied
sciences such as paleo-ecology and bio-
geography (Spaulding et al. 2010).
While diatoms have been previously
recorded from supraglacial habitats (Muel-
DIATOMS FROM CRYOCONITE HOLES IN SPITSBERGEN
114
ler et al. 2001, Van de Vijver et al. 2010b,
Cameron et al. 2012), speculation persists
as to how these communities are assem-
bled. It is possible that diatoms, together
with other micro-organisms present in
microbial mats, can be delivered to the
supraglacial habitats from nearby lakes
and streams by winds (Nkem et al. 2006)
and solely reflect those in transport. On the
other hand, they may constitute inde-
pendently functioning communities with a
composition similar, yet distinct from their
surroundings. To our best knowledge, only
two diatom-focused studies of cryoconite
holes exist at this time. One relates to the
Arctic (Yallop et Anesio 2010), the other
one to the Antarctic (Stanish et al. 2013).
The former one reports cultured diatoms
from cryoconite holes and compares these
with database samples. The latter one com-
pares cryoconite hole communities with
microbial mats from adjacent streams.
Therefore, much work remains for the
diatom-related scientific community in or-
der to gain a better understanding of the
structure and function of these extreme
habitats.
To address such gap in our knowledge,
we characterize diatom communities from
cryoconite hole sediments of the Norden-
skiöld glacier (Adolfbukta, Billefjorden,
Svalbard) and compare these results with
adjacent aquatic habitats that could po-
tentially serve as a source. We hypothesize
that if cryoconite holes are seeded by these
adjacent habitats, supraglacial diatom com-
munities should reflect the diatom commu-
nities in surrounding lakes and ponds. An
alternative to this hypothesis is the poten-
tial importance of other nearby habitats
such as terrestrial areas and streams, and/
or more long range transport from other
sources. To answer this question, a simple
genus-based approach was applied to re-
duce the effects of taxonomic uncertainty,
and the results were analyzed using multi-
variate approaches to determine the simi-
larity of communities between habitats.
Material and Methods
Sample collection
Since 2007, summer research cam-
paigns organized by the Centre for Polar
Ecology (University of South Bohemia in
České Budějovice, Czech Republic) have
annually taken place in Petuniabukta (see
reports of the research activities at web-
site CPE: http://polar.prf.jcu.cz/docs.htm or
http://polar.prf.jcu.cz/index.htm) located in
the central part of Spitsbergen (Svalbard
Archipelago) (see Fig. 1). The fjord is sur-
rounded by a lowland area of marine ter-
races, and steep slopes up to 937 m. Its
climate is characterized by low precipi-
tation rates (about 200 mm y-1), relatively
warm winters (-6.7 to -17°C), and wind
speed ranging from 2.8 to 23.6 m s-1 (at
78° 42’ N and 16° 27’ E) (Rachlewicz et
al. 2007, Láska et al. 2012).
During the 2014 boreal summer season,
12 samples from cryoconite holes were
collected (Fig. 2) along two sampling lines
on the Nordenskiöld tidewater glacier (gla-
cier description in Rachlewicz et al. 2007).
The first line was situated on an altitudinal
gradient from the glacier front upwards,
and the second one from the side marginal
moraine to the glacier center (Fig. 1B and
Fig. 2). These sites were chosen to study
possible patterns in aeolian transport of
diatoms from the surrounding environ-
ments, which should occur from the sea
and/or from the marginal zones towards
the glacier surface. Sediment samples from
small cryoconite holes (3 – 5 cm in diame-
ter, 10 – 15 cm in depth) were collected
with pipettes with enlarged openings, stored
P. VINŠOVÁ et al.
115
Fig. 1. Map of the study sites. Sampling points are indicated by crosses. On the last image,
C refers to cryoconite holes, H to Hørbye lakes, N to Nordenskiöld lakes, and O to Retrettøya
ponds site.
DIATOMS FROM CRYOCONITE HOLES IN SPITSBERGEN
116
in 25 mL tubes, and preserved with 96%
ethanol. A hand-held GPS was used to
determine altitude and location, and pH,
temperature and conductivity were meas-
ured in the field using a HANNA Instru-
ment HI 98129 Waterproof pH/Conduc-
tivity/TDS Tester (Hanna Instruments
Czech s.r.o., Czech Republic). Additional-
ly, the water temperature of one stable
cryoconite hole on Nordenskiöld glacier
surface was measured every hour for 28
days between the 22nd of July and 25th of
August using a temperature datalogger
‘minikin T’ (EMS Brno, Czech Republic),
revealing stable values between – 1°C and
1°C.
During the 2011 and 2013 summer
campaigns, benthic epilithon/epipelon sam-
ples from the littoral zones of freshwater
lakes and ponds were collected for diatom
analysis as described above for cryoconite
holes. For a full species report of fresh-
water diatoms from Petuniabukta, see
Pinseel (2014). Three of these localities
from Pinseel (2014) (Fig. 1C) were se-
lected for comparison with the cryoconite
samples. The first locality, Retrettøya (O)
(known also as ‘roche moutonnée’ or
‘Oblik’), is a peninsula located in front of
the tide-water glacier Nordenskiöld. The
area is rather freshly deglaciated, being
covered by the Nordenskiöld glacier until
about 30 years ago, and therefore open to
organismal colonization. Several ponds on
this peninsula, situated in eroded tectonic
faults later remodeled by glacial erosion
(Pinseel 2014), are located in close vi-
cinity of the sea, surrounded by a large
colony of Arctic terns, Sterna paradisaea
Pontoppidan. Altogether, glacial influence,
the presence of the Arctic terns colony,
and possible sea spray influence make this
peninsula a unique locality in terms of
nutrient sources. The other two localities
are comprised of kettle lakes located in the
frontal moraines of the Nordenskiöld (N)
and Hørbye (H) glaciers, the latter being
located in the northern part of Petunia-
bukta. All samples used in this study (both
lakes and cryoconite holes), together with
their parameters, are listed in Table 1.
Slide preparation and enumeration
For light microscopy analysis, sub-
samples were cleaned by a modified meth-
od described in Van der Werff (1955). The
sub-samples were added 37% H2O2 and
then heated to 80°C for about 1h. The
reaction was completed by addition of
saturated KMnO4. Following digestion and
oxidation, samples were rinsed three times
with distilled water alternated with centri-
fugation (10 minutes at 3500 x g). The
cleaned diatom material was diluted with
distilled water on microscope cover slides,
dried, and mounted in Naphrax®.
In each sub-sample, diatom valves were
identified to the lowest taxonomic level
possible (taxa with uncertain taxonomic
status were indicated with ‘cf.’ (confer:
species identification is uncertain) or ‘sp.’,
and sometimes only genus level was
possible) and enumerated at 1500 x magni-
fication under immersion oil using an
Olympus® BX51 microscope equipped
with Differential Interference Contrast
(Nomarski) optics. Diatoms were very rare
in the cryoconite hole samples, and there-
fore entire microscope slides were counted
for diatom valves. In total, 9 samples were
used for the community analysis, with
2 additional samples (C10 and C11, with
only a few recovered valves) used in the
“inkspot” diagram (for explanation, see
below). Of these, total counts ranged be-
tween 40 and 216 valves per sample. One
sample (C12) did not contain any frustules
and was therefore removed from further
analysis.
P. VINŠOVÁ et al.
117
Study code
(sample code)
Location/
Lake type
Temper-
ature
(°C)
Conduc-
tivity
(µS/cm)
pH
Altitude
a.s.l.(m) GPS N GPS E
Cryoconite holes
C1 (N002)
Glacier
surface 0.5 1 8.30
130 78° 38' 24.7"
16° 58' 37.1"
C2 (N003)
Glacier
surface 0.7 1 8.60
201 78° 38' 24.6"
17° 00' 05.1"
C3 (N004)
Glacier
surface 0.6 1 8.30
244 78° 38' 22.2"
17° 01' 44.6"
C4 (N006)
Glacier
surface 0.5 1 9.10
278 78° 38' 28.3"
17° 05' 11.0"
C5 (N007)
Glacier
surface 0.4 7 9.30
361 78° 38' 29.2"
17° 06' 37.6"
C6 (N008)
Glacier
surface 0.4 4 9.80
393 78° 38' 30.5"
17° 07' 22.7"
C7 (N009)
Glacier
surface 0.3 1 8.50
262 78° 39' 21.4"
17° 03' 40.9"
C8 (N010)
Glacier
surface 0.4 2 8.40
263 78° 39' 01.0"
17° 03' 31.9"
C9 (N011)
Glacier
surface 0.5 12 10.10
267 78° 38' 41.5"
17° 03' 16.2"
C10 (N012)
Glacier
surface 0.5 2 9.20
237 78° 38' 08.6"
17° 03' 01.0"
C11 (N013)
Glacier
surface 0.4 11 8.80
233 78° 37' 51.9"
17° 02' 55.3"
C12 (N005)
Glacier
surface 0.5 2 9.00
271 78° 38' 25.6"
17° 03' 46.9"
Nordenskiöld lakes (Pinseel 2014)
N1 (SP20)
Kettle
lakes 8.3 443 8.52
28 78° 38' 19.2''
16° 49' 35.8''
N2 (SP21)
Kettle
lakes 8.6 329 8.55
24 78° 38' 19.4''
16° 49' 41.1''
N3 (SP22)
Kettle
lakes 8.4 658 8.33
29 78° 38' 17.6''
16° 50' 1.5''
N4 (SP23)
Kettle
lakes 8.7 566 8.60
26 78° 38' 17.6''
16° 50' 5.1''
Hørbye lakes (Pinseel 2014)
H1 (SPH1B)
Kettle
lakes ND ND ND 37 78° 44' 05.7"
16° 26' 52.5"
H2 (SPH1C)
Kettle
lakes 10.9 861 8.18
23 78° 44' 08.1"
16° 26' 51.9"
H3 (SPH2A)
Kettle
lakes 11.7 461 8.39
41 78° 44' 13.4"
16° 26' 27.7"
H4 (SPH2B)
Kettle
lakes 12.0 760 8.13
43 78° 44' 11.2"
16° 26' 15.2"
H5 (SPH2C)
Kettle
lakes 11.6 756 8.20
44 78° 44' 11.8"
16° 26' 10.1"
H6 (SPH3A)
Kettle
lakes 13.3 338 8.41
49 78° 44' 22.0"
16° 25' 04.8"
DIATOMS FROM CRYOCONITE HOLES IN SPITSBERGEN
118
H7 (SPH3B)
Kettle
lakes 12.7 306 8.42
50 78° 44' 23.9"
16° 24' 56.3"
H8 (SPH3C)
Kettle
lakes 14.2 339 8.54
46 78° 44' 21.4"
16° 25' 08.2"
H9 (SPH4B)
Kettle
lakes 11.3 423 8.28
52 78° 44' 34.2"
16° 24' 42.4"
H10 (SPH4C)
Kettle
lakes 14.5 655 8.32
50 78° 44' 34.2"
16° 24' 48.7"
H11 (SPH5A)
Kettle
lakes 13.9 1428 8.12
54 78° 44' 43.9"
16° 24' 27.4"
H12 (SPH5B)
Kettle
lakes 12.5 1805 8.18
57 78° 44' 45.4"
16° 24' 37.0"
H13 (SPH5C)
Kettle
lakes 13.3 1208 8.32
57 78° 44' 47.2"
16° 24' 39.5"
H14-15
(SP41-42)
Kettle
lakes 11.6 1325 8.18
35 78° 44' 05.0"
16° 26' 34.1"
H16 (SP44)
Kettle
lakes 12.7 180 8.45
40 78° 44' 13.0"
16° 26' 14.1"
H17 (SP45)
Kettle
lakes 12.2 218 8.54
50 78° 44' 35.4"
16° 24' 44.9"
H18 (SP46)
Kettle
lakes 9.2 198 9.20
106 78° 45' 14.0"
16° 21' 46.2"
H19-20
(SP47-48)
Kettle
lakes 8.7 182 8.39
107 78° 45' 24.5"
16° 22' 00.8"
Retrettøya ponds (as ‘roche moutonnée/Oblík’ in Pinseel 2014)
O1 (SP62)
Tectonic
related 8.2 592 8.80
20 78° 39' 24.9''
16° 54' 46.5''
O2 (SP63)
Tectonic
related 8.1 256 9.60
7 78° 39' 27.8''
16° 54' 34.3''
O3 (SP64)
Tectonic
related 8.5 377 8.90
24 78° 39' 23.0''
16° 54' 44.4''
O4 (SP65)
Tectonic
related 8.1 281 9.30
18 78° 39' 19.3''
16° 54' 37.8''
O5 (SP66)
Tectonic
related 7.4 510 8.70
20 78° 39' 21.4''
16° 54' 39.9''
O6 (SP67)
Tectonic
related 8.0 319 8.70
28 78° 39' 18.6''
16° 54' 38.5''
O7 (SP68)
Tectonic
related 8.4 174 9.80
24 78° 39' 16.3''
16° 54' 21.9''
O8 (SP69)
Tectonic
related 9.0 347 9.20
7 78° 39' 8.5'' 16° 54' 51.4''
O9 (SP70)
Tectonic
related 9.0 450 9.40
11 78° 39' 14.6''
16° 55' 28.5''
O10
(SP71-72)
Tectonic
related 9.1 147 9.60
26 78° 39' 14.1''
16° 55' 16.4''
O11 (SP73)
Tectonic
related 9.7 133 10.90
17 78° 39' 9.0'' 16° 55' 8.7''
Table 1. List of all samples used in this study and physico-chemical parameters of the study sites.
P. VINŠOVÁ et al.
119
For the identification of diatom species
from cryoconite hole material, we primari-
ly consulted Pinseel (2014). Along with dia-
toms, the presence of Chrysophyte stomato-
cysts (golden brown algae) was also noted,
although abundances were generally very
low. Following Yallop et Anesio (2010),
count data were combined at the genus
level. Due to the widespread practice of
‘force-fitting’ Arctic taxa into their Euro-
pean and North-American relatives (Tyler
1996), or the use of a too broad morpho-
species concept (Mann 1999), a substantial
number of taxa has been incorrectly identi-
fied in the past. As a consequence, many
studies which identified Arctic diatoms to
the species level might not be reliable
(Pinseel 2014). On the contrary, the genus
level is taxonomically more robust and en-
sures consistency between datasets. More-
over, the large amount of debris, together
with an association of cells with mineral
particles, makes the observation of single
small valves rather difficult (Stibal et al.
2015), and sometimes even unfeasible
when looking for diatoms in living sam-
ples (Vonnahme 2014), impeding identi-
fication of diatom valves up to the species
level.
Statistical analyses
To investigate the similarity of the cryo-
conite diatom flora with those of nearby
freshwater habitats, we compared our cryo-
conite hole dataset with diatom counts
from Pinseel (2014), who counted and
identified 400 diatom valves in littoral sam-
ples from freshwater ponds and lakes in
Petuniabukta, using the methods described
above. Relative abundances were first
calculated from diatom counts from both
datasets, and an “inkspot” plot was created
using the rioja R package (Juggins 2012)
to manageably view diatom community
structure among samples.
Diatom communities were statistically
analyzed using the approaches applied by
Stanish et al. (2012, 2013). Briefly, non-
metric multidimensional scaling (NMDS)
analyses were performed to visualize re-
lationships between communities from
different samples, sites, and habitats. Rare
species (< 1.0%) were removed, all data
square-root transformed, and a distance
matrix was calculated based on Bray–
Curtis dissimilarity using the vegan R
package (Oksanen et al. 2011). From this,
a three dimensional model was created,
which produced a Kruskal’s “stress” value
of 9.88%, and strong nonmetric (R2 =
0.99) and linear fits (R2 = 0.94). To dis-
cern which genera drive patterns between
samples, a corresponding NMDS figure
was produced with genera > 5.0% in
relative abundance superimposed.
Lastly, a hierarchical dendrogram was
produced by creating a distance matrix as
described above, utilizing the “average”
clustering method (coeff = 0.81). To test
for significant differences between the
lake and cryoconite communities in gener-
al, the cryoconite hole diatoms communi-
ties were tested against the pooled lake
diatom community data using permuta-
tional multivariate analysis of variance
(PERMANOVA), with α = 0.05. All analy-
ses were performed using the R statistical
environment (R Core Team 2014, see
References – Other sources).
DIATOMS FROM CRYOCONITE HOLES IN SPITSBERGEN
120
Fig. 2. Examples of cryoconite holes (A–B), and an overview of the Nordenskiöld tide-water
glacier (C).
Results
Species composition of the cryoconite holes
A total of 58 diatom taxa (including
species, subspecies, varieties and formas)
belonging to 46 genera were identified in
the cryoconite material (Fig. 3). An ad-
ditional 26 diatom frustules could not be
identified below genus level. Genus rich-
ness of the cryoconite samples ranged
from 9 to 24 with a median of 18. Some
genera were common throughout the sam-
pling sites, such as Pinnularia Ehrenberg
(in all samples), Nitzschia Hassall (in 8 out
of 9 samples), Staurosirella D.M.Williams
(8/9), Gomphonema Ehrenberg (8/9) and
Luticola D.G.Mann (8/9). The dominance
of these genera are also reflected in the
number of counted valves: Nitzschia (17%
of all counted valves), Psammothidium L.
Bukhtiyarova (14%), Pinnularia (12%),
Staurosirella (6%), Gomphonema (5%)
and Luticola (5%).
The most species-rich genera were
Pinnularia, Nitzschia and Eunotia
Ehrenberg. Six Pinnularia taxa could be
identified – P. cf. brebisonii (Kützing)
Rabenhorst, P. obscura Krasske, P. inter-
media (Lagestedt) Cleve, P. schimanskii
Krammer, P. rabenhorstii (Grunow) Kram-
mer, and P. borealis Ehrenberg (Fig. 3).
The lattermost actually represents one of
the most common species complexes (in 8
out of 9 samples) on the site, together with
P. intermedia (7/9), Nitzschia perminuta
complex form 2 (Pinseel 2014; 7/9),
Gomphonema aff. nathorstii (Pinseel
2014; 8/9), and Psammothidium sp1
(Pinseel 2014; 7/9) - see Fig. 3.
P. VINŠOVÁ et al.
121
Several other Nitzschia taxa, such as
N. amphibia Grunow, N. communata
Grunow, N. flexoides Geitler, N. palea
(Kützing) W.Smith and N. sp8 (Pinseel
2014) have been observed in low numbers
(containing less than 2.5 % out of the total
species counts). Four Eunotia taxa have
been identified: E. cf. ambivalens Lange-
Bertalot & Tagliaventi, E. curtagrunowii
Nörpel-Schempp & Lange-Bertalot (Fig.
3), E. islandica Østrup and E. nymanniana
Grunow. Three species out of the most
common taxa also dominated the cryo-
conite sites in terms of relative abun-
dances: Nitzschia perminuta form 2,
Psammothidium sp1 and Pinnularia
borealis complex, represented 12%, 10%
and 5% respectively.
Fig. 3. Selected diatom species observed in cryoconite holes of Nordenskiöld glacier. Scale bar =
10 µm. 1. Humidophila cf. contenta, 2. Chamaepinnularia sp., 3. Rossithidium petersenii,
4. Eunotia curtagrunowii, 5. Achnanthidium minutissimum complex. 6, Staurosirella sp.,
7. Psammothidium sp1., 8. Stephanodiscus cf. minutulus, 9. Gomphonema aff. nathorstii,
10. Nitzschia perminuta forma 1., 11. Pinnularia intermedia, 12. Luticola nivalis, 13. Pinnularia
borealis complex, 14. Hantzschia amphioxys complex.
Local spatial comparison
To examine diatom community simi-
larity between localities, the genus rich-
ness of the cryoconite holes and nearby
lake habitats was compared. All observed
genera, their abbreviations used for analy-
ses and their distribution in between com-
pared localities, can be found in Table 2.
Even though only a low number of diatom
valves could be enumerated from the cryo-
conite samples, overall genus richness was
comparable, and sometimes even greater,
than some of the lake sites (Fig. 4).
DIATOMS FROM CRYOCONITE HOLES IN SPITSBERGEN
122
Fig. 4. Genus richness of cryoconite holes (C), Hørbye lakes (H), Nordenskiöld lakes (N), and
Retrettøya (O) sites.
Fig. 5. Non-metric multidimensional scaling (NMDS) of diatom communities from cryoconite
holes and nearby lakes, indicating separation of the two habitats. C1–C9 for cryoconite holes,
H1–H20 for Hørbye lakes, N1 – N4 for Nordenskiöld lakes, and O1–O11 for Retrettøya. Diatom
genus abbreviations are given in Table 2.
P. VINŠOVÁ et al.
123
Genus C
H
N
O
Achnanthidium (Ach) x x x x
Adlafia (Adl) x x x
Alveovallum (Alv) x
Amphora (Amp) x x
Aulacoseira (Aul) x
Brachysira x
Caloneis (Cal) x x x x
Cavinula x
Chamaepinnularia (Cha) x x x
Cocconeis (Coc) x
Craticula (Cra) x x
Cyclotella (Cyc) x x
Cymbella (Cym) x x x
Cymbopleura (Cym.1) x x x x
Denticula (Den) x x x
Diatoma (Dia) x x x x
Diploneis (Dip) x x x
Encyonema (Enc) x x x x
Encyonopsis (Enc.1) x x x
Epithemia (Epi) x
Eucocconeis (Euc) x x x x
Eunotia (Eun) x
Fallacia x
Fistulifera x
Fragilaria x x
Frustulia (Fru) x
Geissleria
Gomphonema (Gom) x
Gyrosigma x
Halamphora (Hal) x x x
Hannaea (Han) x x
Hantzschia (Han.1) x x x x
Humidophila (Hum) x x x
Hygropetra (Hyg) x
Kobayasiella (Kob) x x x
Luticola (Lut) x x x x
Mayamaea (May) x x x
Melosira x
Microcostatus x
Muelleria (Mue) x x
Navicula (Nav) x x x x
Neidium (Nei) x x x
Nitzschia (Nit) x x x x
Orthoseira (Ort) x
Pinnularia (Pin) x x x x
Placoneis (Pla) x
Planothidium (Pla.1) x
Psammothidium (Psa) x x x x
Pseudostaurosira (Pse) x
Rossithidium (Ros) x x x x
Sellaphora (Sel) x
Simonsenia x
Stauroforma (Sta) x
Stauroneis (Sta.1) x x x
Staurosira (Sta.2) x
Staurosirella (Sta.3) x
Stephanodiscus (Ste) x
Surirella (Sur) x x x
Tabellaria (Tab) x
Table 2. List of diatom genera (with abbreviations) from cryoconite holes (C), Hørbye lakes (H),
Nordenskiöld lakes (N) and Retrettøya ponds (O).
DIATOMS FROM CRYOCONITE HOLES IN SPITSBERGEN
124
Nonmetric multidimensional scaling
(NMDS) of lake and cryoconite hole com-
munities revealed a strong separation be-
tween the two habitat types, both on
NMDS axis 1 and 2 (see Fig. 5). Cryo-
conite hole diatom communities had greater
proportions of the genera Eunotia (not
visible on figure), Aulacoseira Thwaites,
and Gomphonema, all of which strongly in-
fluenced sample orientation on both NMDS
axes. Lake habitats were strongly influ-
enced by Adlafia Moser, Lange-Bertalot
and Metzeltin, Encyonema Kützing, and
Mayamaea Lange-Bertalot on NMDS axis
2. Habitat differences were further influ-
enced by Staurosira Ehrenberg, Stauro-
sirella, Luticola, Pinnularia and Hantzschia
Grunow on axis 1 for the cryoconite holes,
and Encyonema, Cymbella Agardh, Diato-
ma Bory de Saint-Vincent, and Denticula
Kützing for the lakes. The Bray-Curtis
cluster analysis separated communities
into several distinct groups, of which, one
exclusively consisted of all samples from
the cryoconite holes (see Fig. 6). When the
community data were compared with
PERMANOVA, cryoconite hole diatom
communities were significantly different
from the pooled lake samples (df = 43,
F = 15.64, R2 = 0.27, p = 0.001).
Fig. 6. Bray-Curtis cluster analysis. C1–C9 for cryoconite holes, H1–H20 for Hørbye lakes,
N1 – N4 for Nordenskiöld lakes, and O1–O11 for Retrettøya.
Despite such clear split of both habitat
types, some genera were present between
both localities studied as seen in the
‘inkspot’ plot. This diagram, which can be
used to visualize the community structure
among samples using the raw relative
abundance data (Fig. 7), resulted in a clear
separation of three groups. The first con-
sisted entirely of samples from cryoconite
holes. The second group linked both local-
P. VINŠOVÁ et al.
125
ities of the Hørbye and Nordenskiöld mo-
raine kettle lakes. The third group con-
sisted entirely of samples of Retrettøya.
Some diatom genera showed clear differ-
ences between these three groups: e.g.
Encyonema and Adlafia were both abun-
dant in the third group. Nitzschia, although
prevailing in the entire dataset, was clearly
less abundant in the samples of the second
group. Psammothidium was almost equally
abundant in the first two groups, and,
finally, Pinnularia, Luticola, Staurosirella,
together with other genera (lower part of
Fig. 7), clearly separated the first group
from the remaining two.
Fig. 7. An ‘inkspot’ plot visualizing the diatom community structure among samples using relative
abundance data. Separation of three groups is shown. C1–C11 for cryoconite holes, H1–H20 for
Hørbye lakes, N1 – N4 for Nordenskiöld lakes, and O1–O11 for Retrettøya ponds.
DIATOMS FROM CRYOCONITE HOLES IN SPITSBERGEN
126
Discussion
Although numerous studies have re-
ported the presence of diatoms in cryo-
conite holes, many questions remain as to
their origin, viability, community struc-
ture, and assembly. Here, our aim was to
describe and compare the diatom assem-
blages from cryoconite holes of Norden-
skiöld glacier to the communities from
lake habitats in the immediate vicinity
which might serve as a potential source.
We hypothesized that if diatoms are being
transported from surrounding aquatic habi-
tats to cryoconite systems, then cryoconite
communities should be highly similar to
the communities from the habitat of origin.
We found that the diatom communities
of the cryoconite hole sediments were dis-
tinctly different from those from the adja-
cent lake habitats. Not only did cryoconite
sediment contain different genera not ob-
served in lake habitats, but often a higher
number of genera was observed in the cryo-
conite communities compared to the lake
sediment samples, even when only a frac-
tion of the amount of valves was counted.
This may suggest that (i) cryoconite hole
diatom communities have a unique struc-
ture, albeit at low cellular densities, or that
(ii) these cryoconite communities are not
self-sustaining and are dependent on dis-
persed cells, but are derived from more
varied sources than our surveyed lakes.
If these communities were actively re-
producing, it could be argued that this
should be reflected in a community domi-
nated by only a few species or genera. On
the contrary, our results show the presence
of a lot of different genera with a low num-
ber of species and valves within the cryo-
conite holes, which suggest that these as-
semblages are more likely to be formed as
a result of aeolian dispersion and deposi-
tion. While there is much left to learn about
these extreme environments, our study adds
further evidence that the diatom flora from
cryoconite holes is unique and distinct
from adjacent freshwater habitats. Recent-
ly, there is a need for further study that
could help in developing a more complete
understanding of diatom biogeography, dis-
persal, and evolution.
Ecology of individual diatoms
Our results showed that the cryoconite
diatom community composition exhibited
only little similarity to any of the lake dia-
tom communities in the vicinity, despite
the fact that 25 genera occur in both habi-
tat types. These shared genera may indi-
cate that these lakes contribute to the dia-
tom community in the cryoconite holes
(though investigation at a finer resolution
would be necessary to support this claim).
On the other hand, it is clear that at least
one other source is necessary to explain
the higher number of genera in the Norden-
skiöld glacier cryoconite holes, a number
of which are rare or uncommon to the sur-
rounding lakes. Similar results have been
reported by Stanish et al. (2013), who found
that cryoconite hole communities were dis-
tinctly different from adjacent stream habi-
tats in the McMurdo Dry Valleys (though
they certainly shared more taxa than in our
study). Another supporting data come from
Edwards et al. (2013c), who found that
cryoconite bacterial communities signifi-
cantly differed from those from the glacier
margins in Svalbard.
A number of diatom genera found in
cryoconite holes, including Hantzschia,
Humidophila Lowe et al., Orthoseira
Thwaites, Pinnularia and Luticola, are
typically aerophilic genera, known to
thrive mainly in moist terrestrial soil or
moss habitats. Diatom cells from soils can
be easily transported by wind together
P. VINŠOVÁ et al.
127
with sediment or volcanic dust (Dagson-
Waldhauserová et al. 2015), or be attached
to small moss patches able to overgrow
small cryoconite holes and thus becoming
a ‘glacial mouse’ (Vonnahme 2014).
Pinnularia borealis and Hantzschia am-
phioxys (Ehrenberg) Grunow, two species
highly abundant in cryoconite samples, are
frequently reported from polar soils and
also commonly recorded from mosses
(Beyens 1989, Van de Vijver et al. 2003,
Vinocur et Maidana 2010). Terrestrial dia-
toms make good candidates for cryoconite
colonization because they are more able to
resist freezing and desiccation than
freshwater species (Souffreau et al. 2010,
Yallop et Anesio 2010). Abrupt freezing
can be lethal for diatom vegetative cells,
especially for non-terrestrial species (Souf-
freau et al. 2010, 2013). Furthermore, some
of these diatoms could be present in rest-
ing stages, increasing thus the ability to
survive freezing stress (Souffreau et al.
2013).
In our study, rather than more, few
genera such as Adlafia, Encyonema, Navi-
cula and Nitzschia, were present in greater
relative abundances from the Retrettøya
proglacial site, in contrast to the other two
lake localities. According to our field ob-
servations, no direct hydrological connec-
tion exists between ponds on Retrettøya
and the glacier. However, plenty of fresh
glacial sediment, including cryoconite ma-
terial, was present drying on the margin
and mobilized by wind, which blows from
the glacier towards the fjord. Consequent-
ly, the Retrettøya ponds might be supplied
by cryoconite communities, similar as sug-
gested by Vonnahme (2014) for Hørbye
glacier and proglacial ponds in its frontal
moraine.
Species of the genera Achnanthidium
Kütz., Denticula Kütz., Encyonema, and
Eucocconeis Cleve had relatively high
abundances in moraine kettle lakes. These
genera were uncommon or rare in the cryo-
conite samples, most likely suggesting their
preference for more established (aquatic)
habitats. The A. minutissimum (Kütz.)
Czarnecki complex reaches high abun-
dances in various freshwater habitats
across Petuniabukta and actually presents
the most common freshwater diatom com-
plex in the area (Pinseel 2014). Although
this genus was rarely found in the cryo-
conite holes sampled for diatoms (this
study), fresh cryoconite material from the
same area observed by Vonnahme (2014)
in the field laboratory revealed a few via-
ble individuals of Achnanthidium. Psam-
mothidium is another very interesting ge-
nus that connects the cryoconite holes to
the moraine kettle lakes of Nordenskiöld
and Horbye. Moreover, species of this ge-
nus prefer habitats with sandy bottoms
(Round et Bukhtiyarova 1996). This is in
contrast with another highly abundant ge-
nus, Nitzschia, which dominates both the
cryoconite habitats and the Retrettøya
ponds, but usually occurs in lower abun-
dances in the kettle lakes.
Luticola was a rather common genus in
cryoconite holes involved into our study.
Previously, L. ventricosa (Kütz.) Mann
and L. nivalis (Ehrenberg) Mann have
been retrieved alive from frozen cryo-
conite material (Yallop et Anesio 2010).
Luticola nivalis was also observed in the
present study, together with L. frequentis-
sima (see Fig. 3). Species of the genus
Muelleria (Frenguelli) Frenguelli are not
abundant in the Arctic, and only one spe-
cies resembling the cosmopolitan M. ter-
restris (Petersen) Spaulding & Stoermer
was observed in this study. The species
was previously reported from cryoconite
holes by Yallop et Anesio (2010). Muel-
leria is, however, one of the most domi-
nant genera of Antarctic cryoconite diatom
communities, accompanied by species from
genera Humidophila and Luticola (Mueller
et al. 2001, Van de Vijver et al. 2010,
Stanish et al. 2013, Kohler et al. 2015).
DIATOMS FROM CRYOCONITE HOLES IN SPITSBERGEN
128
Insights into microbial dispersal
While we did not check for cell via-
bility in our samples due to the limited
amount of material, we can postulate that
viability may have been low, as recovered
valves were often broken in addition to be-
ing sparse. However, earlier studies have
suggested that at least some cryoconite di-
atom cells are viable. Stanish et al. (2013)
and Vonnahme (2014) have both reported
viable valves by microscopic analysis, and
Yallop et Anesio (2010) were able to cul-
ture 27 diatom genera from cryoconite ma-
terial, even after being frozen for 1-2 years.
Vonnahme (2014) analyzed only a few
samples (n = 3) of fresh material from cryo-
conite holes of Nordenskiöld and Hørbye
glaciers for diatoms content, and reported
following genera and species as living,
although in low numbers: Achnanthidium
sp., Encyonopsis laevis Nägeli and E. sub-
minuta Krammer & E. Reichardt, Humido-
phila sp., Pinnularia cf. obscura, and
Psammothidium cf. marginulatum (Gru-
now) Bukhtiyarova and Round. All of these
were also identified in our study of fixed
material, albeit present in various abun-
dances.
In Antarctica, some diatom species,
such as Muelleria cryoconicola Stanish &
Spaulding (Van de Vijver et al. 2010),
Luticola bradyi Kohler, and L. spainiae
Kohler & Kopalová are thus far found
almost exclusively in cryoconite holes, and
have been suggested to be endemic to
these habitats. According to Kohler et al.
(2015), transportation of diatom cells from
cryoconite holes to surrounding aquatic
habitats might even take place, as sug-
gested for L. bradyi, a species found in a
large population in cryoconite material,
but only very rarely observed in glacial
meltwater streams. These above observa-
tions, together with the differences in cryo-
conite diatom communities from marginal
habitats, provide evidence that while dia-
toms may be seeded from adjacent habi-
tats, at least some survive, live, and pos-
sibly reproduce while in cryoconite holes.
While unique cryoconite diatom communi-
ties are thought to develop in stable cryo-
conite systems found in Antarctica (Sta-
nish et al. 2013), Arctic systems are much
more exposed to the outside world and do
not persist as long due to glacier hydrolo-
gy and melting. Despite this, they can still
be colonized by living diatoms (Yallop et
Anesio 2010), and these habitats might
subsequently select the most adapted spe-
cies to live in.
In our study, many genera observed in
the cryoconite material were rare in the
lakes and ponds of Petuniabukta. For ex-
ample, the genera Aulacoseira, Tabellaria
Ehrenberg, Melosira Agardh, Orthoseira,
and Stephanodiscus Ehrenberg were pres-
ent in the cryoconite holes, but have never
been observed in the lakes and ponds of
Petuniabukta (> 50 lakes studied, in
Pinseel 2014). One Gomphonema taxon
(i.e. Gomphonema aff. nathorstii, Fig. 3)
was recorded as common in almost all
(eight out of nine) cryoconite samples but
was visually absent from the surrounding
lakes. However, the same species has been
observed previously in a temporal pond on
a mountain top in Mimerdalen (Pinseel,
unpublished data), suggesting that this tax-
on prefers emphemeral habitats. Several
other species of Gomphonema were pres-
ent within the cryoconite samples, al-
though absent from nearby lakes, and the
same is true for species of Staurosira,
Staurosirella and Eunotia. Some genera
(e.g. Achnanthidium, Gomphonema, Psam-
mothidium, Staurosirella) may produce
(mucilaginous) stalks (Potapova 2009,
Gesierich et Rott 2012) so they can be at-
tached to solid objects, hypothetically fa-
voring them in aeolian dispersion. It was
striking that no marine species have been
identified from the cryoconite material, de-
spite the immediate vicinity of the sea,
even though direct evidence of marine dia-
tom deposition on ice sheets was previ-
P. VINŠOVÁ et al.
129
ously reported before from the Antarctic
(Budgeon et al. 2012). This is probably be-
cause the prevalent wind direction is in the
opposite way, which reduces the extent of
marine deposition of diatoms compared to
those that are limnoterrestrial in origin.
As light microscope studies of cryo-
conite material are challenging, it is pos-
sible that the lack of records of diatom
species inhabiting cryoconite holes is a
result of under-sampling, as previously
hypothesized by Yallop et Anesio (2010).
In total, some 84 taxa were identified in
our study, and broader sampling efforts
would certainly reveal more taxa. Our
gradient sampling also revealed that sites
closer to the side of the glacier contained
less diatom valves than those situated
more inside. This could further support the
suggestion of aeolian input of cells rather
than the cell input from avalanches of
eroded material that occurs on glaciers
sides (Landford et al. 2011). On the con-
trary, sites that were situated the most to
the center of the Nordenskiöld glacier (C8
and C9) contained less (roughly half) of
the average counted valves. It might be
interesting to sample across the whole ab-
lation zone of the glacier by adding further
sampling sites, as it could reveal wind
patterns on a local scale (i.e. considering
katabatic winds).
Future directions and conclusions
Contrary to the Antarctic diatom flora,
which has been recently revised based on a
more narrow morphology-based species
concept (ex. Van de Vijver et al. 2010a,
2011, 2013, Kopalová et al. 2012, 2013,
Taylor et al. 2014, among others), the Arc-
tic diatom morphological taxonomy is in-
sufficient at this time to make fine-scale
comparisons between habitats. Moreover,
Pinseel (2014) have identified a high num-
ber of new diatom taxa ready to be de-
scribed. However, in this study, we found
the genus-level to be of great use to com-
pare cryoconite holes with adjacent fresh-
water habitats, and it became clear that our
comparison set of localities in the immedi-
ate vicinity was not broad enough to fully
assess ‘the source’ of the cryoconite diatom
flora. It is possible that a more broad sam-
pling effort to include non-aquatic habitats
nearby, as well as more distant localities
such as Iceland (Dagsson-Waldhauserova
et al. 2015) are necessary to discern the
cryoconite diatom flora ‘source’. In any e-
vent, the diatom communities of cryo-
conite holes have the potential to inform
researchers about microbial dispersal pat-
terns through comparing the regional distri-
bution of diatom taxa in between distinct
polar habitats, and deserve further study in
our investigation of the cryosphere.
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