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Cryoconite holes are small, extreme habitats, widespread in the ablation zones of glaciers 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 communities 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, cryoconite 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.
Content may be subject to copyright.
CZECH POLAR REPORTS 5 (2): 112-133, 2015
Received November 4, 2015, accepted December 21, 2015.
*Corresponding author: Petra Vinšová <>
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.
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
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.
Key words: ecology, limnology, lakes, cryosphere, Svalbard, polar region
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-
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
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: or 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.
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.
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
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
P. VINŠOVÁ et al.
Study code
(sample code)
Lake type
a.s.l.(m) GPS N GPS E
Cryoconite holes
C1 (N002)
surface 0.5 1 8.30
130 78° 38' 24.7"
16° 58' 37.1"
C2 (N003)
surface 0.7 1 8.60
201 78° 38' 24.6"
17° 00' 05.1"
C3 (N004)
surface 0.6 1 8.30
244 78° 38' 22.2"
17° 01' 44.6"
C4 (N006)
surface 0.5 1 9.10
278 78° 38' 28.3"
17° 05' 11.0"
C5 (N007)
surface 0.4 7 9.30
361 78° 38' 29.2"
17° 06' 37.6"
C6 (N008)
surface 0.4 4 9.80
393 78° 38' 30.5"
17° 07' 22.7"
C7 (N009)
surface 0.3 1 8.50
262 78° 39' 21.4"
17° 03' 40.9"
C8 (N010)
surface 0.4 2 8.40
263 78° 39' 01.0"
17° 03' 31.9"
C9 (N011)
surface 0.5 12 10.10
267 78° 38' 41.5"
17° 03' 16.2"
C10 (N012)
surface 0.5 2 9.20
237 78° 38' 08.6"
17° 03' 01.0"
C11 (N013)
surface 0.4 11 8.80
233 78° 37' 51.9"
17° 02' 55.3"
C12 (N005)
surface 0.5 2 9.00
271 78° 38' 25.6"
17° 03' 46.9"
Nordenskiöld lakes (Pinseel 2014)
N1 (SP20)
lakes 8.3 443 8.52
28 78° 38' 19.2''
16° 49' 35.8''
N2 (SP21)
lakes 8.6 329 8.55
24 78° 38' 19.4''
16° 49' 41.1''
N3 (SP22)
lakes 8.4 658 8.33
29 78° 38' 17.6''
16° 50' 1.5''
N4 (SP23)
lakes 8.7 566 8.60
26 78° 38' 17.6''
16° 50' 5.1''
Hørbye lakes (Pinseel 2014)
H1 (SPH1B)
lakes ND ND ND 37 78° 44' 05.7"
16° 26' 52.5"
H2 (SPH1C)
lakes 10.9 861 8.18
23 78° 44' 08.1"
16° 26' 51.9"
H3 (SPH2A)
lakes 11.7 461 8.39
41 78° 44' 13.4"
16° 26' 27.7"
H4 (SPH2B)
lakes 12.0 760 8.13
43 78° 44' 11.2"
16° 26' 15.2"
H5 (SPH2C)
lakes 11.6 756 8.20
44 78° 44' 11.8"
16° 26' 10.1"
H6 (SPH3A)
lakes 13.3 338 8.41
49 78° 44' 22.0"
16° 25' 04.8"
H7 (SPH3B)
lakes 12.7 306 8.42
50 78° 44' 23.9"
16° 24' 56.3"
H8 (SPH3C)
lakes 14.2 339 8.54
46 78° 44' 21.4"
16° 25' 08.2"
H9 (SPH4B)
lakes 11.3 423 8.28
52 78° 44' 34.2"
16° 24' 42.4"
H10 (SPH4C)
lakes 14.5 655 8.32
50 78° 44' 34.2"
16° 24' 48.7"
H11 (SPH5A)
lakes 13.9 1428 8.12
54 78° 44' 43.9"
16° 24' 27.4"
H12 (SPH5B)
lakes 12.5 1805 8.18
57 78° 44' 45.4"
16° 24' 37.0"
H13 (SPH5C)
lakes 13.3 1208 8.32
57 78° 44' 47.2"
16° 24' 39.5"
lakes 11.6 1325 8.18
35 78° 44' 05.0"
16° 26' 34.1"
H16 (SP44)
lakes 12.7 180 8.45
40 78° 44' 13.0"
16° 26' 14.1"
H17 (SP45)
lakes 12.2 218 8.54
50 78° 44' 35.4"
16° 24' 44.9"
H18 (SP46)
lakes 9.2 198 9.20
106 78° 45' 14.0"
16° 21' 46.2"
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)
related 8.2 592 8.80
20 78° 39' 24.9''
16° 54' 46.5''
O2 (SP63)
related 8.1 256 9.60
7 78° 39' 27.8''
16° 54' 34.3''
O3 (SP64)
related 8.5 377 8.90
24 78° 39' 23.0''
16° 54' 44.4''
O4 (SP65)
related 8.1 281 9.30
18 78° 39' 19.3''
16° 54' 37.8''
O5 (SP66)
related 7.4 510 8.70
20 78° 39' 21.4''
16° 54' 39.9''
O6 (SP67)
related 8.0 319 8.70
28 78° 39' 18.6''
16° 54' 38.5''
O7 (SP68)
related 8.4 174 9.80
24 78° 39' 16.3''
16° 54' 21.9''
O8 (SP69)
related 9.0 347 9.20
7 78° 39' 8.5'' 16° 54' 51.4''
O9 (SP70)
related 9.0 450 9.40
11 78° 39' 14.6''
16° 55' 28.5''
related 9.1 147 9.60
26 78° 39' 14.1''
16° 55' 16.4''
O11 (SP73)
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.
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
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).
Fig. 2. Examples of cryoconite holes (A–B), and an overview of the Nordenskiöld tide-water
glacier (C).
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.
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).
Fig. 4. Genus richness of cryoconite holes (C), rbye 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.
Genus C
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
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).
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.
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.
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.
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.
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
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).
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-
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.
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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... Such habitats often harbor a very specialized biological community dominated mostly by cyanobacteria, but with an important contribution from microalgae, heterotrophic bacteria, fungi, and viruses (Hodson and others, 2008), and in some cases also small metazoans. Eolian transportation of local terrestrial or aquatic inoculi are believed to be important contributors to the communities that establish (Cameron and others, 2012;Vinšová and others, 2015), forming assemblages that have been compared to the truncated food webs seen in Antarctic lakes (Säwström and others, 2002), with relatively high alpha diversity levels but scarce number of individuals throughout the water column (Cameron and others, 2012). The microhabitat appears to favor species that can withstand the extreme changes in physical and chemical conditions characteristic of such harsh conditions (Spaulding and others, 2010). ...
... As of 2015, 370 microorganism taxa had been reported from cryoconite holes worldwide, 62 of them belonging to different algal groups (Kaczmarek and others, 2016), Diatoms are one of the frequently reported component, of a different floristic composition compared to the surrounding aquatic environments (Stanish and others, 2013). Particularly, cryoconite holes are known to support aerophytic, halophytic, epipelic and bryophilic diatom species, suggesting multiple origins of colonizing cells (Yallop and Anesio, 2010;Vinšová and others, 2015), with assemblages comparable to those found in acidic lakes and moss-inhabiting communities reported from high-latitude locations (Yallop and Anesio, 2010). Most studies describing cryoconite microbiota have been performed in Arctic/Antarctic glaciers, and to date little attention has been paid to equivalent habitats in the Equatorial region. ...
... Low-latitude glaciers are generally smaller and affected by larger diurnal irradiance fluctuations, being also more strongly linked to the terrestrial surrounding land, including anthropogenic perturbations (Cook and others, 2016). Harsh climatic conditions, often combined with low nutrient concentrations and extreme UV radiation levels, usually impede biological processes (Mueller and others, 2001) even though cryoconite holes may harbor true microbial communities structurally and functionally independent from those in their surroundings (Edwards and others, 2013;Vinšová and others, 2015). In any case, little is known about biogeochemical processes within such habitats (Foreman and others, 2007), the processes governing the emergence of a functional communities (Edwards and others, 2013) and the effects of environmental variables on cryoconite communities. ...
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In the ablation zone of glacier habitats, cryoconite holes are known to harbor diverse microbial communities, including unique diatom floras distinct from those of surrounding aquatic and terrestrial systems. Besides descriptive studies, little is known about the diversity of cryoconite dia-toms and their response to environmental stressors, particularly in low-latitude glaciers. This paper documents an extremely diversified diatom community in Antisana Glacier (Ecuador), reporting 278 taxa found in 54 surface holes, although with low individual abundances. Contrary to our expectations, assemblage structure did not respond to water physical or chemical characteristics, nor to cryoconite hole morphology, but to elevation. We demonstrate that elevation is a driver of diatom assemblages. Both alpha diversity (measured as Fisher's index) and species richness (corrected for unequal sample sizes) correlated negatively with elevation, suggesting a replacement toward simplified, poorer communities along this gradient. The taxonomic composition also changed significantly, as revealed by multivariate statistics. In summary, cryoconite holes are sites of high taxonomic diversity composed of taxa that are allochthonous in origin.
... Benthic diatoms, including a high abundance of Psammothidium and Achnanthidium species, exist in supraglacial cryoconite holes on the surface of Antarctic and Arctic glaciers(Yallop and Anesio 2010;Stanish et al 2013;Vinšová et al 2015). Thus, diatom assemblages of GF lakes along the GrIS could be dominated by supraglacial communities arriving via meltwater. ...
Arctic and alpine areas experience rapid environmental changes that are altering nutrient delivery to lakes. I review the current state of lake ecosystem subsidization research and summarize Arctic and alpine lake subsidies, their ecological importance, and ways they are changing. I identify current knowledge gaps in Arctic and alpine lake subsidization research and highlight the importance of Arctic and alpine lakes for ecosystem subsidy research. Meltwater from the Greenland Ice Sheet (GrIS) exports sediment and nutrients to lakes, but the ecological effects of this subsidy remain unclear. To assess the effects, four glacially fed (GF) lakes that receive GrIS meltwater were compared to snow and groundwater fed (SF) lakes. Total phosphorus (TP) was six times higher in GF lakes compared to SF lakes, but biological and chemical assays called into question its bioavailability. GF lakes had higher algal biomass, which may be due to moderately higher DIN in GF lakes compared to SF lakes. Increased nitrogen (N) deposition rates over the past century have affected North American and European alpine lakes by shifting nutrient limitation status and increasing algal biomass. I evaluated predictors of alpine and subalpine lake sensitivity across North American and European mountain ranges. North American lakes were more sensitive to N deposition than European lakes. Local-scale analyses revealed distinction of lake sensitivity controls, such as land cover, bedrock geology, maximum lake depth, and elevation. Altered Arctic lake thermal stratification may lead to hypolimnetic anoxia and sediment phosphorus release. To assess this, I investigated Greenland lakes that exhibit different stratification patterns. Among anoxia-susceptible lakes, morphometric and stratification strength metrics were not able to explain variation of anoxia. Cooler spring temperatures were associated with mid-summer lake anoxia. Increased TP concentration and algal biomass were associated with lake anoxia. My dissertation research summarizes how environmental change is altering lake nutrient subsidization and cycling in remote Arctic and alpine areas. It highlights ecological responses to those alterations. With accelerating rates of warming and increasing N deposition rates in Arctic and alpine areas, this research provides timely insights of how lakes located in cold, remote areas will change in the coming century.
... The high proportion of GF lake benthic diatom abundance may be because of increased diatom productivity in the shallow littoral zones and relatively less productivity in the turbid water columns of GF lakes. Benthic diatoms, including a high abundance of Psammothidium and Achnanthidium species, exist in supraglacial cryoconite holes on the surface of Antarctic and Arctic glaciers (Stanish et al. 2013;Vinšová et al. 2015;Yallop and Anesio 2010). Thus, diatom assemblages of GF lakes along the GrIS could be dominated by supraglacial communities arriving via meltwater. ...
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Meltwater discharge from the Greenland Ice Sheet (GrIS) exports sediment, solutes, total phosphorus (TP), dissolved inorganic nitrogen (DIN), and other macro-and micronutrients to associated aquatic ecosystems. It remains unclear how this meltwater affects the ecology of glacially fed (GF) lakes. We assessed a suite of physical, chemical, and biological features of four GF lakes, and compared them to those of four nearby snow-and groundwater-fed (SF) lakes. We found that TP concentrations were six times higher in GF compared to SF lakes, but microbial extracellular enzyme activities and aluminum, iron, and phosphorus sediment fractions suggested that much of this TP in GF lakes is likely not biologically available. Turbidity was fifteen times higher in GF lakes, and DIN was twice as high than in SF lakes, but these nitrogen differences were not significant. While diatom species richness did not significantly differ between lake types, GF lakes had higher water column chlorophyll a (Chl a). Diatom species distributions across all lakes were strongly associated with turbidity, TP, and dissolved organic carbon (DOC). While cosmopolitan diatom taxa such as Discostella stelligera were found in both lake types, diatom communities differed across lake types. For instance, Fragilaria and Psammothidium species dominated GF lakes, while Achnanthes species and Lindavia ocellata were dominant in SF lakes. In addition to turbidity, the moderate amounts of DIN in GF lakes may play an important role in shaping diatom communities. This is supported by the high abundance in GF lakes of taxa such as Fragilaria tenera and D. stelligera, which reflect nitrogen enrichment in some lakes. Our results demonstrate how GrIS meltwaters alter the ecology of Arctic lakes, and contribute to the growing body of literature that reveals spatial variability in the effects of glacial meltwaters on lake ecosystems.
... amphioxys and P. cf. borealis have also been found in ice cores from high-latitude settings (Donarummo et al. 2003, Harper & McKay 2010. Pinnularia and Luticola are two common genera found in Swedish cryoconite holes (Vinšová et al. 2015). Vinšová and colleagues reported that these genera made up 12% and 5% of their samples, respectively, and P. borealis was one of the most common species encountered. ...
Diatoms in ice cores have been used to infer regional and global climatic events. These archives offer high-resolution records of past climate events, often providing annual resolution of environmental variability during the Late Holocene. Recently, the first low-latitude tropical diatoms were described from the Quelccaya Summit Dome. Here, we document diatoms observed in ice cores from Quelccaya, spanning AD 1300 to 1815, along with those from two additional glaciers (Coropuna, and Sajama glaciers) in the tropical Andes, spanning AD 1764 to 1814. Diatom assemblages recovered from these three sites were rare, but differ in abundance and species composition through time. Assemblages are characterized by cosmopolitan and aerophilic species, mostly pennate diatoms. There were 44 taxa in all, with Pinnularia cf. borealis Ehrenberg being the most common species encountered in the samples. Eleven taxa were found at all three sites. Both Coropuna and Sajama had taxa that were unique to these locations, whereas Quelccaya had no unique taxa. Due to the rarity of diatoms and the cosmopolitan nature of the dominant species, it is not possible to determine their origin, limiting their utility in paleoclimate reconstructions.
... Nordenskiöldbreen is a tidewater outlet polythermal type of glacier with an area of 242 km 2 , located in the northern part of the Billefjorden fjord (Hagen et al. 1993). Further information on the study site and glacier retreat can be found in Plassen et al. (2004), Rachlewicz et al. (2007, van Pelt et al. (2012), Stacke et al. (2013), Vinšová et al. (2015. The main emphasis was given to a land terminating part and a 250-m-long section of a proglacial river on the northwestern flank of the Nordenskiöldbreen snout. ...
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Unmanned Aerial Vehicles (UAV) have technical capabilities to extended usage in various fields of science. The existing UAVs are to be relatively easily accessible in the near future. It is possible to equip them with different sensors but there are still some usage limitations. This paper focuses on demonstrating UAVs usage for research in polar regions. The research in polar regions is very specific and, due to harsh climate, limits the field work with UAVs. The options and limitations are presented in a case study performed in the Nordenskiöldbreen area, Svalbard Archipelago. In the end some derived products suitable for further analysis are presented.
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Direct measurements of glacier hydrological processes are usually restricted to short periods and a limited number of sites due to logistical, financial, and meteorological constraints. As a result, the indirect study of glacier hydrology through remote sensing has gained traction while the accessibility of high-resolution publicly available remote sensing data has also increased. By quantifying the areal extents of key dynamic features of a tidewater glacier (i.e. the evolution of sea ice, supraglacial lakes, meltwater plumes) as proxies of its hydrological cycle using Sentinel-2 observations, a simple alternative amidst the outlined logistical constraints is potentially available. Here we demonstrate the usefulness of Sentinel-2 satellite images as a simple and accessible tool with high temporal coverage for studying glacier hydrology. To make this case for the Nordenskiöld tidewater glacier, the evolution of its supraglacial hydrological system and respective meltwater plumes areal extents were monitored for the 2016-2020 melting seasons. Hydrological connectivity of supra-and subglacial systems and the resulting meltwater plumes are illustrated. Meltwater is stored on the glacier surface at the beginning of the melt season (June) which is observed through the filling of the supraglacial lakes. The stored meltwater is later released (June/July), probably through englacial conduits and moulins, and consequently reaches the subglacial drainage system. The resulting occurrence of meltwater plumes clearly indicates the latter in Adolfbukta at the glacier terminus. This signals the transport of significant volumes of water in contact with the glacier bed. The meltwater plume activity peaks during late July, and its appearance continues until mid-September. The duration of the glacier melt season is reflected through the filling of supraglacial lakes and later in the appearance of meltwater plumes. The temporal pattern of the hydrologic processes is relatively uniform during the study period, contrasting the large variability of sea ice cover duration. The observed behavior of Nordenskiöld's supraglacial lakes is in good agreement with similar tidewater glaciers in Svalbard.
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Cryoconite holes, supraglacial depressions containing water and microbe-mineral aggregates, are known to be hotspots of microbial diversity on glacial surfaces. Cryoconite holes form in a variety of locations and conditions, which impacts both their structure and the community that inhabits them. Using high-throughput 16S and 18S rRNA gene sequencing, we have investigated the communities of a wide range of cryoconite holes from 15 locations across the Arctic and Antarctic. Around 24 bacterial and 11 eukaryotic first-rank phyla were observed in total. The various biotic niches (grazer, predator, photoautotroph, and chemotroph), are filled in every location. Significantly, there is a clear divide between the bacterial and microalgal communities of the Arctic and that of the Antarctic. We were able to determine the groups contributing to this difference and the family and genus level. Both polar regions contain a “core group” of bacteria that are present in the majority of cryoconite holes and each contribute >1% of total amplicon sequence variant (ASV) abundance. Whilst both groups contain Microbacteriaceae, the remaining members are specific to the core group of each polar region. Additionally, the microalgal communities of Arctic cryoconite holes are dominated by Chlamydomonas whereas the Antarctic cryoconite holes are dominated by Pleurastrum . Therefore cryoconite holes may be a global feature of glacier landscapes, but they are inhabited by regionally distinct microbial communities. Our results are consistent with the notion that cryoconite microbiomes are adapted to differing conditions within the cryosphere.
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Introduction The polar regions, both Arctic and Antarctic, show strong evidence of climate change affecting freshwater species, communities, and ecosystems, and are expected to undergo rapid and continued change in the future (IPCC, 2007). Diatoms in the freshwater and brackish habitats of inland waters of the Antarctic provide valuable records of their historic and modern environmental status. Antarctic habitats also contain a unique biodiversity of species many of which are found nowhere else on Earth. In this chapter, we review investigations using diatoms as indicators of environmental change in Antarctic and subantarctic island habitats, including lakes and ponds, streams and seepage areas, mosses and soils, cryoconite holes, brine lakes, and remarkable subsurface glacial lakes. The Antarctic continent holds the vast majority of the Earth’s freshwater, but the water is largely inaccessible because it is in the form of ice. Life is dependent upon liquid water, a substance scarce in Antarctica. Less than 0.4% of the continent is ice free, and it is within these ice-free regions that freshwater lakes and ephemeral streams form, fed by the melting of snow and glacial ice and occasional precipitation. These ice-free regions are located primarily near the Antarctic coastline (Figure 14.1). Of these regions, the “desert oases” of East Antarctica are considered to be the coldest, driest regions on Earth. In the limited parts of these oases where liquid water is available, even if present for only a few short weeks of the year, there is life (McKnight et al., 1999).
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A revision of the freshwater diatom genus Luticola from the McMurdo Sound Region, including the McMurdo Dry Valleys and Cape Royds, Antarctica, was made to contribute to a consistent flora for the entire Antarctic Region. Detailed light and scanning electron microscopic observations, review of pertinent literature, and examination of historical and type material lead to the identification of 12 Luticola species. Four new species and one new combination are proposed, including L. bradyi sp. nov., L. spainiae, sp. nov., L. macknightiae, sp. nov., L. transantarctica, sp. nov., and L. elegans, comb. nov. stat. nov. Several of these taxa were previously identified as part of the L. muticopsis (Van Heurck) D.G.Mann complex; Navicula muticopsis f. evoluta W. & G.S. West, L. muticopsis f. reducta (W. & G.S. West) Spaulding, and N. muticopsis f. capitata Carlson, or mistaken for the similar L. mutica (Kützing) D.G.Mann and L. cohnii (Hilse) D.G.Mann. Morphological features of all new species were compared to the closest morphologically similar taxa, and their ecology and biogeography are discussed. All Luticola species considered here show restricted Antarctic distributions, and 8 of the 12 reported species are known only from the Antarctic continent.
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Due to global change affecting glaciers worldwide, glacial streams are seen as threatened environments deserving specific scientific interest. Glacial streams from the Coast Range and Rocky Mountains in British Columbia and at the border to Alberta were investigated. In particular glacial streams and downstream sites in the Joffre Lakes Provincial Park, a near by mountain river and two large glacial streams in the Rocky Mountains (Kootenay Range, Jasper National Park) were studied. Regardless of a high variability of catchment glaciation (1 to 99%) thin organic biofilms with firmly attached diatom frustules of the genera Achnanthidium, Psammothidium, Encyonema, Gomphonema and fragilaroid taxa were found in ail cases. In spite of fundamentally different geological conditions between the Coast Range sites and the Rocky Mountain sites, the pioneer taxon Achnanthidium minutissimum (with a slimy long ecomorph) was dominating quantitatively in most of the glacier stream samples together with the rheobiontic Hannaea arcus. Individual glacier stream samples were characterized by the dominance of Achnanthidium petersenii and Gomphonema calcifugum/Encyonema latens. The diatom community analysis (cluster analysis) revealed the expected separation of glacier stream sites and sites of the lower segments of the river continuum (e.g., dominance of Diatom ehrenbergii in the mountain river). In the Joffre area, the total species richness of turbid glacial streams close to the glacier mouth was significantly lower than in the more distant sites. The two largest glacial streams in the Rocky Mountains showed divergent results with a remarkable high species richness (43 taxa) at the Athabasca River origin (Columbia Icefield) and low diversity in Illecillewaet river (9 km downstream the glacier mouth). From the biogeographical point of view the dominant taxa comprised mainly widespread pioneer species coping best with the unstable conditions, while the subdominant taxa comprised taxa specific for pristine arctic-alpine or high altitude habitats (e.g., Psammothidium grischunum). Almost 50% of the taxa were classified as oligo- to oligo-mesotraphentic and approximately 20% as endangered or extremely rare.
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Measuring microbial abundance in glacier ice and identifying its controls is essential for a better understanding and quantification of biogeochemical processes in glacial ecosystems. However, cell enumeration of glacier ice samples is challenging due to typically low cell numbers and the presence of interfering mineral particles. We quantified for the first time the abundance of microbial cells in surface ice from geographically distinct sites on the Greenland Ice Sheet (GrIS), using three enumeration methods: epifluorescence microscopy (EFM), flow cytometry (FCM), and quantitative polymerase chain reaction (qPCR). In addition, we reviewed published data on microbial abundance in glacier ice and tested the three methods on artificial ice samples of realistic cell (10(2)-10(7) cells ml(-1)) and mineral particle (0.1-100 mg ml(-1)) concentrations, simulating a range of glacial ice types, from clean subsurface ice to surface ice to sediment-laden basal ice. We then used multivariate statistical analysis to identify factors responsible for the variation in microbial abundance on the ice sheet. EFM gave the most accurate and reproducible results of the tested methodologies, and was therefore selected as the most suitable technique for cell enumeration of ice containing dust. Cell numbers in surface ice samples, determined by EFM, ranged from ~ 2 × 10(3) to ~ 2 × 10(6) cells ml(-1) while dust concentrations ranged from 0.01 to 2 mg ml(-1). The lowest abundances were found in ice sampled from the accumulation area of the ice sheet and in samples affected by fresh snow; these samples may be considered as a reference point of the cell abundance of precipitants that are deposited on the ice sheet surface. Dust content was the most significant variable to explain the variation in the abundance data, which suggests a direct association between deposited dust particles and cells and/or by their provision of limited nutrients to microbial communities on the GrIS.
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This paper presents the first results of measurements of global solar radiation, albedo, ground surface and 2-m air temperature, relative humidity, and wind speed and di- rection carried out in the central part of Spitsbergen Island in the period 2008-2010. The study site was located on the coastal ice-free zone of Petuniabukta (north-western branch of Billefjorden), which was strongly affected by local topography, character of the ground sur- face, and sea ice extent. Temporal analysis of the selected meteorological parameters shows both strong seasonal and inter-diurnal variation affected by synoptic-scale weather systems, channelling and drainage effects of the fjords and surrounding glaciers. The prevailing pat- tern of atmospheric circulation primarily determined the variation in global solar radiation, wind speed, ground surface and 2-m air temperatures. Furthermore, it was found that ther- mal differences between Petuniabukta and the nearest meteorological station (Svalbard Lufthavn) differ significantly due to differences in sea ice concentrations and ice types in the fjords during the winter and spring months.