Fottea, Olomouc, 14(1): 77-90, 2014
77
New species in the genus Muelleria (Bacillariophyta) from the Maritime
Antarctic Region
Bart V a n
de
V
i j v e r 1’2,
Ralitsa Z id a r o v a 3 & Katerina K o p a l o v á 4
1 Botanic Garden Meise, Department o f Bryophyta & Thallophyta, Domein van Bouchout, B-1860 Meise,
Belgium; ^Corresponding author e-mail: vandevijver@br.fgov.be
2University o f Antwerp, Department o f Biology, ECOBE, Universiteitsplein 1, B-2610 Wilrijk, Belgium
d epartm ent o f Botany, Faculty o f Biology, St. "Kliment Ohridski" University o f Sofia, 8 Dragan TzankovBlvd.,
Sofia 1164, Bulgaria
d epartm ent o f Ecology, Faculty o f Science, Charles University in Prague, Vinicná 7, CZ-128 44 Prague 2,
Czech Republic
A bstract: During a survey of the freshwater aquatic and limno-terrestrial diatom flora of the Maritime
Antarctic Region, six taxa belonging to the genus Muelleria that could not be identified were observed. Based
on detailed LM and SEM observations, these six taxa have been described as new taxa: Muelleria desseiniana
sp. no v .,M levkoviana sp. nov.,M nogae sp. nov.,M spauldingiana sp. no v., M subsabbei sp. nov. and AE
undulatoides sp. nov. Comments are made on their taxonomic position and how they can be distinguished from
other species in this genus. Brief notes on the ecology and distribution of the six taxa are added. A compiled list
of all Muelleria taxa described worldwide is included in this paper.
Key words: Antarctic Region, morphology, Muelleria, new species, taxonomy
I n t r o d u c t io n
The genus Muelleria is one of the most species rich
genera in the Antarctic Region despite its rather low
total number of described species ( S p a u l d in g et al.
1999; V a n d e V ij v e r et al. 2010). Up to now, only
29 taxa are recognized worldwide of which 16 have
a distribution restricted to the Antarctic Region.
Table 1 summarizes all collated published taxonomic
information on all Muelleria species described so
far worldwide. Together with the genera Diadesmis,
Luticola and Hantzschia, the genus is one of the
major constituents of the terrestrial (soil) diatom flora,
although populations rarely reach high abundances
( F e r m a n i et al. 2007; K o p a l o v á & V a n d e V ijv e r
2013). V a n d e V ij v e r et al. (2010) elaborately discuss
the success of the genus Muelleria in the Antarctic
Region which may be due to several factors such as
their strong preference for wet or semidry terrestrial
habitats which are abundantly present in this region.
But it is also likely that a comparable diversity within
this genus could be found in the Arctic Region and the
low number of recorded taxa from the Arctic could be
the result of undersampling and/or force-fitting.
Muelleria taxa are easily recognizable in light
microscope (LM) based on the presence of longitudinal
ribs bordering the axial area, typically (strongly)
deflected to even hooked proximal raphe endings
and central striae that are usually more distant than
the other striae in most species. Confusion with other
taxa is only possible with certain species of the genus
Neidium but the latter is very rare in the Antarctic
Region ( H a m il t o n et al. 2013).
The highest Muelleria diversity seems to be
concentrated on the South Orkney (SOI) and South
Shetland (SSI) Islands. S p a u l d in g et al. (1999) and V a n
d e V ij v e r et al. (2010) described nine new taxa from
Signy Island (SOI), Deception Island (SSI) and King
George Island (SSI). During an ongoing revision of the
Antarctic diatom flora of some islands in the Maritime
Antarctic Region, several very small populations
of Muelleria taxa were found on Livingston Island
(SSI) and James Ross Island. Based on LM and SEM
analysis of their structure, six of these taxa could not
be identified using the currently available literature and
are described here as new species. The relationship of
the new taxa in comparison with other Muelleria taxa
from the Antarctic Region and notes on their ecological
preferences are presented.
M
a t e r ia l s a n d
M
ethods
Epilithic, sediment and moss samples were collected from
various terrestrial and freshwater habitats in the following
islands in the southern Atlantic Ocean: Livingston Island
(South Shetland Islands) and James Ross Island (Table 1).
Samples were fixed in the field with 3% formaldehyde. More
information on sampling methods and physico-chemical
78
V
an de
V
ijv e r
et al: New species in the germs Muelleria
Table 1. List o f sam ples and their characteristics used in this paper.
Sample
Collection date Locality
GPS coordinates
Short description
JRI-S1J
22/02/2007
Komárek‘s slopes below
Berry Hill, Ulu Peninsula,
James Ross Island
(not determined)
seepage area
5-1/05
08/01/2005
Hurd Peninsula, Livingston
Island, South Shetland
Island
(not determined)
wet terrestrial mosses
on the edge of a pool
BYM008
9/01/2009
Byers Peninsula, Livingston (not determined)
Island (South Shetland
Islands, large plain near
large penguin rookery
wet, terrestrial mosses
on edge of large lake
BY052
15/01/2009
Byers Peninsula, Livingston 62°38‘20.1“S
Island (South Shetland
61°06‘44.2“W
Islands, between Midge &
Limnopolar Lake
large lake with
algal mats and fine
sediment/sand on
bottom
LIV-BY16A
16/12/2006
Byers Peninsula, Livingston 62°40‘11.3“S
Island (South Shetland
61°08‘45.3“W
Islands, close to the main
plateau
biofilm in a flooded
area between stones
LIV-BY16C
16/12/2006
Byers Peninsula, Livingston 62°40‘11.3“S
Island (South Shetland
61°08‘45.3“W
Islands, close to the main
plateau
biofilm in a flooded
area between stones
analyses can be found in K o p a l o v á & V a n d e V ij v e r (2013)
and K o p a l o v á et al. (2013).
For light microscope (LM) observations, diatom
samples were prepared following the method described
in V a n d e r W e r f f (1955). In total, 6 sample sites and
microhabitats were examined in this study, with all samples
listed in Table 2. Subsamples of the original material were
oxidized using 37% H20 2 and heating to 80 °C for about
1 h. The reaction was further completed by the addition of
saturated K M n04. Following digestion and centrifugation
(three times 10 min at 3700 xg), the material, free of organic
matter, was further diluted with distilled water for sample
mounting to avoid excessive concentrations o f diatom valves
on the slides. A subsample from the organic-free material
was mounted in Naphrax®. The slides were analyzed using
an Olympus BX51 microscope, equipped with differential
interference contrast (DIC) optics (Nomarski) and Colorview
I Soft Imaging System. Observations on Muelleria
levkoviana sp. nov. were made using brightfield optics as
images in DIC produced only overexposed images. Diatom
samples and slides are stored at BR (National Botanic Garden
of Belgium). For scanning electron microscopy (SEM),
aliquots of the oxidized suspensions were filtered through 1pm pore size polycarbonate filters which were cut in small
pieces, fixed on aluminum stubs after air-drying and sputter
coated (Cressington 208HR, Watford, UK) with Au (10 nm).
Observations and photomicrographs were performed with
a Zeiss® Ultra plus SEM at 8 kV at the Natural History
Museum (London, UK). The number of observations is
quite low but can be explained by the very rare occurrence
of Muelleria taxa in the different samples as was indicated
previously in V a n d e V i j v e r et al. (2010).
Identifications and species determinations are based
primarily on S p a u l d in g & S t o e r m e r (1997), S p a u l d in g et
al. (1999), E s p o s it o et al. (2008) and V a n d e V i j v e r et al.
(2010). Terminology follows mostly V a n d e V i j v e r et al.
(2010) except for valve outline terminology that is based on
H e n d e y (1964) and raphe morphology that follows R o u n d et
al. (1990).
A compiled list of all Muelleria taxa known to date
with references to their original description, type locality and
presence in Antarctica is done (Table 2).
R
esu lts a n d
D
is c u s s io n
total of fourteen Muelleria taxa has been found in the
samples. Eight of them had been previously described
( S p a u l d in g et al. 1999; V a n d e V ij v e r et al. 2010): M.
aequistriata V a n d e V ij v e r et S .A . S p a u l d in g , M. algida
S .A . S p a u l d in g et K o c io l e k , M. kristinae V a n d e V ij v e r ,
M. regigeorgiensis V a n d e V ij v e r et S . A . S p a u l d in g , M.
A
Fottea, Olomoitc, 14(1): 77-90, 2014
79
rostrata V a n d e V i j v e r & S.A. S p a u l d i n g , A/. sabbeiV a n
d e V i j v e r et S.A.S p a u l d i n g , AI. tumida V a n d e V i j v e r
et S.A.S p a u l d i n g and Aí. variolata S.A.S p a u l d i n g et
K o c i o l e k . Six taxa could not be identified using the
currently available literature. All taxa were extremely
rare in the samples making observations in LM and
SEM quite difficult. Often only one (sometimes
even damaged or broken) valve could be observed
in SEM and internal views were very hard (and even
impossible) to find. Nevertheless, a sufficient amount
of observations, both in LM and SEM could be made to
justify their description as new for science: Muelleria
desseiniana sp. nov., Aí. levkoviana sp. nov., M. nogae
sp. nov., M. spauldingiana sp. nov., Aí. sub sabbei sp.
i
Í fff íí ff f
nov. and Aí. undulatoides sp.nov.
Muelleria desseiniana sp. nov. (Figs 1-10)
Light microscopy (Figs 1-8): Frustules in girdle view
rectangular (Fig. 1). Valves linear with gibbous central
part and broadly rounded ends (Figs 2-8). Valve
dimensions (n=10): length 32-40 pm, valve width
8.5-10.0 pm. Axial area rather narrow, linear, slightly
widening near the small, elliptical central area. First
row of areolae along the axial area separated from the
remaining areolae by very narrow, but distinct hyaline
line. Raphe filiform, straight with clearly unilaterally
bent, long proximal raphe endings extending almost till
? f Sfi c
Figs 1-10. M uelleria desseiniana, type population from James Ross Island (sam ple JR I-S IJ): (1 -8 ) LM views; (9) SEM external view; (10)
SEM internal view. Scale bar 10 pm.
80
V
an de
V
ijv e r
et al: New species in the genus Muelleria
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Fottea, Olomoitc, 14(1): 77-90, 2014
83
Figs 11-22. M uelleria levkoviana, type population from Livingston Island (sample BYM 008): (11-21) LM views; (22) SEM external view.
Scale bar 10 pm.
Isotypes (designated here): PLP-244 (University
of Antwerp, Belgium), BRM-ZU9/37 (Hustedt
Collection, Bremerhaven, Gennany)
Type locality: Ulu Peninsula, James Ross Island,
sample JRI-S1J (Leg. J. Komárek; coli. date
22/02/2007).
Etymology: The species is named after our colleague
Dr. Steven Dessein, director of the National Botanic
Garden of Belgium in recognition of his scientific
achievements in botany.
and even beyond the first row of areolae. Distal raphe
endings are weakly deflected to the same side of the
valve as the proximal raphe ends. Striae near the central
area (6-9 striae) more spaced than the other striae and
clearly radiate whereas other striae more distant from
the central area only slightly radiate, becoming almost
entirely parallel near the apices. Central striae 13-15
in 10 pm, distal striae 27-29 in 10 pm. Areolae hardly
discernible in LM.
Scanning electron microscopy (Figs 9-10): Striae
composed of usually transapically elongated, short,
slitlike areolae. First row of areolae bordering the axial
area however appearing as short wavy slits, separated
from the rest of the striae by a narrow hyaline area
and gradually becoming longer towards the apices
(Fig. 9). Near the distal raphe ends, one or two large,
rounded canal punctae present. Proximal raphe endings
unilaterally bent, long, extending into the first row of
areolae. Distal raphe endings deflected onto the valve
mantle, not bifurcated (Fig. 9). Internally, clear but short
rectelevatum present between proximal raphe endings.
Longitudinal canals forming narrow ribs running from
central area towards the apices, clearly perforated on
both sides by first row of areolae (Fig. 10). Distal raphe
endings tenninating on short helictoglossae (Fig. 10).
Ecology and distribution: So far the species was
only observed in very small numbers in samples
obtained from Livingston Island (SSI) and James
Ross Island. The largest population was found in one
sample, collected from a seepage area near Berry Hill
on Ulu Peninsula on James Ross Island. The sample
was dominated by several Luticola and Hantzschia
species clearly indicating its aerophilic nature. Due to
confusion with other similar taxa such as M. luculenta
(as it was earlier reported in Zidarova 2008, PI. III, fig.
26) or the larger and coarser M. algida, its geographic
distribution is still unclear but it is not excluded that
the new species is more widespread on the Maritime
Antarctic islands.
Holotype (designated here): BR-4350 (National
Botanic Garden, Meise, Belgium)
Taxonomical remarks: Muelleria desseiniana lias
a unique combination of features that separates it
84
V
a n de
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ijv e r
et al: New species in the genus Muelleria
Figs 23-30. M uelleria nagae, type population from Livingston Island (sample LIV -B Y 16A ): (23-29) LM views; (30) SEM external view.
Scale bar 10 pm.
well from all other known Muelleria taxa. The raphe
is not bifurcated in its distal part, a feature rarely
seen within the genus Muelleria. Muelleria sabbei
shows comparable, though shorter, raphe endings but
has a clearly elliptic-lanceolate valve outline with
broad, convex margins ( V a n d e V i j v e r et al. 2010).
Additionally, the areolae bordering the central area
are small and rounded whereas in Aí. desseiniana, the
areolae fonn long wavy slits. Muelleria varipunctata
S . A . S p a u l d i n g et K o c i o l e k shows a similar valve
outline but has a much coarser striation pattem,
large round areolae near the axial area and different
(=bifurcating) distal raphe endings. Muelleria
lototskayae V a n d e V i j v e r , M. luculenta and M
algida are larger with longer valves and a different
raphe course, usually showing a tenninal bifurcation
( S p a u l d i n g et al. 1999; V a n d e V i j v e r et al. 2010). All
three taxa have small, rounded to short slitlike areolae,
never showing the elongated, wavy areolae typical for
M. desseiniana.
Muelleria levkoviana sp. nov. (Figs 11-22)
Light microscopy (Figs 11-21): Frustules in girdle
view rectangular (Fig. 11). Valves linear with
clearly undulating margins, a gibbous central part
and cuneately rounded ends (Figs 12-21). Valve
dimensions (n=15): length 24-38 pm, valve width
Fottea, Olomoitc, 14(1): 77-90, 2014
85
Figs 31-40. M uelleria spauldingiana, images from various population from Livingston Island: (31, 32, 36) are from the type population
(sam ple BY052); (31-38) LM views; (39) SEM external view o f an entire (broken) valve; (40) SEM external detail o f the valve apex in girdle
view show ing distal raphe fissures, m antle areolae and canal puncta. Scale bar 10 pm except for figure 1 where scale bar = 1 pm.
6.0-7.8 um. Axial area rather narrow, linear, widening
near the small central area. First row of areolae not
separated from the rest of the striae. Raphe filiform,
straight with clearly unilaterally bent, long proximal
raphe endings extending almost till and even beyond
the first row of areolae. Distal raphe endings not well
visible in LM, apparently deflected. Striae near the
central area (5-7 striae) only slightly more spaced than
the other striae and clearly radiate whereas other striae
more distant from the central area only slightly radiate.
gradually becoming parallel near the apices. Central
striae 16-18 in 10 pm, distal striae 20-22 in 10 pm.
Areolae moderately discernible in LM.
Scanning electron microscopy (Fig. 22): Striae
composed of two types of areolae: near the axial
area usually 1-3 rows of apically elongated areolae,
gradually becoming transapically elongated. Central
striae composed of only transapically elongated
areolae. Proximal raphe endings clearly bent, almost
reaching each other at the end, extending till the first
86
V
an de
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et al: New species in the genus Muelleria
Figs 4 1-51 .M u elleria subsabbei, type population from Livingston Island (sam ple L IV -B Y 16C ): (41-50) LM views; (51) SEM external view.
Scale bar 10 pm.
Figs 52-61. M uelleria undulatoides, type population from Livingston Island (sam ple 5-1/05): (52-60) LM views; (61) SEM external view.
Scale bar 10 pm.
row of areolae. Distal raphe endings deflected, clearly
bifurcating due to siliceous flap. Canal puncta not
observed. No observations of the valve interior could
be made due to the rarity of the taxon.
Holotype (designated here): BR-4351 (National
Botanic Garden, Meise, Belgium)
Isotypes (designated here): PLP-245 (University
of Antwerp, Belgium), BRM-ZU9/38 (Hustedt
Collection, Bremerhaven, Gennany)
Type locality: Byers Peninsula, Livingston Island,
South Shetland Islands, sample BYM008 (Leg. B. Van
de Vijver; coli. date 09/01/2009).
Etymology: The species is named after our colleague
Fottea, Olomouc, 14(1): 77-90, 2014
Dr. Zlatko Levkov (University of Skopje, Macedonia)
in recognition of his important contributions in diatom
research.
Ecology and distribution: Muelleria levkoviana
was only found so far in small numbers in one moss
sample on Byers Peninsula (Livingston Island). The
mosses were sampled near the shore of a very large
lake close to a large penguin rookery. Remains of
penguins, such as feathers and partly decomposed
body parts, were visible in and around the lake.
The sample was dominated by Chamaepinnularia
australomediocris ( L a n g e - B e r t a l o t et S c h m i d t ) V a n
d e V i j v e r and various Psammothidium taxa such as P.
papilio ( K e l l o g g et al.) K o p a l o v á et V a n d e V i j v e r ,
P. germainii ( M a n g u i n ) S a b b e and P. incognitum
( K r a s s k e ) V a n d e V i j v e r et B e y e n s .
Taxonomical remarks: Despite the single SEM
observation of the M. levkoviana so far, it is
possible to justify its separation from all other
Muelleria taxa known so far. Similar taxa include M
terrestris ( P e t e r s e n ) S.A.S p a u l d i n g et S t o e r m e r , M.
meridionalis S.A.S p a u l d i n g et S t o e r m e r , M. supra
S . A . S p a u l d i n g et E s p o s i t o , M. australoatlantica and
M. relicta ( M c C a l l ) R e i c h a r d t . Muelleria relicta
is probably the most similar species based on the
structure of the central striae which are only weakly
more distant in comparison with the rest of the striae.
A similar pattern is found in M. levkoviana. Both taxa
have a comparable raphe structure but differ in the
valve outline with M. relicta having less undulating
valve margins and more broadly rounded apices
( M c C a l l 1933; R e i c h a r d t 2004). Moreover, based on
the illustration in R e i c h a r d t (2004, plate 4, fig. 7) it is
obvious that the first row of areolae is separated from
the rest of the striae contrary to M. levkoviana. It is
however unfortunate that no SEM illustrations exist of
M. relicta to allow comparison of the areola shape and
the structure and the presence of canal puncta, absent
in M levkoviana. Muelleria supra has more undulating
valve margins, a larger central valve part, more distantly
spaced central striae and typically large canal puncta
( E s p o s i t o et al. 2008). Muelleria terrestris is smaller
with parallel valve margins while M. meridionalis has
non-bifurcating distal raphe endings and two large
canal puncta. Moreover, the areolae are large and
rounded and never present the typical direction shift
as is typical fo rM levkoviana ( S p a u l d i n g et al. 1999).
Finally, M. australoatlantica has larger central striae,
longer proximal raphe endings and a high number of
canal puncta and non-bifurcating raphe endings ( V a n
d e V i j v e r et al. 2010).
Muelleria nogae sp. nov. (Figs 23-30)
Light microscopy (Figs 23-29): Larger valves linear
with a clearly widened, gibbous central part and
87
broadly rounded apices (Figs 23-24). Smaller valves
more elliptic-lanceolate with almost convex, less
gibbous margins and broadly rounded ends (Figs 2529). Valve dimensions (n=10): length 24-50 pm, valve
width 10.0-14.0 pm. Axial area rather broad, linear
throughout most of its length, clearly widening to form
a large, elliptic central area. First row of areolae not
markedly separated from the rest of the striae. Raphe
filiform, straight with clearly unilaterally bent to even
hooked, long proximal raphe, completing their course
entirely within the large central area, even within
the smallest specimens. Distal raphe endings clearly
deflected. Striae near the central area almost not more
spaced than the other striae and clearly radiate whereas
the other striae more distant from the central area only
slightly radiate, gradually becoming parallel near the
apices, 18-20 in 10 pm. Areolae clearly discernible in
LM, ca. 15-20 in 10 pm.
Scanning electron microscopy (Fig. 30): Striae
composed of large, rounded to transapically elongated,
oval areolae. The smallest areolae present near the
central area. Proximal raphe endings sometimes
asymmetrically bent with one hooked and one simply
bent. Distal raphe endings deflected, bifurcated due to
the presence of a large siliceous flap. Presence of canal
puncta unclear due to possible confusion with normal
areolae.
Holotype (designated here): BR-4352 (National
Botanic Garden, Meise, Belgium)
Isotypes (designated here): PLP-246 (University
of Antwerp, Belgium), BRM-ZU9/39 (Hustedt
Collection, Bremerhaven, Germany)
Type locality: Byers Peninsula, Livingston Island,
South Shetland Islands, sample LIV-BY16A (Leg. M.
Toro; coli. date 16/12/2006).
Etymology: The species is named after Dr. Teresa
Noga (Institute of Botany, Krakow, Poland) who found
the species also on King George Island.
Ecology and distribution: Muelleria nogae is a rare
species found so far only on Livingston Island and on
King George Island ( T e r e s a N o g a , pers. comm.). The
largest population was found in sample LIV-BY16A,
taken from a biofilm in an area flooded by fresh water
between stones on Byers Peninsula close to the main
plateau. The sample is dominated by various taxa of
Luticola, Hantzschia, Pinnularia and Diadesmis.
Taxonomic remarks: Muelleria nogae can hardly be
confused with any other Muelleria species known so
far, based on the unique, large central area. Muelleria
bachmannii ( H u s t e d t ) S .A .S p a u l d i n g et S t o e r m e r
shows a somewhat similar valve outline with a
somewhat enlarged central area but the latter area
never reaches the same size and shape as in M. nogae
( S p a u l d i n g et al. 1999, Fig. 67). Other Muelleria taxa
with a comparable gibbous valve center such as the
88
more narrow M. meridionalis differ sufficiently not to
be mistaken for M. nogae.
Muelleria spauldingiana sp. nov. (Figs 31-40)
Light microscopy (Figs 31-38): Valves linearlanceolate with convex margins and clearly capitate,
broadly rounded apices. Valve dimensions (n=8):
length 38-50 pm, width 8.5-9.5 pm. Axial area rather
narrow, linear, only very slightly widening towards
the very small central area. First row of areolae
bordering the axial area separated from the striae by
a narrow hyaline line. Raphe filiform and straight.
Proximal raphe endings unilaterally bent, relatively
long, terminating before the first row of areolae, never
extending into the first row. Distal raphe endings
deflected to the same side as the proximal endings.
Striae near the central area (4-5 striae) more spaced
than the other striae and clearly radiate whereas other
striae more distant from the central area only slightly
radiate, gradually becoming parallel and even weakly
convergent near the apices. Central striae 13-15 in 10
pm, distal striae 19-21 in 10 pm. Striae rather coarse,
areolae discernible inLM , 18-20 in 10 pm.
Scanning electron microscopy (Figs 39-40):
Striae composed of irregularly (c- and e-) shaped
areolae throughout the entire valve. Largest and most
complex areolae present near the axial area, gradually
becoming smaller and more rounded towards the valve
margins (Fig. 39). Areolae on mantle showing similar
irregular shape (Fig. 40). Proximal raphe endings bent,
terminating before the first row of areolae. Distal raphe
endings bifurcating due to a large siliceous flap. One
large canal punctum present near distal raphe endings
(Fig. 40). Due to the rarity of this taxon, no observations
of the valve interior could be made.
Holotype (designated here): BR-4353 (National
Botanic Garden, Meise, Belgium)
Isotypes (designated here): PLP-247 (University
of Antwerp, Belgium), BRM-ZU9/40 (Hustedt
Collection, Bremerhaven, Germany)
Type locality: Byers Peninsula, Livingston Island,
South Shetland Islands, sample BY052 (Leg. B. Van
de Vijver; coli. date 15/01/2009).
Etymology: The species is named after our colleague
Dr. Sarah A. Spaulding (INSTAAR, Boulder, Colorado,
USA) in recognition of her contributions to Antarctic
diatom research in general and the genus Muelleria
more specifically.
Ecology and distribution: Muelleria spauldingiana
is a very rare species, occasionally observed in very
low numbers in some lakes of Livingston Island (South
Shetland Islands). The largest population was found in
a large lake between Midge and Limnopolar Lake on
the central plateau of Byers Peninsula (although only
a few valves were found). The lake is characterized
by a slightly alkaline pH (7.35) and low specific
V
an de
V
ijv e r
et al: New species in the germs Muelleria
conductance values (< 100 gS.cm ').
Taxonomic remarks: There are hardly any Muelleria
species that show a similar valve outline as present in
M spauldingiana. The most similar taxon seems to
be M. peraustralis (W. et G.S.W e s t ) S .A .S p a u l d i n g
et S t o e r m e r , described in 1911 from the Antarctic
Continent ( W e s t & W e s t 1911, S p a u l d i n g & S t o e r m e r
1997; V a n d e V i j v e r et al. 2012). Both species can
however be separated based on the shape of the areolae,
being slit-like in M. peraustralis but very irregular,
even e-shaped in M. spauldingiana. Moreover, M.
spauldingiana has more capitate apices compared to
the subcapitate apices in M. peraustralis. Muelleria
bachmannii also shows a comparable valve outline
but has very small, rounded areolae ( V a n d e V i j v e r ,
pers. obs.). Muelleria meridionalis is smaller with less
elongated valves, non-bifurcating distal raphe endings
and small, slit-like areolae. All other Muelleria
species lack the typical capitate apices, coarse striation
pattem and irregularly shaped areolae, justifying the
description of M. spauldingiana as a new species.
Muelleria subsabbei sp. nov. (Figs 41-51)
Light microscopy (Figs 41-50): Valves strictly
lanceolate with non-protracted, broadly rounded,
apices. Valve dimensions (n=20): length 18-27 pm,
width 5.5-6.9 pm. Axial area moderately broad, linear,
widening to form a small, elliptic central area. First
row of areolae bordering the axial area separated from
the striae by a narrow hyaline line. Raphe filiform and
straight. Proximal raphe endings short, unilaterally
bent, terminating before the first row of areolae. Distal
raphe endings deflected to the same side as the proximal
endings. Striae near the central area (4-5 striae) more
spaced than the other striae and clearly radiate whereas
other striae more distant from the central area only
slightly radiate, gradually becoming parallel and even
weakly convergent near the apices. Central striae ca.
18 in 10 pm, distal striae 26-28 in 10 pm. Striae rather
coarse, areolae usually but not always discernible in
LM.
Scanning electron microscopy (Fig. 51): Striae
composed of very fine, transapically elongated slit-like
areolae throughout the entire valve. Areolae bordering
the axial area sometimes irregularly shaped, separated
by hyaline zone from the rest of the striae. Proximal
raphe endings bent, terminating before the first row of
areolae. Distal raphe endings not bifurcating, strongly
hooked onto the valve face extending into the first row
of areolae, never continuing onto the mantle. One large
canal punctum present near distal raphe endings. Due
to the rarity of this taxon, no observations of the valve
interior could be made.
Holotype (designated here): BR-4354 (National
Botanic Garden, Meise, Belgium)
Fottea, Olomouc, 14(1): 77-90, 2014
Isotypes (designated here): PLP-248 (University
of Antwerp, Belgium), BRM-ZU9/41 (Hustedt
Collection, Bremerhaven, Germany)
Type locality: Byers Peninsula, Livingston Island,
South Shetland Islands, sample LIV-BY16C (Leg. M.
Toro; coli. date 16/12/2006).
Etymology: The specific epithet refers to the
resemblance to Muelleria sabbei V a n d e V i j v e r et
S . A . S p a u l d i n g but showing lower valve dimensions.
Ecology and distribution: Muelleria subsabbei is
a rare species found so far only in two samples on
Livingston Island. The largest population was found
in sample LIV-BY16C, taken between stones from
a biofilm in a seepage area flooded by fresh water on
Byers Peninsula close to the main plateau. The sample
is dominated by various taxa of Luticola, Hantzschia,
Pinnularia and Diadesmis.
Taxonomic remarks: Muelleria subsabbei is up to
now the smallest Muelleria species observed. Other
small Muelleria species such as M. kristinae V a n d e
V i j v e r show a different pattern of the central striae
(lacking any differentiation with the rest of the striae,
different distal raphe endings (never continuing onto
the valve face) and different valve outlines (linearlanceolate with acutely rounded ends compared to the
strictly lanceolate valves with broadly rounded ends in
M. subsabbei) ( V a n d e V i j v e r et al. 2010). Muelleria
terrestris ( P e t e r s e n ) S . A . S p a u l d i n g et S t o e r m e r
has more linear valves with longer proximal raphe
endings. Unfortunately, SEM images for the latter
species are lacking making a comparison on the distal
raphe endings and the shape of the areolae impossible.
Muelleria sabbei is much larger with clearly convex,
rounded margins but shows comparable distal raphe
endings terminating on the valve face, hence the
nomenclatural connection between both species.
Muelleria undulatoides sp. nov. (Figs 52-61)
Light microscopy (Figs 52-60): Valves strictly linear
with parallel but clearly undulated margins and almost
non-protracted to weakly subcapitate (never rostrate),
broadly rounded apices. Valve dimensions (n=10):
length23-29 pm, width6.0-7.5 pm. Axial area narrow,
linear, slightly widening forming a small, elliptic
central area. First row of areolae bordering the axial
area usually not separated from the striae by a narrow
hyaline line, although valves with larger areolae near
the axial area were observed. Raphe filiform, straight.
Proximal raphe endings short, unilaterally bent,
terminating before the first row of areolae. Distal raphe
endings deflected to the same side as the proximal
endings. Striae near the central area (6-8 striae) very
slightly more spaced than the other striae and clearly
radiate whereas other striae more distant from the
central area only slightly radiate, parallel in the lower
89
1/3 of the valve and even weakly convergent near the
apices. Central striae 18-20 in 10 pm, distal striae
23-26 in 10 pm. Striae rather coarse, areolae usually
discernible in LM.
Scanning electron microscopy (Fig. 61): Only one
eroded valve observed. Striae composed of small,
rounded areolae. Areolae of the central striae smaller.
Presence of canal puncta unclear. Proximal raphe
endings clearly deflected, terminating before the first
row of areolae. Distal raphe endings not bifurcating,
continuing onto the valve mantle.
Holotype (designated here): BR-4355 (National
Botanic Garden, Meise, Belgium)
Isotypes (designated here): PLP-249 (University
of Antwerp, Belgium), BRM-ZU9/42 (Hustedt
Collection, Bremerhaven, Germany)
Type locality: Hurd Peninsula, Livingston Island,
sample 5-1/05 (Leg. R. Zidarova; coli. date
08/01/2005).
Etymology: The specific epithet refers to the undulated
valve outline of the valves. It was deliberately chosen
not to use the epithet ‘undulata’ to avoid confusion
with Navicula gibbula var. undulata K r a s s k e .
Ecology and distribution: So far the species has only
been found in very low numbers in between mosses
covered with thin cyanobacterial mats, collected on the
shore of a small pool on Hurd Peninsula, Livingston
Island, South Shetland Islands.
Taxonomic remarks: Actually, there is only a handful
of Muelleria species showing an undulated valve
outline. K o l b e (1959) discusses the morphology of
Navicula lagerstedtii C l e v e but shows valves that
represented a different taxon, later described as N.
dubitanda. Therefore, several years later, H u s t e d t &
K o l b e described in 1961 Navicula dubitanda (in 1997
latertransferred to M w e / f e r m by S p a u l d i n g et S t o e r m e r )
but the illustrated valves show a species which is much
bigger (up to 52 pm) with very narrow apices contrary
to M. undulatoides that is smaller and has broadly
rounded apices ( H u s t e d t 1961). Navicula gibbula var.
undulata, described in 1938 from Iceland, has a similar
valve outline with undulated margins ( K r a s s k e 1938).
Only 2 specimens could be identified from this species
( L a n g e - B e r t a l o t et al. 1996, plate 17, figs. 25, 25).
Based on the illustrations, the latter taxon apparently
has a finer striation pattern, compared to the coarse
striae in M. undulatoides. Moreover, the overall valve
shape appears more slender and less compacted as is
the case in M. undulatoides. Therefore, we exclude
conspecificity between both taxa despite the lack of a
detailed SEM analysis between both taxa. Muelleria
holmenii ( F o g e d ) S . A . S p a u l d i n g et S t o e r m e r ,
described from Peary Land, Greenland ( F o g e d 1955),
is also only known from a few specimens ( S p a u l d i n g et
al. 1999). Based on the illustrated valves in S p a u l d i n g
90
V
et al. (1999, figs 4, 5), the latter can be separated
from M undulatoides by the narrower rostrate apices,
compared to the more subcapitate, broadly rounded
apices in the new species. Other taxa showing a similar
valve outline are up to now not known. Taxa such as
M. supra or M. meridionalis are sufficiently different
based on their gibbous central part and the differences
in the raphe structure to avoid all confusion ( S p a u l d in g
et al. 1999; E s p o s it o et al. 2008).
A
cknow ledgem ents
Samples on Byers Peninsula (Livingston Island) were taken in
the fram ew ork o f the IPY -L im nopolar Project POL2006-06635
(M inisterio de Ciencia y Tecnología, Spain). Samples on Hurd
Peninsula (Livingston Island) were taken w ith the logistic support
o f the Bulgarian Antarctic Institute. The authors w ould also like
to thank the m em bers o f expeditions to the Czech J.G. Mendel
Antarctic Station for their support and help in the field. This study
has been supported as a long-term research developm ent project
no. RVO 67985939. Mrs. K. K opalová benefitted from an Erasmus
grant during her stay in Belgium and G A U K grant nr. 394211. Part
o f the research was funded w ithin the BELSPO CCAM BIO project
(Belgium ) and by L’O réal-U nesco For Women in Science Program
(Bulgaria).
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© Czech Phycological Society (2014)
Received November 11, 2013
Accepted December 13, 2013