Phytotaxa 710 (1): 001–021 ISSN 1179-3155 (print edition)

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Article PHYTOTAXA
Copyright © 2025 Magnolia Press ISSN 1179-3163 (online edition)

https://doi.org/10.11646/phytotaxa.710.1.1

Diatom diversity in a mountain lake in southern European Alps (Italy)

FRANCESCA CIUTTI1*, SAÚL BLANCO2, MARCO BERTOLI3, ELISABETTA PIZZUL3, PAOLO PASTORINO4,
MARINO PREARO4 & CRISTINA CAPPELLETTI1
1
Technology Transfer Center, Fondazione Edmund Mach, Via Edmund Mach 1, 38098 San Michele all’Adige (TN), Italy
� [email protected]; https://orcid.org/0009-0003-5216-1375
� [email protected]; https://orcid.org/0009-0008-1585-7222
2
Diatom Lab. IMARENABIO. La Serna 58 - 24007 Leon, Spain
� [email protected]; https://orcid.org/0000-0002-9015-2512
3
Department of Life Sciences, University of Trieste, Via L. Giorgieri 10, 34127 Trieste, Italy
� [email protected]; https://orcid.org/0000-0001-8345-6107
� [email protected]; https://orcid.org/0000-0001-9916-5005
4
Istituto Zooprofilattico Sperimentale del Piemonte, Liguria e Valle d’Aosta, Via Bologna 148, 10154 Torino, Italy
� [email protected]; https://orcid.org/0000-0002-0585-1168
� [email protected]; https://orcid.org/0000-0002-2847-6006
* Author for correspondence

Abstract

Mountain lakes, often regarded as sentinels of environmental degradation and climate change, are extreme environments.
Diatoms are widely considered good indicators in freshwater ecosystems. Littoral diatoms in lakes are highly effective in
detecting trophic and thermal conditions and point-source pollution and are considered early warning of potential changes in
lake ecosystems. An extensive study of littoral diatoms of Upper Lake Balma (southern European Alps, Italy) was conducted
in 2021. Sampling was carried out at seven littoral stations over two periods (July and October). The results revealed a high
diversity of taxa, with a total of 116 identified. Plates with figures for 103 taxa and measures (length, width and striae/fibulae
number) for 104 taxa are given. Iconographic studies significantly enhance the description of diatom biodiversity in these
environments.

Key words: diatom Red List, high-elevation lake, shallow lake, Upper Balma Lake

Introduction

Mountain lakes are extreme environments, typically ice-covered for much of the year. They usually display low
nutrients, high radiation, low temperatures and oligotrophic status (Catalan et al. 2006) and are considered sentinels of
environmental degradation and global environmental changes (Rogora et al. 2018, Moser et al. 2019).
Diatoms serve as effective indicators in freshwater ecosystems and littoral diatoms in lakes are indicative of trophic
and thermal conditions (Rivera-Rondón & Catalan 2020), as well as efficient in detecting point-source pollution,
serving as early indicators of ecosystem changes (Cantonati & Lowe 2014, Rimet et al., 2016).
In Europe there are studies on littoral diatom in a large number of mountain lakes (Levkov et al. 2005, Štefková
2006, Buczkó 2016, Feret et al. 2017, Rivera-Rondón & Catalan 2017, Ossyssek et al. 2023, Solak et al. 2023a).
Renewed interest in diatom research has led to the discovery of new species, such as Psammothidium toroi Blanco,
Pla-Rabes, Wetzel & Granados 2017 (Blanco et al. 2017), Punctastriata subalpina C.E.Wetzel & Ector 2020 and
Punctastriata catenata C.E.Wetzel & Ector 2020 (Wetzel & Ector 2020), Sellaphora lucectoriana Solak, S.Blanco,
P.B.Hamilton, Peszek 2023 (Solak et al. 2023b), Achnanthidium pavense V.Vassal, Heudre, C.E.Wetzel & Tudesque
2023 and Achnanthidium ecrinense V.VassL, Heudre, C.F.Wetzel & Tudesque 2023 (Vassal et al. 2023) and Orthoseira
helvetica Peszek, C.T.Robinson, M.Rybak & Kawecka 2023 (Peszek et al. 2023).
Diatom diversity is particularly high in oligotrophic environments including mountain lakes and it is still under-
estimated (Lange-Bertalot & Metzeltin 1996, Rivera-Rondón & Catalan 2017). Besides the high diversity of littoral

Accepted by María Luján García: 11 Jun. 2025; published: 11 Jul. 2025 

Licensed under Creative Commons Attribution-N.C. 4.0 International https://creativecommons.org/licenses/by-nc/4.0/
diatom community in these environments, few iconographic publications are available: Romania (Buczko 2016),
Austria, Germany and Finland (Lange-Bertalot & Metzeltin 1996) and Pyrénées (Rivera-Rondón & Catalan 2017).
For Italy investigations of benthic diatoms in mountain lakes both for the Alps and the Apennines are quite limited
(Tolotti 2001, Marchetto et al. 2004, Angeli & Cantonati 2005, Cantonati et al., 2021, Marchetto et al. 2021, Padula
et al. 2021, Lepori & Tolotti 2023). Aim of the study is to improve the knowledge of littoral diatom assemblages of
mountain lakes in southern European Alps.

Study area

Upper Balma Lake (Elevation: 2214 m a.s.l.; Surface: 1.82 ha; Maximum depth: 2.77 m; perimeter: 774 m) is a high
elevation, small and shallow lake in the Cottian Alps (Piedmont, north-western Alps, Italy). The lake catchment is
mainly composed of ophiolite metamorphic bedrocks. The lake is included in the Special Area of Conservation (SAC)
and Special Protection Area (SPA) IT1110006, called “Orsiera Rocciavrè” and in the Orsiera Rocciavrè Nature Park.
General overview of the study area, including physicochemical water parameters and sampling sites short
description, is reported in Bertoli et al. (2023). High P concentrations (mean value: 46.7 ug/L in July and 54.8 ug/L in
October) and low transparency (Secchi disk 1.1 ± 0.1 m) are the main feature of the lake, with levels which are typical
of calcareous catchment (Boggero et al. 2005, Fjellheim et al. 2009). Anthropogenic impacts include fish introduction
for recreational fishing in the 1970s (Salvelinus fontinalis) and livestock grazing.

Methods

Benthic diatoms were studied during the ice-free period, in July and October 2021. Diatoms were collected in seven
littoral areas from hard substrates (boulders, cobbles) and processed in the laboratory to obtain permanent slides by cold
and hot oxidation with 37% H2O2 and K2Cr2O7 and addition of HCL 1N. Cleaned frustules were mounted in Naphrax
(refraction index 1.65) for LM. For morphological analysis, 500 valves per sample were counted and identified to
species level using Olympus BX51 microscope at 1000x magnification and imaging system Leica DMC4500 and LAS
Version 4.12.0. For each taxon a subsample of specimens was measured for length and width (µm) and striae/fibulae
number.
Taxonomic identification relied on monographic references for high-altitude lakes (Lange-Bertalot & Metzeltin
1996, Buczko 2016, Rivera-Rondón & Catalan 2017), as well as Lange-Bertalot et al. (2017) and national floras
of Poland and Russia (Bąk et al. 2012, Kulikovskiy et al. 2016). Other books used for identification were those of
Süsswasserflora von Mitteleuropa (Krammer & Lange-Bertalot 1991a, 1991b, 1997a, 1997b), Diatoms of Europe
(Krammer 2000, Lange-Bertalot 2001, Krammer 2002, Krammer 2003, Lange-Bertalot et al. 2011, Levkov et al.,
2016), Bibliotheca Diatomologica (Lange-Bertalot & Krammer 1989, Krammer 1997a, b, Lange-Bertalot 1993,
Kociolet et al. 2014) and Iconographia Diatomologica (Lange-Bertalot & Genkal 1999, Werum & Lange-Bertalot
2004, Siver et al. 2005, Zidarova et al. 2016)
Research papers for Monoraphid (Bukhtiyarova 1996, Monnier et al. 2007, Buczkó et al. 2013, Blanco et al.
2017), Brachyraphid (Pavlov & Levkov 2013), Biraphid (Cantonati et al. 2009, Van de Vijver et al. 2012, Novais et al.
2013, Wojtal et al. 2014, Buczkó et al. 2015, Wetzel et al. 2015, Van de Vijver et al. 2021a, 2021b, 2022a, Alibert et
al. 2023) and Araphid diatoms (Flower 2005, Jüttner et al. 2015, García et al. 2021, Heudre et al. 2021, Van de Vijver
et al. 2022b) were also used.
Taxa were cross-referenced with the Diatom Red List (Hofmann et al. 2018) using OMNIDIA software (Ver.
6.1.8, upgrade May 2024) (Lecointe et al. 1993).

Results

Surveys conducted in July and October 2021 across seven littoral stations identified 116 diatom taxa, spanning 9
orders, 24 families, and 37 genera. Table 1 lists all taxa, with references to plates and figures. The number of specimens
measured for length, width and striae/fibule number in 10 µm is indicated (n = x).

 • Phytotaxa 710 (1) © 2025 Magnolia Press CIUTTI et al.

TABLE 1. Taxon list of species found in Upper Balma Lake, with relative abundance, reference to Diatom Red List and
measures. Abundance in July (Jul) and October (Oct): >50%=dominant (d), 10–50%= abundant (a), 5–10%= frequent (f),
1–5%=occasional (o), <1%= rare (r). Diatom Red List (RL) (Hofmann et al. 2018): a) Highly endangered, b) Threatened of
extinction, c) Risk not estimated, d) Early alert, e) Not endangered, f) Not enough data. Number of measured specimens (n)
for length (L) and width (W) (µm) and striae/fibule number in 10 µm (Str/fib).
Taxon Jul Oct RL Plate & Figure n L W Str/Fib
Achnanthidium minutissimum (Kützing) Czarnecki f o Pl.3 - Figs 2–25
1994 sensu lato
Adlafia sp. r Pl.5 - Figs 30–31 3 14.8–18.0 3.3–3.6 24–27
Aulacoseira alpigena (Grunow) Krammer 1991 r c Pl.1 - Figs 1–4
Aulacoseira granulata (Ehrenberg) Simonsen 1979 r e
Aulacoseira nivalis (W.Smith) J.English & Potapova r e Pl.1 - Figs 5–6
2009
Brachysira calcicola Lange-Bertalot 1994 r a Pl.5 - Figs 19–20
Brachysira aff. neoexilis Lange-Bertalot 1994 r Pl.5 - Figs 16–18 5 22.9–27.5 4.4–4.9 –
Brachysira confusa Van de Vijver, R.L.Albert, r r Pl.5 - Figs 11–15 6 18.8–27.4 5.7–6.5 25–28
B.Kennedy & Kusber 2021
Brachysira sp. r Pl.5 - Figs 21–22
Caloneis sp. r Pl.4 - Fig. 6
Caloneis bacillum (Grunow) Cleve 1894 sensu lato r f Pl.4 - Figs 1–3 5 28.6–38.6 6.7–7.2 20–22
Caloneis lauta J.R.Carter 1981 r c Pl.4 - Figs 4–5 3 28.4–35.9 5.5–6.2 16–17
Cavinula pseudoscutiformis (Hustedt) D.G.Mann & r b Pl.6 - Figs 41–44
Stickle 1990
Chamaepinnularia cf. mediocris (Krasske) Lange- r d Pl.6 - Fig. 28 1 9.0 3.0 19
Bertalot 1996
Chemopinnularia sp. r Pl.6 - Figs 24–27 4 6.6–13.0 2.3–2.6 18
Cymbopleura sp. r Pl.6 - Fig. 57 1 41.9 10.1 14
Encyonema brevicapitatum Krammer 1997 o o Pl.6 - Figs 58–62 8 11.9–19.3 4.2–5.0 15–18
Encyonema kalbei Krammer 1997 r f Pl.6 - Figs 65–67 1 16.3 4.3 15
Encyonema lunatum (W.Smith) Van Heurck 1880 r r f Pl.7 - Fig. 2 3 37.7–40.6 6.0–6.5 11–13
Encyonema lunatum var. boreale Krammer 1997 r Pl.7 - Fig. 1 1 42.0 5.4 9
Encyonema minutum (Hilse) D.G.Mann 1990 o o e Pl.6 - Figs 50–54 6 9.6–13.8 4.1–4.6 15–18
Encyonema neogracile Krammer 1997 r b Pl.7 - Figs 3–4 3 28.1–39.3 5.7–6.5 12–13
Encyonema perlangebertalotii Kulikovskiy & r r Pl.6 - Figs 63–64 3 15.1–16.8 5.0–5.2 17–18
Metzeltin 2012
Encyonema perpusillum (A.Cleve) D.G.Mann 1990 r b Pl.6 - Figs 55–56 2 21.9–22.0 3.9–4.0 9–11
Encyonema rostratum Krammer 1997 r Pl.6 - Figs 68–71 3 15.9–16.1 3.6–3.9 14–15
Encyonema silesiacum (Bleisch) D.G.Mann 1990 r r e Pl.6 - Figs 46–49 3 23.1–25.2 7.1–7.3 13–14
Encyonema sp. 1 r r Pl.7 - Figs 5–6 3 32.3–36.0 5.6–5.8 11–12
Encyonema sp.2 r Pl.7 - Figs 7–8 5 19.5–22.5 4.6–5.4 16–17
Encyonema sp.3 r Pl.7 - Fig. 9 1 35.0 7.6 13
Encyonopsis aff. neerlandica Van de Vijver, Verweij, r r Pl.7 - Figs 21–23 4 19.0–22.3 3.7–3.9 25–26
van der Wal & A.Mertens 2012
Eunotia boreoalpina Lange-Bertalot & Nörpel- r c Pl.2 - Figs 1–5 2 19.8–24.2 4.5 16–17
Schempp 1998
Eunotia boreotenuis Nörpel-Schempp & Lange- r c Pl.2 - Fig. 16 2 12–17.4 2.9–3.8 17
Bertalot 1996
Eunotia botuliformis F.Wild, Nörpel & Lange-Bertalot r Pl.2 - Fig. 15 1 9.3 3.2 19
1993
Eunotia exigua (Brébisson ex Kützing) Rabenhorst r r e Pl.2 - Figs 9–12 8 13.3–19.2 2.4–3.1 20–24
1864
Eunotia groenlandica Nörpel-Schempp & Lange- r c Pl.2 - Fig. 32 1 27.8 3.0 14
Bertalot 1996
Eunotia islandica Østrup 1918 r f Pl.2 - Figs 13–14 2 27.5–27.9 6.8–7.0 13–14,5
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TABLE 1. (Continued)
Taxon Jul Oct RL Plate & Figure n L W Str/Fib
Eunotia juettnerae Lange-Bertalot 2011 r c Pl.2 - Fig. 30–31 3 60–77.6 3.3–3.5 15–16
Eunotia minor (Kützing) Grunow 1881 r Pl.2 - Fig. 17 1 35.8 7.0 13
Eunotia neocompacta var. vixcompacta Lange- r c Pl.2 - Figs 6–8 3 15.0–20.1 2.9–3.2 19–22
Bertalot 2011
Eunotia paludosa Grunow 1862 r d Pl.2 - Fig. 33 1 18.1 2.8 22
Eunotia praerupta Ehrenberg 1843 r r a 2 42.2–83.9 13.5–14 13–16
Eunotia pseudogroenlandica Lange-Bertalot & r r c Pl.2 - Fig. 18–21 4 12.1–25.6 2.9–3.2 15–18
Tagliaventi 2011
Eunotia aff. pseudogroenlandica Lange-Bertalot & r r Pl.2 - Fig. 22–26 8 10.2–23.4 3.3–3.8 13–16
Tagliaventi 2011
Eunotia valida Hustedt 1930 r r c Pl.2 - Fig. 27–29 5 44.9–72.0 4.3–5.1 11–13
Eunotia sp.1 r 1 22.5 4.0 14
Eunotia sp.2 r
Fragilaria cf. campyla (Hilse) Van de Vijver, Kusber r 5 47.2–50.2 2.6–3.6 21–22
& D.M.Williams 2022
Fragilaria cf. nanana Lange-Bertalot 1993 a d Pl.1 - Figs 19–23 12 24.2–48.5 2.0–2.3 21–23
Fragilaria radians (Kützing) D.M.Williams & Round r e Pl.1 - Figs 15–18 5 17.9–22.0 2.6–3.0 20–22
1988
Fragilaria sp. r r
Frustulia crassinervia (Brébisson ex W.Smith) Lange- r r d Pl.5 - Figs 32–33 3 35.5–41.6 10.1– –
Bertalot & Krammer 1996 11.5
Geissleria aff. moseri Metzeltin, Witkowski & Lange- r r Pl.5 - Figs 23–25 4 11.0–19.6 5.7–5.9 15–17
Bertalot 1996
Genkalia boreoalpina Wojtal, C.E.Wetzel, Ector, r r c Pl.5 - Figs 34–38 4 13.4–15.0 3.8–4.1 –
Ognjanova-Rumenova & Buczkó 2014
Genkalia digitulus Lange-Bertalot & Kulikovskiy r r b Pl.5 - Figs 39–42 4 9.4–11.8 3.9–4.1 26–30
2012
Gomphonema hebridense W.Gregory 1854 r d Pl.7 - Figs 12–15 9 30.3–34.9 5.5–6.1 14–17
Gomphonema subclavatum (Grunow) Grunow 1884 r r e Pl.7 - Figs 10–11 7 34.0–50.6 7.4–8.8 10–11
Gomphonema varioreduncum Jüttner, Ector, E. o r f Pl.7 - Figs 16–20 15 15.2–26.8 4.0–5.0 12–14
Reichardt, Van de Vijver & E.J.Cox 2013
Humidophila schmassmannii (Hustedt) Buczkó & o o c Pl.6 - Figs 33–36 9 7.2–9.6 2.6–3.1
Wojtal 2015
Humidophila sp. r r Pl.6 - Figs 29–32 7 6.4–10.3 1.9–2.6
Kobayasiella parasubtilissima (H.Kobayasi & r r d Pl.5 - Figs 26–29 3 19.2–20.6 3.7–3.9
T.Nagumo) Lange-Bertalot 1999
Mayamaea cf. fossalis (Krasske) Lange-Bertalot 1997 r e Pl.6 - Fig. 45 1 3.5 2.7 22
Meridion constrictum Ralfs 1843 r r e Pl.1 - Figs 7–9 4 12.5–18.6 5.5–6.4 20–24
Microcostatus maceria (Schimanski) Lange-Bertalot r r b Pl.6 - Figs 37–39 2 13.8 4.2 – 4.3
1999 –14.5
Microcostatus krasskei (Hustedt) J.R.Johansen & Sray r d Pl.6 - Fig. 40 1 8.6 3.4
1998
Microfissurata paludosa Cantonati & Lange-Bertalot r Pl.6 - Fig. 1 2 16.5 4.6
2009 – 17.9
Navicula exilis Kützing 1884 r b Pl.5 - Figs 6–8 4 28.4–36.0 5.9–6.2 14–15
Navicula sp. r Pl.5 - Fig. 9 2 18.8–20.0 5.8–6.0 18–19
Neidium aff. levanderi (Hustedt) Lange-Bertalot & r Pl.5 - Figs 4–5 4 22.4–30.7 4.6–6.4 32–33
Metzeltin 1996
Neidium alpinum Hustedt 1943 r r a 3 20.1–26.6 4.2–5.1
Neidium longiceps (W.Gregory) R.Ross 1947 r r c Pl.5 - Figs 1–3 4 25.3–32.2 5.6–7.3 28–30
Nitzschia acidoclinata Lange-Bertalot 1976 r r d Pl.7 - Figs 32–33 3 19.0–21.2 3.0–3.1 26–28 (8–10)
Nitzschia alpina Hustedt 1943 r b Pl.7 - Fig. 34 1 20.5 3.2 11 (30–31)
Nitzschia bryophila (Hustedt) Hustedt 1943 r r c Pl.7 - Figs 24–27 6 15.5–29 4.1–40.6
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TABLE 1. (Continued)
Taxon Jul Oct RL Plate & Figure n L W Str/Fib
Nitzschia gracilis Hantzsch 1860 r e 3 – 2.7–3.0 – (13–16)
Nitzschia palea var. debilis (Kützing) Grunow 1880 r e Pl.7 - Figs 35–37 4 19.8–29.1 2.7–3.0 – (12–13)
Nitzschia perminuta Grunow 1881 o o e Pl.7 - Figs 28–31 11 18.6–29.0 2.6–3.1 29–32 (10–13)
Odontidium mesodon (Ehrenberg) Kützing 1849 r r e Pl.1 - Figs 28–32 15 11.3–19.2 6.2–7.2 24–26
Pinnularia ammerensis Kulikovskiy, Lange-Bertalot r f Pl.4 - Figs 14–15 7 22.6–36.3 5.4–6.5 11–14
& Metzeltin 2010
Pinnularia borealis Ehrenberg 1843 r e 2 37.0–45.0 8.7–10.8 4–5
Pinnularia cf. lindanedbalovae B.van de Vijver & r r Pl.4 - Figs 12–13 5 22.0–28.5 4.7–5.0 12–14
A.Moravcová 2013
Pinnularia microstauron (Ehrenberg) Cleve 1891 r d Pl.4 - Figs 18–19 7 50.7–63.0 9.1–11.2 9–10
Pinnularia aff. microstauron var. rostrata Krammer r r Pl.4 - Figs 20–23 6 25.2–27.5 5.0–5.7 12
2000
Pinnularia cf. notabilis Krammer 1985 r r c Pl.4 - Figs 16–17 13 55.7–95.0 11.7– 10–11
18.0
Pinnularia silvatica J.B.Petersen 1935 r f 3 17.2–20.6 3.7–3.8 20–22
Pinnularia sinistra Krammer 1992 r r e Pl.4 - Figs 7–11 14 24.0–41.2 4.0–5.1 11–12
Planothidium distinctum (Messikommer) Lange- r r b Pl.3 - Fig. 1
Bertalot 1999
Psammothidium aff. acidoclinatum (Lange-Bertalot) r r Pl.3 - Figs 51–55 10 10.3–14.3 4.9–5.9 25–26
Lange-Bertalot 1999
Psammothidium bioretii (H.Germain) Bukhtiyarova & r e Pl.3 - Fig. 35
Round 1996
Psammothidium bristolicum Bukhtiyarova 1996 o r f Pl.3 - Figs 41–45 12 10.2–14.2 4.4–5.0 25–27
Psammothidium chlidanos (M.H.Hohn & Hellerman) r r b Pl.3 - Figs 31–34 4 10.0–16.4 4.9–5.7
Lange-Bertalot 1999
Psammothidium helveticum (Hustedt) Bukhtiyarova & r r e Pl.3 - Figs 26–30 6 12.4–18.6 5.0–6.7 22–24
Round 1996
Psammothidium helveticum var. minus (Flower & r e Pl.3 - Figs 76–79 5 8.2–10.6 4.8–6.0 22–27
Jones) Buczkó 2016
Psammothidium levanderi (Hustedt) Bukhtiyarova & r c Pl.3 - Figs 80–82 3 8.5–8.7 4.2–5.0 23–26
Round 1996
Psammothidium marginulatum (Grunow) r f Pl.3 - Figs 36–40 6 11.3–12.7 4.5–4.8 25–27
Bukhtiyarova & Round 1996
Psammothidium microscopicum (Cholnoky) S.Blanco o r f Pl.3 - Figs 71–75 5 4.0–4.8 3.1–3.5 28
2016
Psammothidium rechtense (Leclercq) Lange-Bertalot r r a Pl.3 - Figs 46–50 4 11.2–12.5 4.4–4.8 28
1999
Psammothidium scoticum (R.J.Flower & V.J.Jones) o o a Pl.3 - Figs 61–65 11 8.5–10.5 3.6–4.4 26–31
Bukhtiyarova & Round 1996
Psammothidium subatomoides (Hustedt) Bukhtiyarova a o d Pl.3 - Figs 56–60 12 6.1–9.1 3.6–4.7
& Round 1996
Psammothidium toroi Blanco, Pla-Rabes, Wetzel & r r Pl.3 - Figs 66–70 11 6.6–10.7 3.4–4.1 22–26
Granados 2017
Psammothidium sp. r Pl.3 - Figs 83–85 2 8.0–8.5 4.4–4.5 25
Pseudostaurosira parvissima Kociolek 2014 r r Pl.1 - Figs 33–37 8 6.7–12.0 1.9–2.5 18–19
Pseudostaurosira sp. f f 30 8.6–16.4 3.0–5.2 12–14
Sellaphora aff. medioconvexa (Hustedt) C.E.Wetzel r r Pl.6 - Figs 7–10 1 11.8 3.7
2015
Sellaphora aff. multiconfusa (Lange-Bertalot) r r Pl.6 - Figs 11–14 5 9.6–13.9 4.1–4.8 24–26
C.E.Wetzel, Ector, B.Van de Vijver, Compère &
D.G.Mann 2015
Sellaphora pseudopupula (Krasske) Lange-Bertalot r r c Pl.6 - Figs 2–6 4 17.8–20.3 5.4–6.0 22
1996
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Diatom diversity in southern European Alps Phytotaxa 710 (1) © 2025 Magnolia Press • 
TABLE 1. (Continued)
Taxon Jul Oct RL Plate & Figure n L W Str/Fib
Sellaphora sorella (M.H.Hohn & Hellerman) r r Pl.6 - Figs 15–19 11 5.8–10.0 3.0–3.5 17–20
C.E.Wetzel, Ector, B.Van de Vijver Compère &
D.G.Mann 2015
Sellaphora vanlandinghamii (Kociolek) C.E.Wetzel r Pl.6 - Figs 20–23 8 5.7–7.0 3.4–4.0 28
2015
Sellaphora sp. r 1 12.7 3.7 24
Stauroforma exiguiformis (Lange-Bertalot) r r b Pl.1 - Figs 24–27 3 26.8–28.2 3.3–3.6 17–18
R.J.Flower, V.J.Jones & Round 1996
Stauroneis aff. acidoclinata Lange-Bertalot & Werum r e Pl.5 - Fig. 10 2 35.3–38.0 7.5 23–24
2004
Staurosira aff. aventralis Lange-Bertalot & Rumrich o o Pl.1 - Figs 38–42 21 8.0–12.7 3.1–3.9 14–16
2000
Staurosira cf. venter (Ehrenberg) Cleve & J.D.Möller r o e Pl.1 - Figs 43–47 20 5.8–9.6 3.4–4.2 14–15
1879
Staurosirella ansata (M.H.Hohn & J.Hellerman) r r Pl.1 - Figs 48–52 7 8.3–15.8 2.8–3.7 12
J.C.Kingston 2000
Staurosirella sp. r Pl.1 - Figs 53–56 2 4.3–5.1 2.62.7 12–13
Surirella roba Leclercq 1983 r c
Tabellaria flocculosa (Roth) Kützing 1884 o r e Pl.1 - Figs 10–14 9 14.8–25.2 5.6–7.2 14–18

Genera with a higher number of species were Eunotia (16 species), Psammothidium (14), Encyonema (13) and
Pinnularia (8) (Figure 1). The most abundant genera (>1% relative abundance) included Achnanthidium, Encyonema,
Fragilaria, Humidophila, Nitzschia, Psammothidium, Pseudostaurosira and Staurosira in July and October and
Brachysira, Eunotia, Gomphonema and Tabellaria in July. For 27 taxa identification was possible to genus level, both
for low number of valves found (i.e. Adlafia sp., Cymbopleura sp.) and an unclear correspondence with known species
(i.e. Psammothidium aff. acidoclinatum (Lange-Bertalot) Lange-Bertalot). For the latter, further SEM investigations
are needed. The needle-shaped Fragilaria cf. nanana Lange-Bertalot 1993 was the dominant species, with a mean
percentage of 37.0 and 68.7 respectively in July and October. Other abundant species (>1%) were Achnanthidium
minutissimum s.l., Encyonema brevicapitatum Krammer 1997, Encyonema minutum (Hilse) D.G.Mann 1990,
Humidophila schmassmannii (Hustedt) Buczkó & Wojtal 2015, Nitzschia perminuta Grunow 1881, Psammothidium
scoticum (R.J.Flower & V.J.Jones) Bukhtiyarova & Round 1996, Psammothidium subatomoides (Hustedt) Bukhtiyarova
& Round 1996, Pseudostaurosira sp., Staurosira aff. aventralis Lange-Bertalot & Rumrich 2000 and Staurosira cf.
venter (Ehrenberg) Cleve & J.D.Möller 1879 in July and October, and Gomphonema varioreduncum Jüttner, Ector,
E. Reichardt, Van de Vijver & E.J.Cox 2013, Psammothidium bristolicum Bukhtiyarova 1996, Psammothidium
microscopicum (Cholnoky) S.Blanco 2016 and Tabellaria flocculosa (Roth) Kützing 1844 in July. Most of taxa were
found with low abundance (<1%).
The diatom Red List for protection (Hofmann et al. 2018) includes 71 out of 116 species found (Figure 2). Among
them, 24 species are classified as worth of protection as highly endangered (5) [Brachysira calcicola Lange-Bertalot
1994, Eunotia praerupta Ehrenberg 1843, Neidium alpinum Hustedt 1943, Psammothidium rechtense (Leclercq)
Lange-Bertalot 1999 and Psammothidium scoticum (R.J.Flower & V.J.Jones) Bukhtiyarova & Round 1996], threatened
of extinction (10) [Cavinula pseudoscutiformis (Hustedt) D.G.Mann & Stickle 1990, Encyonema neogracile Krammer
1997, Encyonema perpusillum (A.Cleve) D.G.Mann 1990, Genkalia digitulus Lange-Bertalot & Kulikovskiy 2012,
Microcostatus maceria (Schimanski) Lange-Bertalot 1999, Navicula exilis Kützing 1884, Nitzschia alpina Hustedt
1943, Planothidium distinctum (Messikommer) Lange-Bertalot 1999, Psammothidium chlidanos (M.H.Hohn &
Hellerman) Lange-Bertalot 1999 and Stauroforma exiguiformis (Lange-Bertalot) R.J.Flower, V.J.Jones & Round 1996]
and early alert (9) [Chamaepinnularia cf. mediocris (Krasske) Lange-Bertalot 1996, Eunotia paludosa Grunow 1862,
Frustulia crassinervia (Brébisson ex W.Smith) Lange-Bertalot & Krammer 1996, Gomphonema hebridense Gregory
1854, Kobayasiella parasubtilissima (H.Kobayasi & T.Nagumo) Lange-Bertalot 1999, Microcostatus krasskei (Hustedt)
J.R.Johansen & Sray 1998, Nitzschia acidoclinata Lange-Bertalot 1976, Pinnularia microstauron (Ehrenberg) Cleve
1891, P. subatomoides (Hustedt) Bukhtiyarova & Round 1996].

 • Phytotaxa 710 (1) © 2025 Magnolia Press CIUTTI et al.

FIGURE 1. Number of species observed per genera in Upper Balma Lake.

Diatom diversity in southern European Alps Phytotaxa 710 (1) © 2025 Magnolia Press • 
FIGURE 2. Species included in the Diatom Red List (Hofmann et al. 2018).

Discussion

Upper Balma Lake is a circumneutral (pH 7.6 ± 0.07), low-mineralized lake (conductivity: 12.81 ± 1.34 µS cm⁻¹)
situated on siliceous substrate in the Alps. Elevated total phosphorus levels (46.7 ± 1.31 µg L⁻¹ in July) likely result
from nutrient enrichment due to fish stocking and livestock grazing (Bertoli et al. 2023).
The periphytic diatom community comprises 116 taxa, 27 identified to genus level, with dominant genera including
Achnanthidium, Encyonema, Fragilaria, Humidophila, Nitzschia, Psammothidium, Pseudostautosira and Staurosira.
Most of known taxa are typical of oligo to dystrophic, electrolyte poor and weakly acid or circumneutral habitats,
on siliceous substrate, i.e. G. hebridense W.Gregory 1854, N. perminuta Grunow 1881 P. subatomoides (Hustedt)
Bukhtiyarova & Round 1996 and Psammothidium helveticum (Hustedt) Bukhtiyarova & Round 1996 (Lange-Bertalot
et al. 2017, Van Dam et al. 1994).
Among the most represented genera, Eunotia is typical of naturally acid, oligotrophic to dystrophic freshwater
habitats (i.e. Eunotia botuliformis F.Wild, Nörpel & Lange-Bertalot 1993; Eunotia minor (Kützing) Grunow 1881)
and of acid peat bogs and fens which often neighbor mountain lakes (i.e. E. paludosa Grunow 1862) (Cantonati et al.,
2011, Lange-Bertalot et al. 2017, Rimet et al. 2023).
The genus Psammothidium is frequently observed in lentic and lotic freshwater habitats and includes typical
cold stenothermic species (Blanco et al. 2017, Cantonati et al. 2021). Among the 14 species found in the lake, P.
microscopicum (Cholnoky) S.Blanco 2016 is a cryophilic species of oligotrophic lakes (Cantonati et al., 2021) and
Psammothidium levanderi (Hustedt) Bukhtiyarova & Round 1996 and Psammothidium toroi Blanco, Pla-Rabes,
Wetzel & Granados 2017 are typical of oligotrophic and very low mineralized lakes (Kopalová et al. 2016, Blanco et
al., 2017, Feret et al., 2017).
Araphid diatoms Staurosira and Pseudostaurosira are well represented in the lake, with a mean percentage of 9.1
and 11.1 respectively in July and October. Unattached diatoms are typically present in lakes with long water renewal
time, characterized by dominance of high-profile diatoms which compete for light and nutrients (Rimet et al. 2019).
The needle-shaped F. cf nanana Lange-Bertalot 1993 is the dominant species found in the lake, and it has already been
observed related to fish introduction (Sochuliaková et al. 2018, Cantonati et al. 2021).

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PLATE 1. Light micrographs of centric and araphid taxa. Figs 1–4. Aulacoseira alpigena. Figs 5–6. A. nivalis. Figs 7–9. Meridion
constrictum. Figs 10–14. Tabellaria flocculosa. Figs 15–18. Fragilaria radians. Figs 19–23. F. cf. nanana. Figs 24–27. Stauroforma
exiguiformis. Figs 28–32. Odontidium mesodon. Figs 33–37. Pseudostaurosira parvissima. Figs 38–42. Staurosira aff. aventralis. Figs
43–47. S. cf. venter. Figs 48–52. Staurosirella ansata. Figs 53–56. Staurosirella sp. Scale bar: 10 μm.

Diatom diversity in southern European Alps Phytotaxa 710 (1) © 2025 Magnolia Press • 
PLATE 2. Light micrographs of eunotioid taxa. Figs 1–5. Eunotia borealpina. Figs 6–8. E. neocompacta var. vixcompacta. Figs 9–12. E.
exigua. Figs 13–14. E. islandica. Fig. 15. E. botuliformis. Figs 16. E. boreotenuis. Fig 17. E. minor. Figs 18–21. E. pseudogroenlandica.
Figs 22–26. E. aff. pseudogroenlandica. Figs 27–29. E. valida. Figs 30–31. E. juettnerae. Fig. 32. E. groenlandica. Fig. 33. E. paludosa.
Scale bar: 10 μm.

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PLATE 3. Light micrographs of monoraphid taxa. Fig. 1. Planothidium distinctum. Figs 2–6. Achnanthidium minutissimum s.l. forme
A. Figs 7–11. A. minutissimum s.l. forme B. Figs 12–15. A. minutissimum s.l. forme C. Figs 16–20. A. minutissimum s.l. forme D. Figs
21–25. A. minutissimum s.l. forme E. Figs 26–30. Psammothidium helveticum. Figs 31–34. P. chlidanos. Fig. 35. P. bioretii. Figs 36–40. P.
marginulatum. Figs 41–45. P. bristolicum. Figs 46–50. P. rechtense. Figs 51–55. P. aff. acidoclinatum. Figs 56–60. P. subatomoides. Figs
61–65. P. scoticum. Figs 66–70. P. toroi. Figs 71–75. P. microscopicum. Figs 76–79. P. helveticum var. minus. Figs 80–82. P. levanderi.
Figs 83–85. Psammothidium sp. Scale bar: 10 μm.

Diatom diversity in southern European Alps Phytotaxa 710 (1) © 2025 Magnolia Press • 11
PLATE 4. Light micrographs of symmetric biraphid taxa. Figs 1–3. Caloneis bacillum s.l. Figs 4–5. C. lauta. Fig. 6. Caloneis sp. Figs
7–11. Pinnularia sinistra. Figs 12–13. P. cf. lindanedbalovae. Figs 14–15. P. ammerenensis. Figs 16–17. P. cf. notabilis. Figs 18–19. P.
microstauron. Figs. 20–23 P. aff. microstauron var. rostrata. Scale bar: 10 μm.

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PLATE 5. Light micrographs of symmetric biraphid taxa. Figs 1–3. Neidium longiceps. Figs 4–5. Neidium aff. levanderi. Figs 6–8.
Navicula exilis. Fig. 9. Navicula sp. Fig. 10. Stauroneis aff. acidoclinata. Figs 11–15. Brachysira confusa. Figs 16–18. B. aff. neoexilis.
Figs 19–20. B. calcicola. Figs 21–22. Brachysira sp. Figs 23–25. Geissleria aff. moseri. Figs 26–29. Kobayasiella parasubtilissima. Figs
30–31. Adlafia sp. Figs 32–33 Frustulia crassinervia. Figs 34–38. Genkalia boreoalpina. Figs 39–42. G. digitulus. Scale bar: 10 μm.

Diatom diversity in southern European Alps Phytotaxa 710 (1) © 2025 Magnolia Press • 13
PLATE 6. Light micrographs of symmetric and asymmetric biraphid taxa. Fig. 1. Microfissurata paludosa. Figs 2–6. Sellaphora
pseudopupula. Figs 7–10. S. aff. medioconvexa. Figs 11–14. S. aff. multiconfusa. Figs 15–19. S. sorella. Figs 20–23. S. vanlandinghamii.
Figs 24–27. Chamaepinnularia sp. Fig. 28. Chamaepinnularia cf. mediocris. Figs 29–32. Humidophila sp. Figs 33–36. Humidophila
schmassmannii. Figs 37–39. Microcostatus maceria. Fig. 40. M. krasskei. Figs 41–44. Cavinula pseudoscutiformis. Fig. 45. Mayamaea cf.
fossalis. Figs 46–49. Encyonema silesiacum. Figs 50–54. E. minutum. Figs 55–56. E. perpusillum. Fig. 57. Cymbopleura sp. Figs 58–62.
E. brevicapitatum. Figs 63–64. E. perlangebertalotii. Figs 65–67. E. kalbei. Figs 68–71. E. rostratum. Scale bar: 10 μm.

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PLATE 7. Light micrographs of asymmetric biraphid and nitzschioid taxa. Fig. 1. Encyonema lunatum var. boreale. Fig. 2. E. lunatum.
Figs 3–4. E. neogracile. Figs 5–6. Encyonema sp. 1. Figs 7–8. Encyonema sp. 2. Fig. 9. Encyonema sp. 3. Figs 10–11. Gomphonema
subclavatum. Figs 12–15. G. hebridense. Figs 16–20. G. varioreduncum. Figs 21–23. Encyonopsis aff. neerlandica. Figs 24–27. Nitzschia
bryophila. Figs 28–31. N. perminuta. Figs 32–33. N. acidoclinata. Fig. 34. N. alpina. Figs 35–37. N. palea var. debilis. Scale bar: 10
μm.

Diatom diversity in southern European Alps Phytotaxa 710 (1) © 2025 Magnolia Press • 15
 Among the 70 species listed in the Diatom Red List (Hofmann et al. 2018), 24 taxa (34.3%) are recognized as
conservation priorities.
High elevation freshwater ecosystems comprise the least impacted, pristine and extreme environments (Zaharescu
et al. 2016). In the Arctic and Antarctic regions, limnological and paleolimnological studies have shown that diatoms
are informative of environmental changes in the short and long term. These ecosystems are alarm bells, since they
show signs related to climate change first and to the highest degree (Douglas & Smol 2010). Global warming is now
affecting temperate zones as well, but as expected, lakes in these areas show a temporal shift in changes (in some
cases up to a century) (Lotter et al. 2010, Spauldind & Mcknight 2010). The risk of extirpation of a large number
of cold-stenothermal diatom species, in particular from southern refugia has been hypnotized by Bahls (2017) as a
consequence of climate change.
Extensive studies on the littoral diatoms of high elevation lakes, conducted through multiple sampling stations and
campaigns, can reveal the hidden biodiversity of these environments (Cantonati et al. 2021). Single-sample surveys in
other regions revealed the presence of 355 species in 83 lakes of the Pyrenees (Rivera-Rondón & Catalan 2017), 127
species in 34 lakes from the territory of the Tatra Mountain (Štefková 2006) and 326 species in 62 natural lakes above
1300 m a.s.l. in the French Alps (Feret et al. 2017).
Even if cosmopolitan species play a fundamental role in diatom assemblages, geological and geographical
peculiarities have been observed, and diatoms may frequently develop local forms (Cantonati et al. 2001) and presence
of endemic species and sub-species has been observed (Bahls 2017, Rimet et al. 2023), together with the high number
of threatened taxa (Cantonati et al. 2022).
In the Upper Balma Lake we found a high number of taxa (116), most of which with low abundances (<1%).
Light microscopy (LM) observations allow identification to genus level for 27 taxa (24%). The observation in high
elevation freshwater ecosystems of species or varieties not yet described has been underlined by other authors (Buczko
2016; Rivera-Rondón & Catalan 2017; Safiallah et al. 2020) and recent studies on high elevation, lentic and lotic
environments led to description of new species, e.g. Sellaphora lucectoriana Solak, S.Blanco, P.B.Hamilton, Peszek
2023 (Solak et al. 2023b) and Orthoseira helvetica Peszek, C.T.Robinson, M.Rybak & Kawecka 2023 (Peszek et al.
2023) or new records for local floras (Kheiri et al. 2018, Singh Rana et al. 2022).
Traditional morphological analysis of diatoms will be increasingly supported by genetic investigations (DNA
metabarcoding), which thus will provide valuable contribution in detecting hidden biodiversity (e.g. Kang et al. 2021;
Rimet et al. 2023). Iconographic publications can play a valuable role in enhancing the description of diatom diversity
within these extreme environments (Lange-Bertalot & Metzeltin 1996, Buczko 2016, Rivera-Rondón & Catalan 2017,
Kheiri et al. 2018).

Conclusions

High elevation lentic and lotic freshwater ecosystems are characterized by harsh environmental conditions, supporting
highly specific aquatic communities, rich in endemic and threatened and yet undescribed taxa. Both direct (e.g.,
eutrophication, fish introduction) and indirect (e.g., climate change, acidification, microplastics) anthropogenic
impacts can affect these pristine ecosystems. Research on biodiversity is critical for understanding ecosystem alteration
mechanisms.
Together with genetic studies, iconographic publications can play a valuable role in enhancing the description of
diatom diversity within these unique environments.

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