Biota Neotropica 19(2): e20180568, 2019
www.scielo.br/bn

Periphytic diatoms from an oligotrophic lentic system, Piraquara I reservoir, Paraná 
state, Brazil

Angela Maria da Silva-Lehmkuhl1,2* , Priscila Izabel Tremarin3, Ilka Schincariol Vercellino4 & 

Thelma A. Veiga Ludwig5

1Universidade Federal do Amazonas, Estrada Parintins Macurany, 1805, 69152-240, Parintins, AM, Brasil
2Universidade Estadual Paulista Julio de Mesquita Filho, 13506-900, Rio Claro, SP, Brasil

3Acqua Diagnósticos Ambientais Ltda., Curitiba, PR, Brasil
4Centro Universitario São Camilo, São Paulo, SP, Brasil

5Universidade Federal do Paraná, Av. Cel. Francisco H. dos Santos, Jardim das Américas, 81531980, 
Curitiba, PR, Brasil

*Corresponding author: Angela Maria da Silva-Lehmkuhl, e-mail: angela_ecologia@yahoo.com.br

SILVA-LEHMKUHL, A.M., TREMARIN, P.I., VERCELLINO, I.S., LUDWIG, T.A.V. Periphytic diatoms from 
an oligotrophic lentic system, Piraquara I reservoir, Paraná state, Brazil. Biota Neotropica. 19(2): e20180568. 
http://dx.doi.org/10.1590/1676-0611-BN-2018-0568

Abstract: Knowledge of biodiversity in oligotrophic aquatic ecosystems is fundamental to plan conservation 
strategies for protected areas. This study assessed the diatom diversity from an urban reservoir with oligotrophic 
conditions. The Piraquara I reservoir is located in an Environmental Protection Area and is responsible for the 
public supply of Curitiba city and the metropolitan region. Samples were collected seasonally between October 
2007 and August 2008. Periphytic samples were obtained by removing the biofilm attached to Polygonum 
hydropiperoides stems and to glass slides. The taxonomic study resulted in the identification of 87 diatom taxa. The 
most representative genera regarding the species richness were Pinnularia (15 species) and Eunotia (14 species). 
Five species were registered for the first time in Brazil and seven in the State of Paraná. Taxonomic and ecological 
comments of the species registered are provided.
Keywords: artificial substrate, diatom ecology, macrophytes, reservoirs.

Diatomáceas perifíticas de um sistema lêntico oligotrófico, reservatório do 
Piraquara I, estado do Paraná, Brasil

Resumo: O conhecimento da biodiversidade em ecossistemas aquáticos oligotróficos é fundamental para planejar 
estratégias de conservação de áreas protegidas. Este estudo teve como objetivo conhecer a diversidade de 
diatomáceas em um reservatório urbano com condições oligotróficas. O reservatório do Piraquara I está inserido em 
uma Área de Proteção Ambiental em região de manancial e é responsável pelo abastecimento público de Curitiba 
e região metropolitana. Foram realizadas coletas estacionais entre os meses de outubro de 2007 e agosto de 2008. 
As amostras perifíticas foram obtidas pela remoção do biofilme aderido a caules de Polygonum hydropiperoides e 
de lâminas de vidro. O estudo taxonômico resultou na identificação de 87 táxons. Os gêneros mais representativos, 
considerando a riqueza de espécies, foram Pinnularia (15 espécies) e Eunotia (14 espécies). Cinco espécies de 
diatomáceas foram registradas pela primeira vez no Brasil e sete no estado do Paraná. Comentários taxonômicos 
e ecológicos também são apresentados para as espécies encontradas.
Palavras-chave: substrato artificial, ecologia de diatomáceas, macrófitas, reservatório.

ISSN 1676-0611 (online edition)

Inventory

http://dx.doi.org/10.1590/1676-0611-BN-2018-0568 http://www.scielo.br/bn

http://www.scielo.br/bn
https://orcid.org/0000-0002-0356-9572
https://orcid.org/0000-0003-0576-3499


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Silva-Lehmkuhl, A.M. et al.

Biota Neotrop., 19(2): e20180568, 2019

http://www.scielo.br/bn http://dx.doi.org/10.1590/1676-0611-BN-2018-0568

Introduction
Diatoms are considered one of the most representative groups of 

microalgae due to the number of described species. Approximately 
12,000 species of diatoms have been described hitherto (Guiry 2012) 
although the estimated number of species is thought to be at least 30,000 
(Mann e Vanormeligen 2013). Diatoms are excellent bioindicators of 
water quality, being sensible to environmental pollution and a number of 
anthropogenic pressures, such as the increase in nutrient concentrations, 
acidification, land use, and the presence of toxic chemicals dissolved 
in water (Leskinen & Hällfors 1990, Lowe & Pan 1996, Lobo et al. 
2002, Rimet et al. 2015).

Diatoms release mucilage through raphe, rimoportulae and apical 
pore fields, facilitating the frustule adhesion to different types of 
substrates. This strategy contributes to diatoms representativeness 
in the periphytic community (Round et al. 1990, Lowe 1996). The 
species composition of periphytic assemblages can be influenced by 
the substrate micro-topography (Murdock and Dodds 2007, Souza 
& Ferragut 2012). Inventories using substrates with different surface 
types usually hold a great algal diversity (Ács et al. 2000). Therefore, 
diatom inventories should be carried out in a great number of different 
environments and using diverse substrates to maximize the recovery 
of new species, extend the distribution of known species and better 
define their ecological preferences, which increase the reliability of 
environmental diagnostics.

Man-made lentic environments with oligotrophic conditions are rare 
in urban areas and are examples of environmental health. Unfortunately, 
human activities in the surrounding watershed have been accelerating 
the processes of eutrophication, affecting the water quality and the 
biodiversity, leading to the loss of important ecological functions 
(Tundisi 2003, Torrisi et al. 2010). Due to the undesirable impact of 
these activities on the water physical and chemical conditions, efforts 
have been generally focused on studying eutrophic environments rather 
than on preserving oligotrophic water bodies. For this reason, there 
is usually a gap in the knowledge of the algal diversity of protected 
areas, which creates exceptional opportunities to study the remaining 
oligotrophic environments (Kociolek & Stoermer 2009).

Recent studies in oligotrophic Brazilian reservoirs contributed to 
the description of new diatom species in the genera Kurtkrammeria 
and Encyonema (Marquardt et al. 2016, Marquardt et al. 2017) and 
extended the geographic distribution of the already known species 
(Canani et al. 2011). Paleolimnological studies with diatoms are 
important in providing a further understanding of biodiversity, detecting 
the response of the assemblages over time. Generally, a decrease in the 
diatom diversity during the eutrophication process can be observed 
when comparing environments that were oligotrophic but are currently 
eutrophised (Faustino et al. 2016, Wengrat et al. 2017).

Previous diatom inventories were carried out in urban reservoirs of 
the Iguaçu river basin along a trophic gradient ranging from mesotrophic 
(Passaúna, Bertolli et al. 2010 and Piraquara II, Marra et al. 2016), 
eutrophic (Iraí, Silva et al. 2010) to hypereutrophic (Itaqui, Faria et al. 
2010). The Piraquara I reservoir, which was surveyed in the present 
study, is connected to Piraquara II reservoir. The macrophyte Polygonum 
hydropiperoides Michaux and glass slides were used as substrates to 
assess the periphytic diatoms in these reservoirs.

The present study aimed at assessing the composition of periphytic 
diatom assemblages from the oligotrophic reservoir Piraquara I. We 

provided measures, illustrations, taxonomical comments and ecological 
data based on scientific literature, in order to contribute to the knowledge 
of Brazilian diatoms from oligotrophic environments.

Material and Methods

The Piraquara I Reservoir (25°30’24.16’’ S, 49°1’29.4’’ W) is a 
man-made public water supply of the Iguaçu River basin located in an 
Environmental Protection Area which is in a transition zone between 
the dense and mixed ombrophilous forest. The Cayuguava river was 
dammed to construct the reservoir in 1979 (Figure 1) (Guimarães 2008). 
The reservoir has a 3.3 km2 of flooded area, 7 meters in depth, a flow rate 
of 600 l/s, and a water residence time of 438 days (Júnior et al. 2005).

The physical and chemical data were provided by the Paraná 
Environmental Institute (IAP) (unpublished data) and by the water 
company of Paraná State (SANEPAR) (unpublished data). The Trophic 
State Index (TSI) was calculated according to Lamparelli (2004) 
(Zorzal-Almeida et al. 2017).

Samplings were carried out seasonally (i.e. in spring, summer, 
autumn, and winter) from October 2007 to August 2008, at two sampling 
stations. Periphytic samples were obtained from natural and artificial 
substrates. As natural substrates, we used stems of the macrophyte 
Polygonum hydropiperoides Michaux that were collected near the 
reservoir margin. The periphyton was obtained by scraping the stems 
with a razor blade wrapped in foil. Glass slides (7.5 cm x 2.5 cm) were 
used as artificial and inert substrates fixed to woody frames that were 
left submerged in the water column for 30 days. The artificial substrates 
were placed at about 80 meters far from the margin of the reservoir to 
prevent being eventually trapped in the macrophytes by the wind action.

Substrate fragments were fixed in Transeau’s solution (6:3:1) 
(Bicudo & Menezes 2017). The samples were treated by oxidation 
according to the method of Simonsen (1974) modified by Moreira-
Filho & Valente-Moreira (1981). The material was air-dried onto glass 
slides and mounted in Naphrax® resin. Observations, measurements, 
and diatom photomicrographs were performed at 1000× magnification 
in an Olympus BX40 microscope equipped with a DP71 Olympus 
camera. Diatom identification was performed up to the lowest level 
of taxonomic hierarchy based on recent and classic literature (e.g. 
Lange-Bertalot 1993, 1999, 2001, Lange-Bertalot & Metzeltin 1996, 
Metzeltin & Lange-Bertalot 1998, 2002, 2005, 2007, Bahls 2015). For 
each taxon, we included information on morphometry, remarks on the 
autoecology (when available in literature) as well as on the occurrence 
in the samples (see also table 2). Taxonomic comments were provided 
only for poorly known species in Brazil. We considered that a species is 
recorded for the first time in Brazil and in the state of Paraná if it was not 
previously recorded in a published article. The studied materials were 
deposited at the herbarium of the State University of Paraná (UPCB) 
under the numbers 63371 to 63374 (glass slides) and 63375 to 63378 
(macrophyte P. hydropiperoides).

Results and Discussion

The Piraquara I reservoir was classified as oligotrophic by the 
TSI index, characterized by low concentrations of phosphorus and 
nitrogen, high transparency of the water column (up to 3.2 meters), 
and a slightly acid to nearly neutral pH. Summer showed the highest 



3

Periphytic diatoms from an oligotrophic reservoir

Biota Neotrop., 19(2): e20180568, 2019

http://dx.doi.org/10.1590/1676-0611-BN-2018-0568 http://www.scielo.br/bn

Figure 1. Sampling site in Piraquara I reservoir, Paraná state, Brazil. Artificial Substrate (AS) and Natural Substrate (NS). (shapefile map from SUDERHSA 
2007).

values of temperature and accumulated rainfall occurred in the summer 
sample (27°C and 326.9 mm, respectively). The physical and chemical 
water parameters are shown in Table 1.

We identified 88 infrageneric taxa that belong to 35 genera. 
Pinnularia Ehrenberg (15 spp.) and Eunotia Ehrenberg (14 spp.) were 
the most representative diatoms in both substrates. The macrophyte 
P. hydropiperoides showed the highest richness (77) and the highest 
number of exclusive species (35) that represented 45.4% of the total 
substrate richness, whereas the artificial substrate presented 52 species 
and 10 exclusive taxa, which corresponded to 19.2% of the total 
substrate richness.

The species first recorded in Brazil were Achnanthidium 
neomicrocephalum Lange-Bertalot & Staab, Craticula riparia var. 
mollenhaurei Lange-Bertalot, Frustulia australocrassinervia Casa, 
Mataloni & Van de Vijver, Gomphonema spiculoides Gandhi and 
Stauroneis neohyalina Lange-Bertalot & Krammer. Seven diatoms are 
recorded for the first time in Paraná State: Craticula acidoclinata Lange-
Bertalot & Metzeltin, Fragilaria billingsii Wengrat, Wetzel & Morales, 
Eunotia longicamelus Costa, Bicudo & Wetzel, Eunotia pseudoimplicata 
Lange-Bertalot, Costa & Wetzel, Humidophila biscutella (Moser, 
Lange-Bertalot & Mezeltin) Lowe, Kociolek, Johansen, Van de Vijver, 

Table 1. Physical and chemical characteristics of the Piraquara I reservoir registered seasonally between October 2007 and August 2008.

Abiotic Variables spring
(October/2007)

summer
(February/2008)

autumn
(May/2008)

winter
(August/2008)

Temperature (°C) 20.8 27 18.6 16
Accummulated rainfall (mm) 96.6 326.9 200.5 104.7
Transparency (m) 3.2 2.6 2.9 2.3
Conductivity (μS.cm-1) 24.0 24.5 22.5 24.5
Turbidity (NTU) 1.9 1.9 1.5 2.0
pH 6.85 6.75 6.7 6.0
NO3 (μg.L-1) 95 40 45 35
NH4

+ (μg.L-1) 120 175 180 100
TN (μg.L-1) 1500.0 2400.0 800 600
TP (μg.L-1) 12 15 17 17
OD (mg.L-1) 6.05 5.35 7.77 7.7
Chl-a (μg.L-1) 3.06 3.09 1.55 1.55
TSI 50.9 51.6 50.3 50.3



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Lange-Bertalot & Kopalová, Nitzschia semirobusta Lange-Bertalot and 
Pinnularia viridiformis var. minor Krammer.

Pinnularia and Eunotia (15 and 14 species, respectively) presented 
a higher species richness in Piraquara I reservoir. The presence of 
these genera was favored by the slight acidity, low conductivity and 
water oligotrophy conditions of the studied reservoir (Round et al. 
1990, Metzeltin & Lange-Bertalot 1998, Spaulding & Edlund 2009, 
Costa et al. 2017). Eunotia was also the most representative genus in 
a eutrophic reservoir (Silva et al. 2010) located near to the Piraquara 
I. However, only three Eunotia species (E. naegelii, E. minor and E. 
subarcuatoides), which have a wide tolerance to trophic gradients (Van 
Dam et al. 1994, Silva et al. 2010, Costa et al. 2017) and they are able 
to occur under oligotrophic and eutrophic conditions, were common 
to both reservoirs.

The genera Navicula, Gomphonema, and Nitzschia, which are 
commonly registered in nutrient-rich environments (Goldsborough & 
Robinson 1996), showed a high species number in both substrates at the 
mesotrophic and hypereutrophic reservoirs located nearby to Piraquara 
I (Bertolii et al. 2010, Faria et al. 2010, Marra et al. 2016).

The richest assemblage was obtained from Polygonum 
hydropiperoides (natural substrate), 77 species, and can be explained 
by the glabrous or pubescent petiole of this macrophyte (Melo 2008), 
which offers a greater architectural complexity than the smooth and 
uniform surface of glass slides. The roughness difference between 
substrates usually does not contribute to an increase in algal biomass, 
but it is an important factor in selecting the species composition of the 
periphyton (Burkholder 1996, Souza & Ferragut 2012).

The diatom species richness found in the oligotrophic Piraquara 
I reservoir was lower than in the mesotrophic reservoirs Piraquara 
II and Passaúna (135 and 106 taxa, respectively), the eutrophic 
reservoir Iraí (96 taxa) and the hypereutrophic reservoir Itaqui (124 
taxa) (Bertolli et al. 2010, Silva et al. 2010, Faria 2010, Marra et al. 
2016). The Piraquara II is a mesotrophic reservoir located downstream 
from Piraquara I. Both reservoirs shared the presence of 31 diatom 
species, of which 11 were recorded from oligotrophic to mesotrophic 
environments and 20 species had a wide trophic tolerance. The diatom 
assemblage had more species that are tolerant in Piraquara II than in 
Piraquara I. Although eutrophication might lead to a loss in diatom 
diversity (Wengrat et al. 2017), the species richness in shallow lakes 
might be also influenced by the presence, abundance and diversity of 
macrophytes (Sayer et al. 1999, Bicudo et al. 2007). Piraquara II and 
Itaqui had more diatom richness than the other reservoirs, which can 
be related to the fact that in the former reservoir were sampled three 
different species of macrophytes (Marra et al. 2016), and that the latter 
reservoir was almost totally covered by the floating macrophyte Pistia 
stratiotes L. (Faria et al. 2010, Faria et al. 2013).

Summer showed the highest accumulated rainfall (326.9 mm) during 
the sampling period and the lowest species number in both substrates. 
High rainfall lead to the detachment of the periphyton from the substrates 
(Casartelli et al. 2016), which certainly contributed to the less number 
of species registered in Piraquara I. We registered nine diatom species 
(Spicaticribra kingstonii, Eunotia intermedia, Eunotia parasiolii, 
Eunotia pseudoimplicata, Encyonopsis subminuta, Cymbopleura 
naviculiformis, Planothidium incuriatum, Capartogramma crucicula 
and Nitzschia nana) that occurred in spring on both substrates, but were 

not recorded in summer. These species release mucilage and remain 
attached to the substrates and are therefore commonly observed from 
the periphytic samples (Round et al. 1990, Santos et al. 2011, Montoya-
Moreno et al. 2011). In summer, we registered temporary losses of 
species that were retrieved again during the sampling period. As shown in 
Table 2, the planktonic and benthic diatoms registered were Aulacoseira 
ambigua, Cyclotella meneghiniana, Fragilaria recapitellata, Fragilaria 
sp., Ulnaria delicatissima, Eunotia intricans, Eunotia paratridentula, 
Encyonopsis schubartii, Achnanthidium minutissimum, Achnanthidium 
sp., Sellaphora tridentula, Pinnularia brauniana, Pinnularia divergens 
var. media, Pinullaria obscura, Pinnularia pseudosudetica, Navicula 
leptostriata, Nitzschia acidoclinata and Nitzschia semirobusta.

Diatoms taxonomy and ecology

Family STEPHANODISCACEAE
Spicaticribra kingstonii Johansen, Kociolek & Lowe
Figure: 2A
Valve diameter: 14.8-15.2 µm; areolae: 13 in 10 µm; fultoportulae: 3-5 
per valve; rimoportulae: 1-3 per valve.
In Brazil, this species was described by Ludwig et al. (2008) as 
Thalassiosira rudis Tremarin, Ludwig, Becker & Torgan and later 
synonimized with S. kingstonii by Rivera & Cruces (2013).
Ecology: previously registered in Brazilian tropical and subtropical 
reservoirs with oligotrophic to eutrophic conditions (Bartozek et al. 
2018, Silva et al. 2010, Bertolli et al. 2010), and neutral pH with 
moderate conductivity (Ludwig et al. 2008). In our samples, it occurred 
in circumneutral pH (6.85), low conductivity (24 μS cm-1) and 
oligotrophic conditions, only in spring (temperature < 21°C),
Occurrence in samples: artificial substrates (UPCB 63371).

Cyclotella meneghiniana Kützing
Figure: 2B
Valve diameter: 10-12 µm; striae: 16 in 10 µm; rimoportulae: 1-2 per 
valve.
Ecology: it is recorded in oligotrophic to eutrophic environments, 
with a higher abundance in eutrophic waters (Bartozek et al. 2018, 
Silva et al. 2010) and in acidic waters (van Dam et al. 1994). In our 
samples, it occurred in slightly acid to circumneutral pH (6-6.85), low 
conductivity (24-24.5 μS cm1) and oligotrophic conditions, in spring 
and winter (temperature < 21°C).
Occurrence in samples: artificial (UPCB 63374) and natural (UPCB 
63375; UPCB 63378) substrates.

Discostella stelligera (Cleve & Grunow) Houk & Klee
Figure: 2C
Valve diameter: 12-21 µm; striae: 9.6-18.3 in 10 µm.
Ecology: common in oligo-mesotrophic environments (Yang & 
Dickman 1993, Potapova & Charles 2007), and in alkaline waters with 
high conductivity (Bartozek et al. 2018). It was common in our samples, 
in slightly acid to circumneutral pH (6-6.85), low conductivity (24-24.5 
μS cm-1) and oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63371; UPCB 63372; UPCB 
63373; UPCB 63374) and natural (UPCB 63375; UPCB 63376; UPCB 
63377; UPCB 63378) substrates.



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Periphytic diatoms from an oligotrophic reservoir

Biota Neotrop., 19(2): e20180568, 2019

http://dx.doi.org/10.1590/1676-0611-BN-2018-0568 http://www.scielo.br/bn

Table 2. Diatom species and seasonal occurrence on artificial and natural substrates in Piraquara I reservoir.

Species
Artificial substrates natural substrates

spring
(October/2007)

summer
(February/2008)

autumn
(May/2008)

winter
(August/2008)

spring
(October/2007)

summer
(February/2008)

autumn
(May/2008)

winter
(August/2008)

Spicaticribra kingstonii X - - - - - - -

Cyclotella meneghiniana - - - X X - - X

Discostella stelligera X X X X X X X X

Aulacoseira ambigua X - X X X X X X

A. tenella - - X X X X X X

Fragilaria billingsii X X X X X X X X

F. capucina - X X X - X X X

F. recapitellata X X X X X - - X

F. tenera X X X X - X X -

Fragilaria sp. X - X X - - - X

Staurosira construens - - - - - - - X

Staurosirella pinnata - - - X - - - -

Ulnaria delicatissima X X - X X - - X

U. ulna - - - - - X - -

Eunotia desmogonioides X X X - X X X X

E. intermedia X - - - - - - -

Eunotia intricans - - - - X - X -

E. longicamelus - - - - - - - X

E. minor - - - - - - - X

E. naegelii - - - - - - - X

E. parasiolii - - - - X - - -

E. paratridentula X - X - X X - X

E. pseudoimplicata X - - - X - - -

E. pseudosudetica X X X - X X X X

E. pyramidata f. capitata - - - - - - X -

E. rabenhorstii var. monodon - - - - - X X -

E. subarcuatoides - - - - X X X X

E. yanomami - - - - - - X -

Placoneis elginensis - - X - - - - -

Encyonema neogracile X X X X X X X X

E. silesiacum X - - - X X - X

Encyonopsis cf. reichardtii - X X - - X X X

E. schubartii X X X - X - X -

E. subminuta X - - - - - - -

Gomphonema gracile X X X X X X X X

G. lagenula - - - - - - X X

G. pantropicum - - - X - - - -

G. spiculoides X X X X X X X X

Cymbopleura naviculiformis - - - - X - - -

A. minutissium X - - X X - X X

A. neomicrocephalum - - - X - - - -

Achnanthidium sp. X - X - X X X X

Planothidium incuriatum - - - - X - - -

Humidophila biscutella - - X X - - - X

Luticola simplex - - - - - - - X

Luticola permuticoides - - - - - X - -



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Species
Artificial substrates natural substrates

spring
(October/2007)

summer
(February/2008)

autumn
(May/2008)

winter
(August/2008)

spring
(October/2007)

summer
(February/2008)

autumn
(May/2008)

winter
(August/2008)

Frustulia australocrassinervia X X X X X X X X

F. undosa - - X X - X X -

Brachysira neoexilis X X X X X X X X

Neidium affine - - - - - - X -

Sellaphora rectangularis - - X X - - X -

S. tridentula - - - - X - X -

Pinnularia borealis - - - X X - - -

P. brauniana X - X X X - X X

P. divergens var. divergens - - - - - - X -

P. divergens var. media - - - - X - X X

P. grunowii - - X - X X - -

P. obscura - - - - X - X -

P. pseudogibba X - - X X X X -

P. rumrichae - - X - X X - X

P. schoenfelderi - - - - - - X -

P. subcapitata var. elongata - - - X - - - -

P. subcapitata var. subrostrata - - X - X X X -

P. subgibba var. undulata - - X - X X X -

P. submicrostauron - - X - - X X -

P. viridiformis - - - - X X - X

P. viridiformis var. minor - - - - - - - X

Chamaepinnularia 
brasilianopsis - - - - - - X -

C. mediocris - X - - - - X X

Kobayasiella sp. - - - - - - X -

Navicula cryptocephala X X X - X X - X

N. cryptotenella X X X X X X X X

N. leptostriata - - - X X - X -

N. veneta - - - - - - X -

Capartogramma crucicula X - - - - - - -

Stauroneis neohyalina - - X - - X - X

S. gracilis - - - - - - X -

S. phoenicenteron - - X X - - X -

Craticula acidoclinata - - - - - - X -

C. riparia var. mollenhaurei - - - - - - X -

C. submolesta - - - - - - X -

Nitzschia acidoclinata - - X X X - - X

N. nana - - - - X - - -

N. semirobusta X - - X X X X X

Hantzschia amphyoxis - - - - - X - -

Rhopalodia gibberula var. 
vanheurckii - - - - - - - X

Surirella grunowii X - - - - - X -

Iconella curvula - - - - - X - -

Total of species 30 18 34 31 42 35 49 41

Continuation Table 2.



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Periphytic diatoms from an oligotrophic reservoir

Biota Neotrop., 19(2): e20180568, 2019

http://dx.doi.org/10.1590/1676-0611-BN-2018-0568 http://www.scielo.br/bn

Figure 2. A. Spicaticribra kingstonii; B. Cyclotella meneghiniana; C. Discostella stelligera; D–E. Aulacoseira ambigua; F–G. Aulacoseira tenella; H–I. Fragilaria 
billingsii; J. Fragilaria capucina; K–L. Fragilaria recapitellata; M–O. Fragilaria tenera; P. Staurosirella pinnata; Q–R. Fragilaria sp., S–T. Staurosira construens; 
U–V. Eunotia parasiolii; W–X. Eunotia pseudosudetica; Y–Z. Eunotia paratridentula; AA–AB. Eunotia intermedia; AC. Eunotia pyramidata f. capitata; AD. Eunotia 
rabenhorstii var. monodon; AE. Ulnaria delicatissima; AF. Ulnaria ulna; AG–AH. Eunotia desmogonioides; AI. Eunotia intricans; AJ–AK. Eunotia pseudoimplicata; 
AL–AN. Eunotia longicamelus; AO–AP. Eunotia minor; AQ. Eunotia subarcuatoides; AR. Eunotia naegelii; AS. Eunotia yanomami.



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Biota Neotrop., 19(2): e20180568, 2019

http://www.scielo.br/bn http://dx.doi.org/10.1590/1676-0611-BN-2018-0568

Family AULACOSEIRACEAE
Aulacoseira ambigua (Grunow) Simonsen
Figures: 2D–2E
Valve diameter: 4-9.4 µm; striae: 8-17 in 10 µm; areolae: 8-17 in 10 µm.
Ecology: this species has a wide ecological tolerance, registered in 
oligotrophic (Stenger-Kovács et al. 2007), mesotrophic (Bicudo et al. 
2016) and meso-eutrophic (Gómez & Licursi 2001, Ivanov & Kirilova 
2004, Dela-Cruz et al. 2006) environments. In our samples, it occurred 
in slightly acid to circumneutral pH (6-6.85), low conductivity (24-24.5 
μS cm-1) and oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63371; UPCB 63373; UPCB 
63374) and natural (UPCB 63375; UPCB 63376; UPCB 63377; UPCB 
63378) substrates.

Aulacoseira tenella (Nygaard) Simonsen
Figures: 2F–2G
Valve diameter: 6.3-8 µm; striae: 15 in 10 µm.
Ecology: common in oligotrophic to mesotrophic environments (Bicudo 
et al. 2016, Siver & Kling 1997). In our samples, it occurred in slightly 
acid to circumneutral pH (6-6.85), low conductivity (24-24.5 μS cm-1) 
and oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63373; UPCB 63374) and 
natural (UPCB 63375; UPCB 63376; UPCB 63377; UPCB 63378) 
substrates.

Family FRAGILARIOPHYCEAE
Fragilaria billingsii Wengrat, Wetzel & Morales
Figures: 2H–2I
Length: 66-73.6 µm; width (central region): 2-2.3 µm; striae: 22 in 
10 µm.
Fragilaria billingsii is characterized by long and narrow valves with a 
bilaterally inflated central area. The striae are distinct and the specimes 
found in the present study have a higher striae density than those 
registered by Wengrat et al. (2016) (striae 17-20 in 10 µm).
Ecology: previously registered only for meso-eutrophic environments, 
with slightly acid pH and low conductivity (Wengrat et al. 2016) In our 
samples, it occurred in slightly acid to circumneutral pH (6-6.85), low 
conductivity (24-24.5 μS cm-1) and oligotrophic conditions.
This is the first record for Paraná state.
Occurrence in samples: artificial (UPCB 63371; UPCB 63372; UPCB 
63373; UPCB 63374) and natural (UPCB 63375; UPCB 63376; UPCB 
63377; UPCB 63378) substrates.

Fragilaria capucina Desmazières
Figure: 2J
Length: 27-34.2 µm; width: 2.8-2.9 µm; striae: 15-16 in 10 µm.
The taxonomy of Fragilaria capucina is very confusing. Based on the 
type materials, there are three different morphotypes of F. capucina 
documented in the literature. One of the morphotypes has lanceolate 
valves; the second one has linear to lanceolate valves that are shorter 
than in the other two morphotypes, and the third one has a linear valve 
with a rhombic central area and slightly rostrate valve ends (Tuji & 
William 2006, Krammer & Lange-Bertalot 1991a).
Ecology: probably cosmopolitan, found in oligo-mesotrophic (Delgado 
et al. 2015) to mesotrophic (vam Dam et al. 1994) and eutrophic 

environments (Hofmann 1994). Considered as a planktonic, benthic 
and terrestrial species (Delgado et al. 2015). In our samples, it occurred 
attached to artificial substrates and macrophytes, in slightly acid to 
circumneutral pH (6-6.75), low conductivity (24-24.5 μS cm-1) and 
oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63372; UPCB 63373; 
UPCB 63374) and natural (UPCB 63376; UPCB 63377; UPCB 63378) 
substrates.

Fragilaria recapitellata Lange-Bertalot & Metzeltin
Figures: 2K–2L
Length: 22.2-30.4 µm; width: 2-4 µm; striae: 16-18 in 10 µm.
This species has lanceolate valves with capitate ends and a unilateral 
central area (Metzeltin et al. 2009).
Ecology: this species is fairly common in alpha-mesosaprobic and 
eutrophic conditions (Cantonati et al. 2017, van Dam et al. 1994). In 
contrast to previous studies, it was common in our samples, indicating 
a tolerance to oligotrophic waters, poor in nutrients, with slightly acid 
to circumneutral pH (6-6.85), and low conductivity (24-24.5 μS cm-1). 
Although it is cosmopolitan and often present in periphyton samples 
from lakes, rivers, and streams of temperate regions (Delgado et al. 
2015), it is rarely found in tropical regions (Silva et al. 2010, Tremarin 
et al. 2009).
Occurrence in samples: artificial (UPCB 63371; UPCB 63372; 
UPCB 63373; UPCB 63374) and natural (UPCB 63375; UPCB 63378) 
substrates.

Fragilaria tenera (W. Smith) Lange-Bertalot
Figures: 2M–2O
Length: 38.8-51.8 µm; width: 2.3-2.8 µm; striae: 15 in 10 µm.
The specimens analyzed in the present study are shorter, with a lower 
striae density, and the absence of a central area. Hofmann et al. (2013) 
referred specimens of 30-100 µm long, 2-3 µm wide and with a striae 
density of 17-20 in 10 µm, and the often absence of a central area.
Ecology: this species was registered for oligo-mesotrophic environments 
(van Dam et al. 1994). In our samples it occurred attached to the 
substrates in slightly acid to circumneutral pH (6-6.85), low conductivity 
(24-24.5 μS cm-1) and oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63371; UPCB 63372; 
UPCB 63373; UPCB 63374) and natural (UPCB 63376; UPCB 63377) 
substrates.

Fragilaria sp.
Figures: 2Q–2R
Length: 24.5-31.7 µm; width: 2.3-3.1 µm; striae: 16-17 in 10 µm.
Fragilaria sp. resembles Fragilaria austriaca (Grunow) Lange-Bertalot 
in the outline but has wider valves (3-4 µm) and a lower striae density 
(12-15 µm). Fragilaria pararumpens Lange-Bertalot, Hofmann & 
Werum has similar dimensions to Fragilaria sp., but a more lanceolate 
valve with subcapitate ends (Cantonati et al. 2017).
Ecology: In our samples, it occurred only in temperatures < 21°C, 
slightly acid to circumneutral pH (6-6.75), low conductivity (22.5-24.5 
μS cm-1) and oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63371; UPCB 63373; UPCB 
63374) and natural (UPCB 63378) substrates.



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Periphytic diatoms from an oligotrophic reservoir

Biota Neotrop., 19(2): e20180568, 2019

http://dx.doi.org/10.1590/1676-0611-BN-2018-0568 http://www.scielo.br/bn

Family STAUROSIRACEAE
Staurosira construens Ehrenberg
Figures: 2S–2T
Length: 15.7 µm; width: 8.6 µm; striae: 18 in 10 µm.
Ecology: this species is registered for meso-eutrophic environments 
(van Dam et al. 1994) and is considered as tolerant to different water 
conditions (Hofmann 1994). In our samples, it occurred in slighltly acid 
pH (6), low conductivity (24.5 μS cm-1) and oligotrophic conditions, 
only in winter (temperature 16°C).
Occurrence in samples: natural substrates (UPCB 63378).

Staurosirella pinnata (Ehrenberg) Williams & Round.
Figure: 2P
Length: 8-14.4 µm; width: 3.3-3.8 µm; striae: 8-11 in 10 µm.
Ecology: this species is considered as tolerant (Hofmann 1994) to 
indifferent (van Dam et al. 1994) to the trophic state. In our samples, it 
occurred in slighltly acid pH (6), low conductivity (24.5 μS cm-1) and 
oligotrophic condition, only in winter (temperature 16°C).
Occurrence in samples: artificial substrates (UPCB 63374).

Family ULNARIACEAE
Ulnaria delicatissima (Smith) Aboal & Silva
Figure: 2AE
Length: 62.4-109.5 µm; width: 2.4-4.7 µm; striae: 13-16 in 10 µm.
Ecology: common in oligo-mesotrophic environments (van Dam et 
al. 1994, Hofmann 1994). In our samples, it occurred in slighltly acid 
to circumneutral pH (6-6.85), low conductivity (24-24.5 μS cm-1) and 
oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63371; UPCB 63372; UPCB 
63374) and natural (UPCB 63375; UPCB 63378) substrates.

Ulnaria ulna (Nitzsch) Compère.
Figure: 2AF
Length: 90-189 µm; width: 4-4.8 µm; striae: 9-14 in 10 µm.
Ecology: considered as indifferent to tolerant to the trophic state of the 
environment (van Dam et al. 1994, Hofmann 1994). In our samples, it 
occurred in circumneutral pH (6.75), low conductivity (24.5 μS cm-1) 
and oligotrophic conditions, only in summer (temperature of 27°C and 
accumulated rainfall 326.9 mm).
Occurrence in samples: natural substrates (UPCB 63376).

Family EUNOTIACEAE
Eunotia desmogonioides Metzeltin & Lange-Bertalot
Figures: 2AG–2AH
Length: 99-159.3 µm; width: 3.5-5 µm; striae: 17-20 in 10 µm.
This species has a valve outline similar to Eunotia flexuosa (Brébisson) 
Kützing, although with narrower valves and a higher striae density. 
Eunotia rabenhorstiana var. elongata (Patrick) Metzeltin & Lange-
Bertalot is similar to E. flexuosa but has wider valves with more inflated 
ends (Costa et al. 2017).
Ecology: previously registered for Brazilian oligo- to mesotrophic 
reservoirs, with low conductivity and slightly acidic to neutral pH 
(Costa et al. 2017). In our samples, it occurred in slightly acid to 
circumneutral pH (6-6.85), low conductivity (22.5-24.5 μS cm-1) and 
oligotrophic conditions.

Occurrence in samples: artificial (UPCB 63371, UPCB 63372, UPCB 
63373) and natural (UPCB 63375, UPCB 63376, UPCB 63377, UPCB 
63378) substrates.

Eunotia intermedia (Krasske ex Hustedt) Nörpel & Lange-Bertalot
Figures: 2AA–2AB
Length: 13.3-16.8 µm; width: 3.1-3.3 µm; striae: 14-15 in 10 µm.
Ecology: previously registered for slightly acidic waters with low 
conductivity (Lange-Bertalot et al. 2011). In our samples, it occurred in 
circumneutral pH (6.85), low conductivity (24 μS cm-1) and oligotrophic 
conditions, only in spring (temperature < 21°C).
Occurrence in samples: artificial substrate (UPCB 63371).

Eunotia intricans Lange-Bertalot & Metzeltin
Figure: 2AI
Length: 35-51.7 µm; width: 4.5-5.1 µm; striae: 11-13 in 10 µm.
Eunotia intricans is similar to Eunotia parasiolii Metzeltin & Lange-
Bertalot in the valve outline, although it has a lower striae density 
(6.5-11/10 µm, according to Metzeltin & Lange-Bertalot 1998). In 
Piraquara reservoir, the specimens were longer than the type specimen 
(17-48 µm, according to Lange-Bertalot & Metzeltin 2009), but have 
a similar length to other specimens registered in Brazilian reservoirs 
(Costa et al. 2017).
Ecology: previously registered for oligo-mesotrophic environments, 
with a preference for acid waters with low conductivity (Costa et al. 
2017). In our samples, it occurred in circumneutral pH (6.7-6.85), low 
conductivity (22.5-24 μS cm-1) and oligotrophic conditions, in spring 
and autumn (temperature < 21°C).
Occurrence in samples: natural substrates (UPCB 63375, 63377).

Eunotia longicamelus Costa, Bicudo & Wetzel
Figure: 2AL–2AN
Length: 16.2-52.6 µm; width: 5.4-5.26 µm; striae: 11 in 10 µm; valvar 
undulations: 2-4.
Similar to Eunotia camelus Ehrenberg, but differs in the type and number 
of undulations on the dorsal margin (two undulations subdivided into 
two more) and the presence of wider valves (length: 22-30 µm; width: 
6-8.8 µm; striae density 8-12 in 10 µm and two undulations) (Costa 
et al. 2017, Reichardt 1995).
Ecology: previously registered for mesotrophic environments, with 
a preference for slightly acidic waters (pH 6.6) with medium values 
of conductivity (71 µS cm-1) (Costa et al. 2017). In our samples, it 
occurred only in winter (temperature 16°C) and expanded the occurrence 
to oligotrophic environments with slightly acid pH (6), and low 
conductivity (24.5 μS cm-1).
This is the first record for Paraná state.
Occurrence in samples: natural substrates (UPCB 63378).

Eunotia minor (Kützing) Grunow.
Figures: 2AO–2AP
Length: 17-24.7 µm; width: 3.8 µm; striae: 17-18 in 10 µm.
Ecology: considered as tolerant to different trophic states, ranging 
from oligo- to eutrophic (van Dam et al. 1994, Hofmann 1994). In our 
samples, it occurred in slightly acid pH (6), low conductivity (24.5 μS 
cm-1) and oligotrophic conditions, only in winter (temperature 16°C).
Occurrence in samples: natural substrates (UPCB 63378).



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Eunotia naegelii Migula.
Figure: 2AR
Length: 40-48.4 µm; width: 1.8-3 µm; striae: 21-24 in 10 µm.
Ecology: considered as acidophile; registered for ultra-oligotrophic to 
oligotrophic environments (Costa et al. 2017, van Dam et al. 1994). In 
our samples, it occurred in slightly acid pH (6), low conductivity (24.5 
μS cm-1) and oligotrophic conditions, only in winter (temperature 16°C).
Occurrence in samples: natural substrates (UPCB 63378).

Eunotia parasiolii Metzeltin & Lange-Bertalot.
Figures: 2U–2V
Length: 18.5-32.6µm; width: 3-4.7 µm; striae: 9-12 in 10 µm.
Ecology: previously registered for acidic waters (pH 4.4-5.3) with a low 
conductivity (9.4-12.6 µS cm-1) (Ferrari et al. 2007) to slightly acidic 
(pH 6.6-6.8) and oligotrophic environments (Costa et al. 2017). In our 
samples, it occurred in circumneutral pH (6.85), low conductivity (24 μS 
cm-1) and oligotrophic conditions, only in spring (temperature < 21°C).
Occurrence in samples: natural substrates (UPCB 63375).

Eunotia paratridentula Lange-Bertalot & Kulikovskyi.
Figures: 2Y–2Z
Length: 13-17.6 µm; width: 2.8-4 µm; striae: 14-16 in 10 µm; 
undulations: 3-4.
Previously reported for Paraná State as Eunotia muscicola Krasske var. 
tridentula Nörpel & Lange-Bertalot (Tremarin et al. 2009). It differs 
from E. muscicola Krasske by the lower striae density; and from E. 
paramuscicola Krstić, Levkov & Pavlov by the broadly protracted 
apices (Krstić et al. 2013).
Ecology: considered as acidophile (van Dam et al. 1994); registered 
for mesotrophic environments (Marra et al. 2016). Our study expanded 
its occurrence to environments with pH ranging from slightly acid 
to circumneutral (6-6.85), low conductivity (22.5-24.5 μS cm-1) and 
oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63371; UPCB 63373) and 
natural (UPCB 63375; UPCB 63376; UPCB 63378) substrates.

Eunotia pseudoimplicata Lange-Bertalot, Costa & Wetzel
Figures: 2AJ–2AK
Length: 28.7-43 µm; width: 4.2-5.3 µm; striae: 14-16 in 10 µm.
Ecology: oligo-mesotrophic reservoirs, with slightly acid to neutral 
waters (Costa et al. 2017). In our samples, it occurred in circumneutral 
pH (6.85), low conductivity (24 μS cm-1) and oligotrophic conditions, 
only in spring (temperature < 21°C).
This is the first record for Paraná state.
Occurrence in samples: artificial (UPCB 63371) and natural (UPCB 
63375) substrates.

Eunotia pseudosudetica Metzeltin, Lange-Bertalot & García-Rodríguez
Figures: 2W–2X
Length: 30.4-33.26 µm; width: 4.9-5.4 µm; striae: 12 in 10 µm.
Ecology: this species occurred in slightly acidic to circumneutral 
pH (6-6.85), low conductivity (22.5-24 μS cm-1) and in oligotrophic 
conditions, in agreement with previously studies (Costa et al. 2017), 
but can also be registered for nutrient-rich environments (Bertolli et 
al. 2010, Faria et al. 2010, Marra et al. 2016). Usually found in the 

periphyton, but can be also registered from phytoplankton (Bartozek 
et al. 2013) and surface sediments (Costa et al. 2017).
Occurrence in samples: artificial (UPCB 63371; UPCB 63372; UPCB 
63373) and natural (UPCB 63375; UPCB 63376; UPCB 63377; UPCB 
63378) substrates.

Eunotia pyramidata f. capitata Krasske
Figure: 2AC
Length: 25.5 µm; width: 4.6 µm; striae: 15 in 10 µm; undulations: 3.
It differs from Eunotia rabenhorstii var. triodon Cleve & Grunow by 
the presence of a more pronounced mark in the central undulation 
(Costa et al. 2017).
Ecology: previously registered for moody pools with sphagnum moss 
(Krasske 1948), and from samples of surface sediments collected in 
environments with acid pH and poor in nutrients (Costa et al. 2017). 
In our samples, it occurred in circumneutral pH (6.7), low conductivity 
(22.5 μS cm-1) and oligotrophic conditions, only in autumn (temperature 
18.6°C).
Occurrence in samples: natural substrates (UPCB 63377).

Eunotia rabenhorstii var. monodon Cleve & Grunow
Figure: 2AD
Length: 20 µm; width: 6.2 µm; striae: 17 in 10 µm.
Ecology: previously registered for a mesotrophic reservoir in Paraná 
state (Bertolli et al. 2010, Marra et al. 2016) and for ultraoligotrophic 
to eutrophic reservoirs with slightly acidic to alkaline waters (pH 6.2-
7.3) in São Paulo state (Costa et al. 2017). In our samples, it occurred 
in circumneutral pH (6.7-6.75), low conductivity (22.5-24.5 μS cm-1) 
and oligotrophic conditions, in summer and autumn (temperature from 
18.6-27°C).
Occurrence in samples: natural substrates (UPCB 63376; UPCB 
63377).

Eunotia subarcuatoides Alles, Nörpel & Lange-Bertalot
Figure: 2AQ
Length: 13.5-17.3 µm; width: 2.4-2.9 µm; striae: 23-24 in 10 µm.
Similar to the small forms of Eunotia bilunaris (Ehrenberg) 
Schaarschmidt, but with a lower density of striae (13-17 in 10 µm) that 
are less conspicuous (Costa et al. 2017). The specimens found herein 
presented a less width than the type material (3-4.5 µm, according 
to Alles et al. 1991), but presented a similar width to the specimens 
(2.5-3.3 µm) registered by Costa et al. (2017) in Brazilian reservoirs.
Ecology: considered as acidobiontic (van Dam et al. 1994). In Brazil, it 
was previously registered for ultraoligo to eutrophic conditions (Costa et 
al. 2017, Silva et al. 2010), with acid to alkaline pH (Costa et al. 2017). 
In our samples, it occurred in slightly acid to circumneutral pH (6-6.85), 
low conductivity (22.5-24.5 μS cm-1) and oligotrophic conditions.
Occurrence in samples: natural substrates (UPCB 63375; UPCB 
63376; UPCB 63377; UPCB 63378).

Eunotia yanomami Metzeltin & Lange-Bertalot
Figure: 2AS
Length: 120 µm; width: 13.5 µm; striae: 15 in 10 µm; undulations: 4.
Similar to Eunotia zygodon Ehrenberg in the valve outline, but E. 
zygodon has a smaller size (less than 55 μm in length, according to 
Metzeltin & Lange-Bertalot 1998).



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http://dx.doi.org/10.1590/1676-0611-BN-2018-0568 http://www.scielo.br/bn

Ecology: previously registered for oligotrophic to mesotrophic 
environments (Marra et al. 2016, Costa et al. 2017), with slightly acid 
waters (pH 5.4-6.4) and low to medium conductivity (12-70 μS cm-1) 
(Bicca & Torgan 2009). In our samples, it occurred in circumneutral 
pH (6.7), low conductivity (22.5 μS cm-1) and oligotrophic conditions, 
only in autumn (temperature 18.6°C).
Occurrence in samples: natural substrates (UPCB 63377).

Family GOMPHONEMATACEAE
Placoneis elginensis (Gregory) Cox
Figure: 3A
Length: 34.7 µm; width: 10 µm; striae: 13 in 10 µm.
Ecology: previously registered for highly alkaline waters, and for 
mesotrophic and eutrophic environments (van Dam et al. 1994, 
Hofmann 1994, Poulíčková et al. 2008). In our samples, it occurred in 
circumneutral pH (6.7), low conductivity (22.5 μS cm-1) and oligotrophic 
conditions, only in autumn (temperature 18.6°C).
Occurrence in samples: artificial substrates (UPCB 63373).

Encyonema neogracile Krammer.
Figures: 3B–3C
Length: 35.5-56.6 µm; width: 6.5-7.5 µm; dorsal striae: 14 in 10 µm; 
ventral striae: 15-16 in 10 µm.
Ecology: this species was frequent in our samples and occurred in 
slightly acid to circumneutral pH (6-6.85), low conductivity (22.5-24.5 
μS cm-1) and oligotrophic conditions, but was also previously registered 
for oligo-mesotrophic and mesotrophic environments (van Dam et al. 
1994, Marra et al. 2016).
Occurrence in samples: artificial (UPCB 63371; UPCB 63372; UPCB 
63373; UPCB 63374) and natural (UPCB 63375; UPCB 63376; UPCB 
63377; UPCB 63378) substrates.

Encyonema silesiacum (Bleisch) Mann.
Figure: 3D
Length: 17.4-21.5 µm; width: 5-7.5 µm; dorsal striae: 11-17 in 10 µm; 
ventral striae: 12-16 in 10 µm.
Ecology: widely distributed, considered as a circumneutral species (van 
Dam et al. 1994), occurring in waters ranging from oligo to eutrophic 
(van Dam et al. 1994, Hofmann 1994, Bertolli et al. 2010, Silva et al. 
2010, Marra et al. 2016). This study expands its occurrence to slightly 
acid pH (6-6.85), low conductivity (24-24.5 μS cm-1) and oligotrophic 
conditions.
Occurrence in samples: artificial (UPCB 6371) and natural (UPCB 
63375; UPCB 63376; UPCB 63378) substrates.

Encyonopsis cf. reichardtii Krammer
Figures: 3Q–3S
Length: 36.5-40 µm; width: 5.8-6 µm; maximum ratio L/W= 6.8; dorsal 
striae: 18-20 in 10 µm; ventral striae: 17-21 in 10 µm.
The specimens from Piraquara I reservoir presented the valve outline, 
striation pattern, and isolated punctum near the central striae similar 
to Encyonopsis reichardtii Krammer. As the type specimens have a 
slightly more rhombo-lanceolate contour, wider valves (7-7.7 µm) and 
a maximum ratio L/W of 6.1 (Krammer 1997b: 131) we preferred to 
identify the taxon as conferatum.

Ecology: this species occurred in slightly acid to circumneutral pH (6-
6.75), low conductivity (22.5-24.5 μS cm-1) and oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63372; UPCB 63373) and 
natural (UPCB 63376; UPCB 63377; UPCB 63378) substrates.

Encyonopsis schubartii (Hustedt) Krammer
Figure: 3E
Length: 29.5-33.6 µm; width: 6-6.4 µm; dorsal striae: 9-11 in 10 µm; 
ventral striae: 9-12 in 10 µm.
Ecology: previously registered for oligotrophic (Krammer 1997b), and 
mesotrophic environments, with acid pH (Wengrat et al. 2015). In our 
samples, it occurred in circumneutral pH (6.7-6.85), low conductivity 
(22.5-24.5 μS cm-1) and oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63371; UPCB 63372; UPCB 
63373) and natural (UPCB 63375; UPCB 63377) substrates.

Encyonopsis subminuta Krammer & Reichardt.
Figure: 3F
Length: 15.3-21.3 µm; width: 3.7-4.8 µm; striae: 26 in 10 µm.
Ecology: common in oligo- and mesotrophic environments, and rare 
in electrolyte-poor environments (Bertolli et al. 2010, Cantonati et al. 
2017). In our samples, it occurred in circumneutral pH (6.85), low 
conductivity (24 μS cm-1) and oligotrophic conditions, only in spring 
(temperature 20.8°C).
Occurrence in samples: artificial substrate (UPCB 63371).

Gomphonema gracile Ehrenberg
Figure: 3J–3K
Length: 28-43 µm; width: 5.8-7.2 µm; striae: 14-16 in 10 µm.
The Gomphonema gracile species-complex includes representatives 
with valves of diverse forms, sizes and measures. Gomphonema 
naviculoides Smith is very similar to G. gracile in the valve outline 
and striae density, but they differ in the width range, being of 4.2-9 µm 
in G. gracile and of 7.8-9.5 µm in G. naviculoides (Reichardt 2015a).
Ecology: typical of environments with low conductivity and nutrients 
(Krammer & Lange-Bertalot 1986). It occurred in all our samples, in 
slightly acid to circumneutral pH (6-6.85), low conductivity (22.5-24.5 
μS cm-1) and oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63371; UPCB 63372; UPCB 
63373; UPCB 63374) and natural (UPCB 63375; UPCB 63376; UPCB 
63377; UPCB 63378) substrates.

Gomphonema lagenula Kützing
Figures: 3L–3M
Length: 17.2-21.7 µm; width: 4.5-4.8 µm; striae: 18-19 in 10 µm
Ecology: considered as tolerant (Hofmann 1994), this species is widely 
distributed in environments ranging from oligotrophic to hypereutrophic 
(Bertolli, et al. 2010, Silva et al. 2010, Faria et al. 2010, Marra et al. 
2016, Bartozek et al. 2018). In our samples, it occurred in slightly acid 
to circumneutral pH (6-6.7), low conductivity (22.5-24.5 μS cm-1) and 
oligotrophic conditions, in autumn and winter (temperature < 19°C).
Occurrence in samples: natural substrates (UPCB 63377; UPCB 
63378).



12

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Figure 3. A. Placoneis elginensis; B–C. Encyonema neogracile; D. Encyonema silesiacum; E. Encyonopsis schubartii; F. Encyonopsis subminuta; G–I. Gomphonema 
spiculoides; J–K. Gomphonema gracile; L–M. Gomphonema lagenula; N–O. Gomphonema pantropicum; P. Cymbopleura naviculiformis; Q–S. Encyonopsis 
cf. reichardtii; T–W. Achnanthidium minutissimum; X–AD. Achnanthidium neomicrocephalum; AE–AH. Achnanthidium sp.; AI–AJ. Planothidium incuriatum; 
AK. Humidophila biscutella; AL. Luticola permuticoides; AM. Luticola simplex; AN–AP. Frustulia australocrassinervia; 3AQ–3AS. Frustulia undosa; AT–AV. 
Brachysira neoexilis; AW–AX. Neidium affine.



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http://dx.doi.org/10.1590/1676-0611-BN-2018-0568 http://www.scielo.br/bn

Gomphonema pantropicum Reichardt
Figure: 3N–3O
Length: 35.2-38 µm; width: 6-6.3 µm; striae: 10-11 in 10 µm.
Valves lanceolate, with constricted round apices. Spaced and convergent 
striae towards the middle part of the valves and parallel to the apices.
In Brazilian environments, Gomphonema pantropicum is often 
misidentified as Gomphonema subtile Ehrenberg (Osorio et al. 2017, 
Marra et al. 2016, Marquardt & Bicudo 2014). Gomphonema subtile 
presents the valves with broadly capitate ends, a long and narrow “neck” 
and flatly-rounded apices (Reichardt 2015b).
Ecology: distributed in South America and in tropical Africa, India, 
and Southeast Asia (Reichardt 2015b); the ecology of this species is 
little known. In Brazil, it was previously registered from epiphytic 
samples in the North and South region (Medeiros et al. 2018, Azevedo 
et al. 2018). In our samples, it occurred in slightly acid pH (6), low 
conductivity (24.5 μS cm-1) and oligotrophic conditions, only in winter 
(temperature 16°C).
Occurrence in samples: artificial substrates (UPCB 63374).

Gomphonema spiculoides Gandhi
Figures: 3G–3I
Length: 54.2-69 µm; width: 7.3-8 µm; striae: 14 in 10 µm.
This species presents valve ends clearly apiculate, a central area weakly 
delimited, a narrow axial area, undulate and lateral raphe, and striae 
slightly radiate.
The specimens recorded in the present study have striae slightly radiate 
and a less width than the specimens found by Karthick & Kociolek 
(2012), resembling the Gandhi’s original draw and description. 
Gomphonema spiculoides can be mistaken with large valves of 
Gomphonema gracile Ehrenberg.
Ecology: this species has a low tolerance to alkaline conditions and 
it was previously registered for oligotrophic and slightly acid waters 
(Riato et al. 2017). It occurred in all our samples, in slightly acid to 
circumneutral pH (6-6.85), low conductivity (22.5-24.5 μS cm-1) and 
oligotrophic conditions.
This is the first record for Brazil.
Occurrence in samples: artificial (UPCB 63371; UPCB 63372; UPCB 
63373; UPCB 63374) and natural (UPCB 63375; UPCB 63376; UPCB 
63377; UPCB 63378) substrates.

Family CYMBELLACEAE
Cymbopleura naviculiformis (Auerswald) Krammer
Figure: 3P
Length: 32.1 µm; width: 9 µm; dorsal striae: 13 in 10 µm; ventral 
striae: 15 in 10 µm.
Ecology: previously registered for oligotrophic to eutrophic waters 
(Bertolli et al. 2010, Silva et al. 2010, Marra et al. 2016, Cantonati et 
al. 2017). In our samples, it occurred in circumneutral pH (6.85), low 
conductivity (24 μS cm-1) and oligotrophic conditions, only in spring 
(temperature 20.8°C).
Occurrence in samples: natural substrates (UPCB 63375).

Family ACHNANTHIDIACEAE
Achnanthidium minutissimum (Kützing) Czarnecki
Figures: 3T–3W
Length: 9.6-15.4 µm; width: 2.6-2.8 µm; indistinct striae.

Ecology: very wide range of ecological preferences (Cantonati et al. 
2017). In our samples, it occurred in temperature < 21°C, slightly acid 
to circumneutral pH (6-6.85), low conductivity (22.5-24.5 μS cm-1) and 
oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63371; UPCB 63374) and 
natural (UPCB 63375; UPCB 63377; UPCB 63378) substrates.

Achnanthidium neomicrocephalum Lange-Bertalot & Staab
Figures: 3X–3AD
Length: 21.6-25.3 µm; width: 2.5-2.8 µm; indistinct striae.
Valves linear-lanceolate with subcapitate apices. Axial area narrow. 
Central area variable in shape.
Achnanthidium neomicrocephalum has narrower valves than 
Achnanthidium gracillimum (Meister) Lange-Bertalot (3.4-4.5 µm) 
and A. caledonicum (Lange-Bertalot) Lange-Bertalot (2.7-3.8 µm). 
Achnanthidium caledonicum presents broader valve ends than A. 
neomicrocephalum.
Ecology: previously registered for oligotrophic environments, mainly 
springs and lakes (Cantonati et al. 2017, Cantonati & Lange-Bertalot 
2010). In our samples, it occurred in slightly acid pH (6) and low 
conductivity (24.5 μS cm-1), and oligotrophic condition, only in winter 
(temperature 16°C).
This is the first record for Brazil
Occurrence in samples: artificial substrates (UPCB 63374).

Achnanthidium sp.
Figures: 3AE–3AH
Length: 13.3-15.4 µm; width: 2.4-2.5 µm; indistinct striae.
Valves linear-lanceolate, with parallel margins and capitate apices. 
Narrow and linear axial area. Central area bordered by one striae on 
both sides. Resembles Achnanthidium tropicocatenatum Marquardt, 
Wetzel & Ector although the central portion is not inflated (Marquard 
et al. 2017).
Ecology: this species was frequent in ours samples, and occurred in 
slightly acid to circumneutral pH (6-6.85), low conductivity (22.5-24.5 
μS cm-1) and oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63371; UPCB 63373) and 
natural (UPCB 63375; UPCB 63376; UPCB 63377; UPCB 63378) 
substrates.

Planothidium incuriatum Wetzel, Van de Vijver & Ector
Figures: 3AI–3AJ
Length: 20 µm; width: 7 µm; striae: 14 in 10 µm.
Planothidium incuriatum is widely reported in Brazil as P. biporomum 
(Hohn & Hellerman) Lange-Bertalot, characterized by the rostrate 
apices that differ from the capitate to subcapitate apices of P. biporomum. 
Planothidium incuriatum is also misidentified as P. rostratum (Østrup) 
Round & Bukhtiyarova but it can be distinguished by its smaller 
dimensions (Wetzel et al. 2013).
Ecology: widely distributed in a broad range of environmental conditions 
(Wetzel et al. 2013). In our samples, it occurred in circumneutral pH 
(6.85), low conductivity (24 μS cm-1) and oligotrophic conditions, only 
in spring (temperature 20.8°C).
Occurrence in samples: natural substrates (UPCB 63375).



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Family DIADESMIDACEAE
Humidophila biscutella (Moser, Lange-Bertalot & Mezeltin) Lowe, 
Kociolek, Johansen, Van de Vijver, Lange-Bertalot & Kopalová
Figure: 3AK
Length: 9.5 µm; width: 2.3 µm; indistinct striae.
Valves linear with concave margins and broadly rounded apices; the 
central area is elliptical and not forming fascia. It is possible that 
Humidophila biscutella has been previously reported as Humidophila 
contenta (Grunow) Lowe, Kociolek, Johansen, Van de Vijver, Lange-
Bertalot & Kopalová. It differs from the latter species by the absence 
of wide apices and of the central area with fascia (Moser et al. 1998). 
Humidophila deceptionensis Kopalová, Zidarova & Van de Vijver is 
similar to H. biscutella, but has wider valves (2.7-3.2 µm) (Kopalová 
et al. 2015). In Brazil, it was previously registered for reservoirs in São 
Paulo state (Marquardt et al. 2018).
Ecology: the ecology of this species id little known. In our samples, it 
occurred only in temperatures < 19°C, slightly acid to circumneutral 
pH (6-6.75), low conductivity (22.5-24.5 μS cm-1) and oligotrophic 
conditions.
This is the first record for Paraná state.
Occurrence in samples: artificial (UPCB 63373; UPCB 63374) and 
natural (UPCB 63378) substrates.

Luticola permuticoides Metzeltin & Lange-Bertalot
Figure: 3AL
Length: 13.5-19.1 µm; width: 6.4-7.8 µm; striae: 19 in 10 µm.
Ecology: found in epiphytic samples collected in rivers with slightly 
acidic waters (Straube et al. 2017). In our samples, it occurred in slightly 
acid pH (6), low conductivity (24.5 μS cm-1) and oligotrophic conditions, 
only in winter (temperature 16°C).
Occurrence in samples: natural substrates (UPCB 63378).

Luticola simplex Metzeltin, Lange-Bertalot & García-Rodríguez
Figure: 3AM
Length: 12.8-16 µm; width: 4.8-5 µm; striae: 24 in 10 µm.
Ecology: previously recorded from sediment samples of subfossil 
assemblages in oligotrophic conditions and from modern assemblages 
in eutrophic conditions (Faustino et al. 2016). In our samples, it 
occurred in circumneutral pH (6.75), low conductivity (24.5 μS cm-1) 
and oligotrophic conditions, only in summer (temperature of 27°C and 
accumulated rainfall of 326.9 mm).
Occurrence in samples: natural substrates (UPCB 63376).

Family AMPHIPLEURACEAE
Frustulia australocrassinervia Casa, Mataloni & Van de Vijver
Figures: 3AN–3AP
Length: 32-43.2 µm; width: 8-9.6 µm; delicate striae, ratio 3.8-4.6.
This taxon has lanceolate valves, with slightly triundulate margins, 
protracted and rostrate apices. It differs from Frustulia crassinervia 
(Brébisson) Lange-Bertalot & Krammer by the presence of wider valves 
(11.5-12.5 µm) (Casa et al. 2018).
Ecology: previously registered for acidic waters with low conductivity 
(Casa et al. 2018). It occurred in all samples, in slightly acid to 
circumneutral pH (6-6.85), low conductivity (22.5-24.5 μS cm-1) and 
oligotrophic conditions.
This is the first record for Brazil.

Occurrence in samples: artificial (UPCB 63371; UPCB 63372; UPCB 
63373; UPCB 63374) and natural (UPCB 63375; UPCB 63376; UPCB 
63377; UPCB 63378) substrates.

Frustulia undosa Metzeltin & Lange-Bertalot
Figures: 3AQ–3AS
Length: 24.5-44.1 µm; width: 7.5-10 µm; delicate striae.
It differs from Frustulia crassinervia (Brébisson) Lange-Bertalot & 
Krammer by the presence of a narrower and longer tri-undulated valve 
(Metzeltin & Lange-Bertalot 1998).
Ecology: previously registered for a mesotrophic reservoir (Marra et 
al. 2016). In our samples, it occurred in slightly acid to circumneutral 
pH (6-6.75), low conductivity (22.5-24.5 μS cm-1) and oligotrophic 
conditions.
Occurrence in samples: artificial (UPCB 63373; UPCB 63374) and 
natural (UPCB 63376; UPCB 63377) substrates.

Family BRACHYSIRACEAE
Brachysira neoexilis Lange-Bertalot
Figures: 3AT–3AV
Length: 16-32 µm; width: 4-5.6 µm; striae: difficult to distinguish
Wolfe & Kling (2001) have suggested that Brachysira neoexilis is 
a synonym of Brachysira microcephala (Grunow) Compère. Some 
authors consider that both species differ significantly (Cantonati et al. 
2017) since the latter species presents clearly capitate apices and usually 
wider valves than B. neoexilis.
Ecology: considered as a species distributed in oligotrophic and acidic 
conditions (Lange-Bertalot & Moser 1994, Cantonati et al. 2017), and 
previously registered for mesotrophic and eutrophic reservoirs (Bertolli 
et al. 2010, Silva et al. 2010, Marra et al. 2016). It occurred in all our 
samples, in slightly acid to circumneutral pH (6-6.85), low conductivity 
(22.5-24.5 μS cm-1) and oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63371; UPCB 63372; UPCB 
63373; UPCB 63374) and natural (UPCB 63375; UPCB 63376; UPCB 
63377; UPCB 63378) substrates.

Family NEIDIACEAE
Neidium affine (Ehrenberg) Pfitzer
Figures: 3AW–3AX
Length: 42.9-46.1 µm; width: 10-10.8 µm; striae 25-26 in 10 µm.
Ecology: previously registered for oligotrophic and mesotrophic 
environments (Hofmann 1994, Marra et al. 2016). In our samples, it 
occurred in circumneutral pH (6.7), low conductivity (22.5 μS cm-1) 
and oligotrophic conditions, only in autumn (temperature of 18.6°C).
Occurrence in samples: natural substrates (UPCB 63377).

Family SELLAPHORACEAE
Sellaphora rectangularis (Gregory) Lange-Bertalot & Metzeltin
Figure: 4C
Length: 31.3-48.8 µm; width: 7.1-11.7 µm; striae: 21-27 in 10 µm.
Ecology: previously registered for oligotrophic to eutrophic 
environments (van Dam et al. 1994, Lange-Bertalot & Metzeltin 
1996). In our samples, it occurred in autumn and winter samples 
(temperature < 19°C). Thys study extends its occurrence to slightly 
acid to circumneutral pH (6-6.7), and low conductivity (22.5-24.5 μS 
cm-1) conditions.



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Periphytic diatoms from an oligotrophic reservoir

Biota Neotrop., 19(2): e20180568, 2019

http://dx.doi.org/10.1590/1676-0611-BN-2018-0568 http://www.scielo.br/bn

Figure 4. A–B. Sellaphora tridentula; C. Sellaphora rectangularis; D. Pinnularia borealis; E. Pinnularia brauniana; F. Pinnularia divergens var. divergens; 
G. Pinnularia divergens var. media; H. Pinnularia grunowii; I–J. Pinnularia obscura; K. Pinnularia pseudogibba; L–M. Hantzschia amphioxys; N. Pinnularia 
schoenfelderi; O. Kobayasiella sp.; P–Q. Capartogramma crucicula; R–T. Nitzschia semirobusta; U–V. Nitzschia acidoclinata; W. Nitzschia nana; X. Pinnularia 
viridiformis var. viridiformis; Y. Pinnularia viridiformis var. minor; Z. Pinnularia rumrichiae; AA. Pinnularia subcapitata var. elongata; AB. Pinnularia subcapitata 
var. subrostrata; AC–AD. Pinnularia subgibba var. undulata; AE. Pinnularia submicrostauron; AF. Navicula cryptocephala; AG. Navicula cryptotenella; AH. 
Navicula leptostriata; AI. Navicula veneta; AJ–AK. Chammaepinularia brasilianopsis; AL–AP. Chammaepinnularia mediocris.



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Occurrence in samples: artificial (UPCB 63373; UPCB 63374) and 
natural (UPCB 63377) substrates.

Sellaphora tridentula (Krasske) C.E.Wetzel
Figure: 4A–4B
Length: 17.5-20.8 µm; width: 4-4.6 µm; striae: difficult to distinguish.
Ecology: considered as an acidophile species (van Dam et al. 1994). In 
our samples, it occurred in spring and autumn (temperature < 21°C). 
This study extends the occurrence of this species to circumneutral 
pH (6.7-6.85), low conductivity (22.5-24 μS cm-1) and oligotrophic 
conditions.
Occurrence in samples: natural substrates (UPCB 63375; UPCB 
63377).

Family PINNULARIACEAE
Pinnularia borealis Ehrenberg var. borealis.
Figure: 4D
Length: 31-31.2 µm; width: 7-7.2 µm; striae: 7 in 10 µm.
Ecology: considered as indifferent to environmental conditions 
(Hofmann 1994) P. borealis is common in oligo and mesotrophic 
environments (van Dam et al. 1994, Bertolii et al. 2010), while rarely 
registered for eutrophic conditions (Silva et al. 2010). In our samples, it 
occurred in temperature s < 21°C, slightly acid to circumneutral pH (6-
6.85), low conductivity (24-24.5 μS cm-1) and in oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63374) and natural (UPCB 
63375) substrates.

Pinnularia brauniana (Grunow) Studnicka.
Figure: 4E
Length: 52.2-60 µm; width: 8.5-9.6 µm; striae: 11-12 in 10 µm.
Ecology: previously registered for oligotrophic, mesotrophic and 
eutrophic conditions (Van Dam et al. 1994, Silva et al. 2010, Marra 
et al. 2016). According to Krammer (1992), P. brauniana is not 
considered commonly found in waters with low conductivity and pH 
< 6. This study extends its occurrence to temperatures < 21°C, acid to 
circumneutral pH (6-6.85), low conductivity (22.5-24.5 μS cm-1), and 
oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63371; UPCB 63373; 
UPCB 63374) and natural (UPCB 63375; UPCB 63377; UPCB 63378) 
substrates.

Pinnularia divergens Smith var. divergens
Figure: 4F
Length: 74.5 µm; width: 12.5 µm; striae: 12 in 10 µm.
Ecology: in agreement with previously studies (van Dam et al. 1994, 
Cantonati et al. 2017), we found this species in pH < 7, low conductivity 
(22.5 μS cm-1) and oligotrophic conditions only in autumn (temperature 
18.6°C).
Occurrence in samples: natural substrates (UPCB 63377).

Pinnularia divergens var. media Krammer
Figure: 4G
Length: 39-54.5 µm; width: 7.7-9.5 µm; striae: 14-15 in 10 µm.
Ecology: according to previous studies, this species prefers oligotrophic 
environments (van Dam et al. 1994, Cantonati et al. 2017) but can also 
occur in eutrophic waters (Silva et al. 2010). In our samples, it occurred 

in in temperatures < 21°C, slightly acid to circumneutral pH (6-6.85), 
low conductivity (24-24.5 μS cm-1) and in oligotrophic conditions.
Occurrence in samples: natural substrates (UPCB 63375; UPCB 
63377; UPCB 63378).

Pinnularia grunowii Krammer
Figure: 4H
Length: 31-41 µm; width: 6-7.5 µm; striae: 14 in 10 µm.
Ecology: previously registered for eutrophic environments, with low 
to moderate electrolytes and pH > 8 (Krammer 2000) and also for 
mesotrophic waters (Bertolli et al. 2010). In our samples, it occurred in 
circumneutral pH (6.7-6.85) and low conductivity (22.5-24.5 μS cm-1), 
and oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63373) and natural (UPCB 
63375; UPCB 63376) substrates.

Pinnularia obscura Krasske
Figures: 4I–4J
Length: 20.5-21.5 µm; width: 4-4.5 µm; striae: 16 in 10 µm.
Ecology: considered as a species tolerant to environments with pH < 
4 (Ciniglia et al. 2007). In our samples, it occurred in circumneutral 
pH (6.7-6.85), low conductivity (22.5-24 μS cm-1), and oligotrophic 
conditions, in spring and autumn (temperature < 21°C).
Occurrence in samples: natural substrates (UPCB 63375; UPCB 
63377).

Pinnularia pseudogibba Krammer
Figure: 4K
Length: 54 µm; width: 9 µm; striae: 11 in 10 µm.
Pinnularia pseudogibba differs from Pinnularia subgibba Krammer 
var. subgibba by the presence of lanceolate valves with subrostrate 
apices (Krammer 2000).
Ecology: previously registered for oligotrophic environments (Krammer 
2000). In our samples, it occurred in slightly acid to circumneutral pH (6-
6.85), low conductivity (24-24.5 μS cm-1) and oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63371; UPCB 63374) and 
natural (UPCB 63375; UPCB 63376; UPCB 63377) substrates.

Pinnularia rumrichae Krammer.
Figure: 4Z
Length: 45-46.5 µm; width: 7.5-8 µm; striae: 13 in 10 µm.
Ecology: in our samples, it occurred in slightly acidic to circumneutral 
pH (6-6.85), low conductivity (24-24.5 μS cm-1) and in oligotrophic 
conditions.
Occurrence in samples: artificial (UPCB 63373) and natural (UPCB 
63375; UPCB 63376; UPCB 63378) substrates.

Pinnularia schoenfelderi Krammer.
Figure: 4N
Length: 23 µm; width: 4.8 µm; striae: 16 in 10 µm.
Ecology: previously registered for oligotrophic environments with low 
to moderate conductivity (Krammer 2000), and also for mesotrophic 
waters (Bertollit et al. 2010). In our samples, it occurred in circumneutral 
pH (6.7), low conductivity (22.5 μS cm-1) and oligotrophic conditions, 
only in autumn (temperature 18.6°C).
Occurrence in samples: natural substrates (UPCB 63377).



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Periphytic diatoms from an oligotrophic reservoir

Biota Neotrop., 19(2): e20180568, 2019

http://dx.doi.org/10.1590/1676-0611-BN-2018-0568 http://www.scielo.br/bn

Pinnularia subcapitata var. elongata Krammer.
Figure: 4AA
Length: 43.5 µm; width: 6 µm; striae: 16 in 10 µm.
Ecology: commonly associated with sphagnum moss (Krammer 1992). 
In our samples, it occurred in slightly acid pH (6), low conductivity (24.5 
μS cm-1) and oligotrophic conditions, only in winter (temperature 16°C).
Occurrence in samples: artificial substrates (UPCB 63374).

Pinnularia subcapitata var. subrostrata Krammer
Figure: 4AB
Length: 38.5-46.5 µm; width: 6-6.5 µm; striae: 13 in 10 µm.
Ecology: widespread species; common in waters with medium 
electrolyte concentrations (Krammer 1992). In our samples, it occurred 
in circumneutral pH (6.7-6.85), low conductivity (22.5-24.5 μS cm-1) 
and oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63373) and natural (UPCB 
63375; UPCB 63376; UPCB 63377) substrates.

Pinnularia subgibba var. undulata Krammer
Figures: 4AC–4AD
Length: 65.1-68.5 µm; width: 7-7.8 µm; striae: 11-12 in 10 µm.
Ecology: previously registered for waters with neutral pH and high 
conductivity (Noga et al. 2014). In Brazil, it was recorded from 
planktonic samples in Guaraguaçu River (Tremarin et al. 2010) and 
from epiphytic samples in a mesotrophic reservoir of the South region 
(Marra et al. 2016). This study expanded its occurrence to circumneutral 
pH (6.7-6.85), low conductivity (22.5-24.5 μS cm-1) and oligotrophic 
conditions.
Occurrence in samples: artificial (UPCB 63373) and natural (UPCB 
63375; UPCB 63376; UPCB 63377) substrates.

Pinnularia submicrostauron Schroeter
Figure: 4AE
Length: 38.4-43.7 µm; width: 5-6.6 µm; striae: 12-13 in 10 µm.
Ecology: previously registered as epiphytic in non-polluted 
environments, with acid pH (3.95-5.89) and medium to high 
conductivity (46-94 μS cm-1) (Krammer 1992, Veselá 2009, Szulc et 
al. 2014). In our samples, it occurred in circumneutral pH (6.7), low 
conductivity (22.5-24.5 μS cm-1) and oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63373) and natural (UPCB 
63376; UPCB 63377) substrates.

Pinnularia viridiformis var. viridiformis Krammer
Figure: 4X
Length: 131 µm; width: 17.3 µm; striae: 10 in 10 µm.
Ecology: this species is frequently sampled especially in oligotrophic 
and mesotrophic environments, with low to moderate conductivity 
(Krammer 2000, Bertolli et al. 2010), being more abundant in acid 
pH (Van de Vijver & Beyen 1997, Veselá 2014). In our study, it was 
only recorded from samples of periphyton associated with aquatic 
macrophytes (75%), in slightly acid to circumneutral pH (6-6.85), low 
conductivity (24-24.5 μS cm-1) and oligotrophic conditions.
Occurrence in samples: natural substrates (UPCB 63375; UPCB 
63376; UPCB 63378).

Pinnularia viridiformis var. minor Krammer.
Figure: 4Y
Length: 52-82.5 µm; width: 11-12.5 µm; striae: 10-12 in 10 µm.
Pinnularia viridiformis var. minor differs from Pinnularia viridis 
(Nitzsch) Ehrenberg by the valve outline and by the presence of a semi 
complex raphe (Krammer 2000).
Ecology: considered as cosmopolitan (Krammer 2000); it is rare in 
waters with neutral to alkaline pH and in moderate to high conductivity 
(Cremer et al. 2004, Noga et al. 2014). It was previously registered for 
streams with acid pH and low conductivity, in Central Brazil (França 
et al. 2017). In our study, it was recorded from samples of periphyton 
associated with aquatic macrophytes, in temperatures of 16°C, acid 
pH (6), low conductivity (24.5 μS cm-1), and oligotrophic conditions.
This is the first record for Paraná state.
Occurrence in samples: natural substrates (UPCB 63378).

Order NAVICULALES
Family incertae sedis
Chamaepinnularia brasilianopsis Metzeltin & Lange-Bertalot
Figures: 4AJ–4AK
Length: 13.5-13.9 µm; width: 3.5-4.1 µm; striae: 24 in 10 µm.
Ecology: previously described for the Neotropical region (Metzeltin & 
Lange-Bertalot 1998), in mesotrophic waters (Marra et al. 2016). with 
circumneutral to slightly alkaline pH, and conductivity between 24.3 
and 145 µS cm-1 (Sala et al. 2015). In our study, it was only registered 
in autumn, from samples of macrophytes, in temperatures of 18.6°C, 
circumneutral pH (6.7), low conductivity (22.5 μS cm-1) and oligotrophic 
conditions.
Occurrence in samples: natural substrates (UPCB 63377).

Chamaepinnularia mediocris (Krasske) Lange-Bertalot & Krammer
Figures: 4AL–4AP
Length: 10-14.7 µm; width: 2.5-3 µm; striae: 21-24in 10 µm.
Ecology: considered as characteristic of oligotrophic environments 
(van Dam et al. 1994, Hofmann 1994), but also cited for mesotrophic 
and eutrophic waters (Silva et al. 2010, Marra et al. 2016). It is a good 
indicator of environments rich in minerals, with moderate to high 
conductivity, and acid pH (Fránková et al. 2009), and also associated 
with the process of deglaciation (Fell et al. 2018). In our study, it 
occurred in summer (temperature of 27°C and high acummulated 
rainfall), in waters with slightly acid to circumneutral pH (6-6.75), low 
conductivity (22.5-24.5 μS cm-1), and oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63372) and natural (UPCB 
63377; UPCB 63378) substrates.

Kobayasiella sp.
Figure: 4O
Length: 39.8 µm; width: 7.4 µm; striae: 15 in 10 µm.
Kobayasiella sp. presents linear-elliptic valves with subcapitate ends. 
The axial area is linear and straight. The central area is elliptic and 
delimited by irregular striae that are radiate towards the valve center.
The valve outline is similar to Kobayasiella acidophila Vanhoutte, 
Kobayasiella subtilissima (Cleve) Lange-Bertalot and Kobayasiella 
tapajoensis Metzeltin & Lange-Bertalot, but these three latter taxa 



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present smaller size (24.7-31.2 µm, 23-27 µm, and 23-34 µm, 
respectively) and a higher striae density (34-42, 36-40 and 34-38 in 
10 µm, respectively). Kobayasiella sp. is also similar to Kobayasiella 
floridana Metzeltin & Lange-Bertalot and Kobayasiella krasskei 
(Metzeltin & Lange-Bertalot) Lange-Bertalot, but these species have 
larger valves (54-83 µm and 44-50 µm, respectively) and higher striae 
density (28-32 and 27-28 in 10 µm, respectively) (Metzeltin & Lange-
Bertalot 2007, 1998, Vanhoutte et al. 2004).
Occurrence in samples: natural substrates (UPCB 63377).

Family NAVICULACEAE
Navicula cryptocephala Kützing
Figure: 4AF
Length: 23.2-39.2 µm; width: 4.8-6 µm; striae: 12-18 in 10 µm.
Ecology: cosmopolitan; registered for oligotrophic to eutrophic 
environments, but more abundant in mesotrophic waters (Bertolli et al. 
2010, Bartozek et al. 2018). In our study, it occurred in slightly acid to 
circumneutral pH (6-6.85), low conductivity (22.5-24.5 μS cm-1), and 
oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63371; UPCB 63372; 
UPCB 63373) and natural (UPCB 63375; UPCB 63376; UPCB 63378) 
substrates.

Navicula cryptotenella Lange-Bertalot.
Figure: 4AG
Length: 20-35 µm; width: 4.5-5.6 µm; striae: 12-18 in 10 µm.
Ecology: widely distributed in oligotrophic to eutrophic conditions 
(Bertolii et al. 2010, Silva et al. 2010, Marra et al. 2016, Bartozek et 
al. 2018, Santos & Ferragut 2018), but most commonly associated 
with low TP concentrations (Wang et al. 2019). It occurred in all of our 
samples, in slightly acid to circumneutral pH (6-6.85), low conductivity 
(22.5-24.5 μS cm-1), and oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63371; UPCB 63372; UPCB 
63373; UPCB 63374) and natural (UPCB 63375; UPCB 63376; UPCB 
63377; UPCB 63378) substrates.

Navicula leptostriata Jørgensen
Figure: 4AH
Length: 33-36.3 µm; width: 4.5-4.8 µm; striae: 17-18 in 10 µm.
Ecology: previously registered for environments with moderate to high 
nutrient concentrations (van Dam et al. 1994, Silva et al. 2010, Ma et 
al. 2014). In our study, it occurred in temperatures < 21°C, slightly acid 
and circumneutral pH (6-6.85), low conductivity (22.5-24.5 μS cm-1), 
and oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63374) and natural (UPCB 
63375; UPCB 63377) substrates.

Navicula veneta Kützing
Figure: 4AI
Length: 17 µm; width: 4 µm; striae: 18 in 10 µm.
Ecology: commonly registered for environments rich in nutrients (van 
Dam et al. 1994, Hofmann 1994, Silva et al. 2010, Marra et al. 2016, Casco 
et al. 2018). In our samples, it only occurred in autumn (temperature of 
18.6°C). This study expanded its occurrence to circumneutral pH (6.7), 
low conductivity (22.5 μS cm-1) and oligotrophic conditions.
Occurrence in samples: natural substrates (UPCB 63377).

Capartogramma crucicula (Grunow ex Cleve) Ross
Figures: 4P–4Q
Length: 27.5 µm; width: 8.5 µm; striae: 21 in 10 µm.
Ecology: the ecological optimum of this species is difficult to define 
(Beyene et al. 2014). It was previously registered for the oligotrophic 
paleophase (Faustino et al. 2016) and for modern mesotrophic conditions 
(Marra et al. 2016). In our study, it occurred in circumneutral pH (6.85), 
low conductivity (24 μS cm-1) and oligotrophic conditions, only in spring 
(temperature of 20.8°C).
Occurrence in samples: artificial substrates (UPCB 63771).

Family STAURONEIDACEAE
Stauroneis neohyalina Lange-Bertalot & Krammer
Figures: 5A–5B
Length: 43-52 µm; width: 8-8.7 µm; striae: difficult to distinguish 
under the LM.
Stauroneis neohyalina differs from Stauroneis gracilior (Rabenhorst) 
Reichardt by the presence of subcapitate apices, narrower valves and 
a higher striae density. The striae are difficult to distinguish under the 
light microscope and the areolae are inconspicuous (Reichardt 1995, 
Lange-Bertalot & Metzeltin 1996). In this species, the proximal ends 
of the raphe are straight and not inflated and teardrop-shaped as in S. 
gracilior (Cantonati et al. 2017).
Widely recorded in different worldwide regions (Guiry & Guiry 2018).
This is the first record for Brazil.
Ecology: previously registered for oligotrophic and acidic environments 
(van Dam et al. 1994, Sienkiewicz 2005, Van de Vijver et al. 2005, Krstić 
et al. 2012). In our samples, it occurred in slightly acid to circumneutral 
pH (6-6.75), low conductivity (22.5-24.5 μS cm-1) and oligotrophic 
conditions, in summer (temperature of 27°C).
Occurrence in samples: artificial (UPCB 63373) and natural (UPCB 
63376; UPCB 63378) substrates.

Stauroneis gracilis Ehrenberg
Figure: 5C
Length: 83.5 µm; width: 13.5 µm; striae: 23 in 10 µm.
Ecology: considered as characteristic of oligotrophic environments 
(Dixit & Smol 1995); it is rare in mesotrophic and eutrophic conditions 
(Silva et al. 2010, Marra et al. 2016) and has its optimum in conditions 
of low conductivity (Pestryakova et al. 2018). In our study, it only 
occurred from samples of aquatic macrophytes, in circumneutral pH 
(6.7), low conductivity (22.5 μS cm-1) and oligotrophic conditions, in 
autumn (temperature of 18.6°C).
Occurrence in samples: natural substrates (UPCB 63377).

Stauroneis phoenicenteron (Nitzsch) Ehrenberg
Figure: 5D
Length: 96.5-116.5 µm; width: 18.2-22.5 µm; striae: 18-19 in 10 µm.
Ecology: considered as characteristic of meso-eutrophic environments 
(van Dam et al. 1994, Silva et al. 2010, Bertolii et al. 2010), it has its 
optimum in conditions of intermediate conductivity (Pestryakova et al. 
2018) and acid to circumneutral pH (Ma et al. 2014). In this samples, it 
occurred in slightly to circumneutral pH (6-6.7), low conductivity (22.5-
24.5 μS cm-1) and oligotrophic conditions, in autumn (temperature < 21°C).
Occurrence in samples: artificial (UPCB 63373; UPCB 63374) and 
natural (UPCB 63377) substrates.



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Figure 5. A–B. Stauroneis neohyalina; C. Stauroneis gracilis; D. Stauroneis phoenicenteron; E. Craticula acidoclinata; F. Craticula riparia var. mollenhaurei; 
G–H. Craticula submolesta; I. Surirella grunowii; J. Iconella curvula; K. Rhopalodia gibberula var. vanheurckii.

Craticula acidoclinata Lange-Bertalot & Metzeltin
Figure: 5E
Length: 65-1-74.2 µm; width: 17.9-18.5 µm; striae: 15 in 10 µm; 
areolae: 24 in 10 µm.
Craticula acidoclinata differs from Craticula ambigua (Ehrenberg) 
Mann by the presence of subrostrate apices and broadly rounded 
ends in the valves (distinctly protracted and rostrate in C. ambigua). 
Craticula cuspidata (Kützing) Mann has wider valves with narrower 
apices (Lange-Bertalot & Metzeltin 1996).
Ecology: registered for acid environments, poor in nutrients (Cantonati 
et al. 2017). It was previously found in Rio Tapajós, Brazil (Metzeltin 
& Lange-Bertalot 1998). In our samples, it occurred in circumneutral 
pH (6.7), low conductivity (22.5 μS cm-1) and oligotrophic conditions, 
in autumn (temperature < 21°C).
This is the first record for Paraná state.

Occurrence in samples: natural substrates (UPCB 63377).

Craticula riparia var. mollenhaurei Lange-Bertalot.
Figure: 5F
Length: 33.5 µm; width: 7 µm; striae: 24 in 10 µm.
Craticula riparia var. mollenhaurei resembles Craticula riparia 
(Hustedt) Lange-Bertalot var. riparia Lange-Bertalot in the valve outline 
but differs from the latter by the presence of longer and wider valves 
(35-50 µm and 8-10.5 µm) and a lower striae density (15-18 in 10 µm). 
Craticula riparia var. riparia is common in eutrophic environments 
with high conductivity (Lange-Bertalot 2001, van Dam et al. 1994).
Ecology: previously registered for slightly acid to neutral waters, with 
low conductivity (Lange-Bertalot 2001, Lange-Bertalot 1993). In our 
samples, it occurred in circumneutral pH (6.7), low conductivity (22.5 
μS cm-1) and oligotrophic conditions, in.autumn (temperature < 21°C).



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This is the first record for Brazil.
Occurrence in samples: natural substrates (UPCB 63377).

Craticula submolesta (Hustedt) Lange-Bertalot
Figures: 5G–5H
Length: 16.8-17.1 µm; width: 4-4.3 µm; striae: 22-24 in 10 µm.
Ecology: considered as characteristic of oligotrophic environments 
(Lange-Bertalot & Metzeltin 1996), it is rarely found in mesotrophic 
conditions (Marra et al. 2016). In our samples, it occurred in 
circumneutral pH (6.7), low conductivity (22.5 μS cm-1) and oligotrophic 
conditions, in autumn (temperature < 21°C).
Occurrence in samples: natural substrates (UPCB 63377).

Family BACILLARIACEAE
Nitzschia acidoclinata Lange-Bertalot
Figure: 4U–4V
Length: 21-38.7 µm; width: 2.5-3 µm; indistinct striae; fibulae: 13-14 
in 10 µm.
Ecology: previously registered for oligo-mesotrophic environments 
(van Dam et al. 1994, Hofmann 1994). In our samples, it occurred in 
temperatures < 21°C, slightly acid to circumneutral pH (6-6.85), low 
conductivity (22.5-24.5 μS cm-1) and oligotrophic conditions.
Occurrence in samples: artificial (UPCB 63373; UPCB 63374) and 
natural (UPCB 63375; UPCB 63378) substrates.

Nitzschia nana Grunow
Figure: 4W
Length: 58.3, width: 5.2 µm; indistinct striae; fibulae: 10 in 10 µm.
Ecology: it avoids saprobic environments (Cantonati et al. 2017). In our 
samples, it occurred in circumneutral pH (6.85), low conductivity (24 
μS cm-1) and oligotrophic conditions, in spring (temperature < 21°C).
Occurrence in samples: natural substrates (UPCB 63375)

Nitzschia semirobusta Lange-Bertalot.
Figures: 4R–4T
Length: 16-20 µm; width: 4.5 -5 µm; striae: 16-18 in 10 µm; fibulae: 
7 in 10 µm.
Nitzschia semirobusta differs from Nitzschia amphibia Grunow by the 
presence of longer fibulae and from Nitzschia amphibioides Hustedt 
by having a greater striae density. Denticula kuetzingii Grunow is 
distinguished from Nitzschia semirobusta by the presence of fibulae 
that are extended completely and with similar thickness from margin 
to margin in the first species (Lange-Bertalot 1993), while in the latter 
species the fibulae decrease in thickness towards the margin.
Ecology: Due to the taxonomic problems and misidentification, 
the ecological preference of N. semirobusta is unclear. Therefore, 
information on its occurrence and ecological preferences is of high 
relevance for delimiting the ecological requirements and tolerances of 
N. semirobusta. It was frequently registered for oligo- to mesotrophic 
reservoirs in São Paulo state (Bartozek et al. 2018). In our samples, it 
occurred in slightly acid to circumneutral pH (6-6.85), low conductivity 
(22.5-24.5 μS cm-1) and oligotrophic conditions.
This is the first record for Paraná state.
Occurrence in samples: artificial (UPCB 63371; UPCB 63374) and 
natural (UPCB 63375; UPCB 63376; UPCB 63377; UPCB 63378) 
substrates.

Hantzschia amphioxys (Ehrenberg) Grunow
Figure: 4L–4M
Length: 23.5-45 µm; width: 5-5.6 µm; indistinct striae; fibulae: 8-9 
in 10 µm.
Ecology: common in soil and aerial habitats (Jahn et al. 2014), and 
considered as tipically indifferent to trophic conditions (van Dam et 
al. 1994, Hofmann 1994). It is rarely found in waters poor in nutrients, 
being more common in mesotrophic environments (Bertolli et al. 
2010, Bartozek et al. 2018, Cuña-Rodríguez et al. 2018) and eutrophic 
conditions (Silva et al. 2010). Previous studies associated this species 
with cold waters (Darling et al. 2017), however, in our samples it only 
occurred in summer (temperature 27°C), in circumneutral pH (6.75), 
low conductivity (24.5 μS cm-1) and oligotrophic conditions.
Occurrence in samples: natural substrates (UPCB 63376).

Family RHOPALODIACEAE
Rhopalodia gibberula var. vanheurckii O. Müller
Figure: 5K
Length: 35.1-36.3 µm; width: 8-8.5 µm; striae: 14-19 in 10 µm; 
transapical ribs: 4-5 in 10 µm.
Ecology: considered as an alkalinophilous species (van Dam et 
al. 1994). In our samples, it occurred in slightly acid pH (6), low 
conductivity (24.5 μS cm-1) and oligotrophic conditions, only in winter 
(temperature 16°C),
Occurrence in samples: natural substrates (UPCB 63378).

Family SURIRELLACEAE
Surirella grunowii Kulikovskiy, Lange-Bertalot & Witkovski.
Figure: 5I
Length: 62.4-76.1 µm; width: 11.5-11.6 µm; striae: difficult to 
distinguish; alar canals: 3-4 in 10 µm.
Surirella linearis var. constricta Grunow was synonymized with S. 
grunowii by Kulikovskiy et al. (2010).
Ecology: previously registered for oligo-mesotrophic environments 
(van Dam et al. 1994). In our samples, it occurred in circumneutral 
pH (6.7-6.85), low conductivity (22.5-24 μS cm-1) and oligotrophic 
conditions, in spring and autumn seasons (temperature < 21°C).
Occurrence in samples: artificial (UPCB 63371) and natural (UPCB 
63377) substrates.

Iconella curvula (Smith) Ruck & Nakov
Figure: 5J
Length: 92 µm; width: 4 µm; indistinct striae; alar canals: 7 in 10 µm.
Ecology: cited as Stenopterobia curvula (Smith) Krammer, it was 
registered for oligotrophic (van Dam et al. 1994) and mesotrophic 
environments (Marra et al. 2016). In our samples, it occurred in 
circumneutral pH (6.75), low conductivity (24.5 μS cm-1) and 
oligotrophic conditions, only in summer (temperature 27°C).
Occurrence in samples: natural substrates (UPCB 63376).

Conclusions

The inventory performed at the Piraquara I reservoir brings an 
important contribution to the diatom diversity from oligotrophic 
environments. We registered the first occurrence of five diatom species 
in Brazil and seven in the state of Paraná, contributing to the geographic 



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Periphytic diatoms from an oligotrophic reservoir

Biota Neotrop., 19(2): e20180568, 2019

http://dx.doi.org/10.1590/1676-0611-BN-2018-0568 http://www.scielo.br/bn

distribution and the ecology of the identified species. Finally, fourteen 
periphytic species extended their ecological limits of pH, conductivity 
and trophic preferences.

Acknowledgments

To “Conselho Nacional de Desenvolvimento Científico e 
Tecnológico” (CNPq) for the master´s scholarship (AMSL), scientific 
productivity scholarship (TAVL) and financial support for the project 
(CTHidro-555397/2006-8). To IAP and SANEPAR for providing 
chemical data and field sampling support and to Laboratório de Ficologia 
UFPR for facilitating the use of the optical microscope.

Author Contributions

Angela Maria da Silva-Lehmkuhl: has substantial contribution in 
the concept and design of the study; data collection; data analysis and 
interpretation; manuscript preparation and to critical revision, adding 
intelectual content.

Priscila Izabel Tremarin: Has substantial contribution to data 
analysis and interpretation, and to critical revision, adding intelectual 
content.

Ilka Schincariol Vercellino: Has substantial contribution to data 
analysis and interpretation, and to critical revision, adding intelectual 
content.

Thelma A. Veiga Ludwig: has substantial contribution in the 
concept and design of the study; data collection; data analysis and 
interpretation; manuscript preparation and to critical revision, adding 
intelectual content.

Conflicts of Interest

The authors declare that they have no conflict of interest related to 
the publication of this manuscript.

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