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Ramos et al 2018 - Some new, rare and interesting desmids from bromeliad phytotelmata in Brazil

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Phytotaxa 346 (1): 059–077
http://www.mapress.com/j/pt/
Copyright © 2018 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Thomas Pröschold: 8 Mar. 2018; published: 29 Mar. 2018
https://doi.org/10.11646/phytotaxa.346.1.3
59
Some new, rare and interesting desmids from bromeliad phytotelmata in Brazil
GERALDO JOSÉ PEIXOTO RAMOS1, CARLOS EDUARDO DE MATTOS BICUDO2 & CARLOS WALLACE 
DO NASCIMENTO MOURA1
1Programa de Pós-Graduação em Botânica, Universidade Estadual de Feira de Santana, Av. Transnordestina, s/nº, Novo Horizonte, 
44036-900, Feira de Santana, BA, Brazil
2Núcleo de Ecologia, Instituto de Botânica, Av. Miguel Estéfano, nº 3687, 04301-902 São Paulo, SP, Brazil
E-mail: geraldojpr@gmail.com (corresponding author)
Abstract
Studies of the microalgae in bromeliad phytotelmata in northeastern Brazil revealed the occurrence of some interesting 
desmids—including new and rare taxa, besides new records for South America. Additionally, we reconsidered some taxa 
previously recorded in the study area, providing new information and ecological data, as well as SEM images. A total of 
19 desmids were identified, 18 from Serra da Jiboia (granite outcrops) and one from Itaberaba (in the “Caatinga” dryland 
domain). Taxonomic notes and ecological characteristics of the studied taxa are included, as well as a checklist of all of the 
desmids recorded in Brazilian bromeliad phytotelmata. 
Key words: algae, Bromeliaceae, Desmidiaceae, Mesotaeniaceae, taxonomy, Zygnematophyceae
Introduction
Desmids comprise a diverse group of freshwater microalgae generally found in shallow, nutrient-poor, acidic to slightly 
alkaline aquatic environments (Brook 1981, Coesel & Meesters 2007). Those conditions are typical of bromeliad 
phytotelmata, making them ideal environments for the development of those algae. Recent taxonomic studies, based 
mainly on light and scanning electron microscopy, continue to reveal new and remarkable desmid species in Europe, 
Africa, and South America (Šťastný 2010; Šťastný & Kouwets 2012; Van Geest & Coesel 2012; Coesel & Van Geest 
2014, 2016; Coesel et al. 2017), even though phytotelmata environments are often overlooked by desmidiologists.
 One of the first reports of the occurrence of microalgae in bromeliad tanks in Brazil was published early in the 
last century, recording two taxa (Cosmarium sp. and Pleurotaenium sp.) in the tanks of Nidularium sp. in Cabo Frio, 
Rio de Janeiro State (Kolkwitz 1933). Later, in the 1970s, Lyra (1976) listed four desmids (Micrasterias sp, Euastrum 
sp., Closterium sp., and Cosmarium sp.) during a survey of microalgae (especially diatoms) in bromeliad tanks in 
Pernambuco State in northeastern Brazil. None of those studies, however, were focused specifically on desmids, and 
only in the last two decades those algae have become subject to specific research approaches, as in the studies of 
Sophia (1999) and Sophia et al. (2004), who carried out floristic and ecological surveys of bromeliad tanks in Rio de 
Janeiro State.
 Studies of desmids from bromeliad phytotelmata in Bahia State are even more recent, with the first survey (Ramos 
et al. 2011) at Serra da Jiboia. Since then, additional taxonomic studies have been undertaken in that area (and elsewhere 
in the state), producing records of rare and new desmids species (Ramos et al. 2017a, b). 
 The present study expanded our knowledge of desmids from phytotelmata in two areas in Bahia State, identifying 
some new and rare taxa for Brazil and South America. Additionally, other species previously recorded for those 
microhabitats were re-evaluated and new information added—with new identifications, SEM images, and refined 
ecological data. We provide here an updated checklist of the desmids recorded in Brazilian bromeliad phytotelmata.
RAMOS ET Al.60 • Phytotaxa 346 (1) © 2018 Magnolia Press
Material and Methods
Study area. The present study was undertaken in two areas in Bahia State, Brazil: Monte da Pioneira at Serra da Jiboia 
(12°51’ S, 39° 28’ W), an area of granite rock outcrops in the municipality of Santa Teresinha; and Fazenda Itaberaba, 
in the municipality of Itaberaba (12°30’ S, 40°04’ W), located in semiarid “ Caatinga” vegetation.
 Sampling and analyses. We examined material from 81 samples (out of a total of 160 collected; 80 in each 
area) from the tanks of two bromeliad species: Aechmea cf. lingulatoides Leme & H.E.Luther (1998: 187) (Fazenda 
Itaberaba), and Alcantarea nahoumii (Leme) J.R.Grant (1995: 91) (Serra da Jiboia). The bromeliad tanks were sampled 
with the aid of a syringe (50 mL) and a polyethylene hose. During the collections, abiotic variables of water such as 
pH, water temperature (T), conductivity (C), and total dissolved solids (TDS) were measured using a Hanna HI98130 
portable probe, and dissolved oxygen (DO) was determined using an Instrutherm device (MO-910).
 All of the materials were examined under an Olympus LX35 Optical Microscope and photographed using a 
MicroPublisher—QImaging MP5.0-RTV-CLR-10-C digital camera (Figs 1–112). After analysis, the samples were 
preserved in Transeau’s solution, following Bicudo & Menezes (2017), and subsequently incorporated into the liquid 
collection of the State University of Feira de Santana herbarium (HUEFS).
 A table (Table 1) demonstrating the frequencies of occurrence of the taxa and water acidity is provided. Five 
frequency categories were used: (1) very frequent—F ≥ 75%; (2) frequent—50% ≤ F ≤ 75%; (3) occasional—15% ≤ 
F ≤ 50%; (4) rare—5% ≤ F ≤ 15%; and, (5) very rare—F ≤ 5%. Percentage refers to total of bromeliads sampled in 
each area which the desmid species occurred. The following categories were defined in terms of acidity, following 
Šťastný (2010): acidophilous (occurring at pH 7.5). As an 
overview of all the desmids known to occur in bromeliad phytotelmata in Brazil, a second table (Table 2) is provided 
with information concerning the vegetation formation in which the bromeliads occur, the bromeliad genera, and the 
locality (by Brazilian state). Species previously recorded for Serra da Jiboia (Ramos et al. 2011) are designated by (*), 
but additional information was added, such as optical photographs with other taxonomically useful views, scanning 
electron microscopy (SEM) details, and ecological data. New identifications were made for some taxa, with additional 
remarks. 
 The preparation of the material for SEM examination followed the protocols of Ramos et al. (2017a).
Results & Discussion
Actinotaenium mooreanum (W.Archer) Teiling (1954: 408) (Figs 68–72)
Cells solitary, approximately 2 times longer than wide, widely elliptic, apices widely rounded; chloroplast stellate, two 
pyrenoids; zygospore subrectangular, dark brown, with rounded-acuminate papillae. Cells 16–20 μm long, 8–9 μm 
wide; zygospore 17–22 μm long, 11–14 μm wide.
 Material examined: BRAZIL, Bahia, Santa Teresinha, Serra da Jiboia. 14-I-2015, G.J.P. Ramos et al. s/n (HUEFS 
224661), 18-VII-2015, G.J.P. Ramos et al. s/n (HUEFS 224679), 11-IX-2015, G.J.P. Ramos et al. s/n (HUEFS 224705), 
20-XI-2015, G.J.P. Ramos et al. s/n (HUEFS 224721).
 Habitat: bromeliad tanks (Alcantarea nahoumii); T 26.1 (± 3.9 ºC). pH 5.9 (± 0.6). C 0.24 (± 0.18 mS.cm-1). TDS 
0.11 (± 0.09 ppt). DO 6.4 (± 0.8 mg.L-1).
 Notes: Actinotaenium mooreanum is reported here for the first time for South America; with previous records 
only from Europe and Oceania (Guiry & Guiry 2017). During our analysis, we commonly found zygospores of this 
species—usually in groups associated with the mucilaginous sheaths of cyanobacteria. According to Růžička (1981), 
this species prefers slightly acidic to neutral environments (pH 4.9–7.5), similar to conditions found in bromeliad 
tanks.
Closterium cornu var. minus Irénée-Marie (1952: 7) (Figs 1–2)
Cells semilunate, 9 to 11 times longer than wide, slightly curved, not inflated in the medianregion; poles acuminate-
truncate; cell walls smooth; chloroplast axial, pyrenoids 2 to 4 in series. Cell dimensions: 44–59 µm long, 4–6 µm 
wide, 2–3 µm apices. 
BROMELIAD PHyTOTELMATA IN BRAZIL Phytotaxa 346 (1) © 2018 Magnolia Press • 61
 Material examined: BRAZIL, Bahia, Santa Teresinha, Serra da Jiboia. 14-I-2015, G.J.P. Ramos et al. s/n (HUEFS 
224653), 18-VII-2015, G.J.P. Ramos et al. s/n (HUEFS 224680), 11-IX-2015, G.J.P. Ramos et al. s/n (HUEFS 224697), 
20-XI-2015, G.J.P. Ramos et al. s/n (HUEFS 224710)
 Habitat: bromeliad tanks (Alcantarea nahoumii); water temperature 26.5 (± 2.8 ºC). pH 5.9 (± 0.6). conductivity 
0.25 (± 0.19 mS.cm-1). total dissolved solids 0.13 (± 0.09 ppt). dissolved oxygen 6.5 (± 2.3 mg.L-1).
 Notes: Closterium cornu var. minus is a rare taxon, with only one record for the type locality (Québec, Canada) 
(Irénée-Marie 1952). The variety identified here differs by having smaller cell dimensions (up to 60 μm) than those 
generally reported for the typical variety (95–190 μm) (Prescott et al. 1975). All specimens from the phytotelmata had 
cell dimensions similar to C. cornu var. minus. This variety was considered as occasional in the bromeliads from Serra 
da Jiboia (Table 1).
Cosmarium amoenum var. jiboensis G.J.P.Ramos, C.E.M.Bicudo & C.W.N.Moura var. nov. (Figs 3–8)
Cells 1.6–1.7 times longer than wide, deep median constriction with a linear, closed sinus, oblong in apical view, 
elliptic-oblong in lateral view; semicells semicircular, with rounded basal angles, slightly convex lateral margins, 
rounded apical angles, apical margin slightly convex to subtruncate; cell wall granulate, granules arranged in vertical 
series in the basal region of the semicells, sometimes in double rows, granules arranged in transverse series in the 
apical region, 3–5 pores surrounding the granules (SEM view); chloroplast with 2 pyrenoids; cells 44–47.5 μm long, 
26.5–28 μm wide, 10–12 μm isthmus.
 Holotype: material numbered (HUEFS 224658!) deposited at Herbário da Universidade Estadual de Feira de 
Santana, population partially illustrated here in LM (Figs 3–6) and SEM (Figs. 7–8).
 Paratypes (here designated): HUEFS 224676, HUEFS 224702, HUEFS 224718 (Herbário da Universidade 
Estadual de Feira de Santana, Feira de Santana, Brazil)
 Type locality and habitat: Collected at “Serra da Jiboia” Santa Teresinha, Bahia state, Brazil. 12° 51’ 6.6” S, 
39°28’ 35.5” W. Sample collected by G.J.P.Ramos, C.W.N. Moura and M.A. Santos (HUEFS 224658) on 14 January 
2015. Cosmarium amoenum var. jiboensis occurred in samples collected in bromeliad tanks (Alcantarea nahoumii); T 
27.5 (± 2.8 ºC). pH 5.9 (± 0.4). C0.25 (± 0.18 mS.cm-1). TDS 0.15 (± 0.12 ppt). DO 7.4 (± 3.9 mg.L-1).
 Etymology:—Due to Serra da Jiboia, type locality
 Notes:—Cosmarium amoenum var. jiboensis is morphologically similar to C. amoenum var. mediolaeve Nordstedt 
(1887:160) however this latter differs by having no granules in midregion of the semicells and elliptic cells in apical 
view. The newly described variety was found to be widely distributed in the bromeliads at Serra da Jiboia during all of 
the sampling periods, suggesting that it is well-adapted to phytotelmata conditions.
Cosmarium bahianum G.J.P.Ramos, C.E.M.Bicudo & C.W.N.Moura (2017: 67) (Figs 9–13)
Cell dimensions: 21–24 µm length, 18–19 µm width, 8.8–10 µm isthmus.
 Material examined: BRAZIL, Bahia, Santa Teresinha, Serra da Jiboia. 14-I-2015, G.J.P. Ramos et al. s/n (HUEFS 
224661), 18-VII-2015, G.J.P. Ramos et al. s/n (HUEFS 224675), 11-IX-2015, G.J.P. Ramos et al. s/n (HUEFS 224695), 
20-XI-2015, G.J.P. Ramos et al. s/n (HUEFS 224714)
 Habitat: bromeliad tanks (Alcantarea nahoumii); water temperature 25.8 (± 2.7 ºC). pH 5.4 (± 0.7). conductivity 
0.16 (± 0.15 mS.cm-1). total dissolved solids 0.08 (± 0.07 ppt). dissolved oxygen 8.0 (± 3.3 mg.L-1).
 Notes: This species was recently described from material collected in bromeliads at Parque das Dunas (Sand Dunes 
Park), Salvador, Bahia State, Brazil (Ramos et al. 2017a), and this second record occurred in very much the same type 
of habitat. Although the species was found to be widely distributed among bromeliads in the study area, we did not 
encounter zygospores, nor the subtriangular morphotype described from the material in the type locality. Nonetheless, 
the characteristic ellipsoid morphotype, as well as cell walls coarsely punctate, the presence of a mucilaginous sheath, 
cell dimensions, and the type of environment in which it was found in Serra da Jiboia, are in full agreement with the 
type material from Parque das Dunas in Salvador, Bahia.
 For now, C. bahianum was not found in other types of environments, but its frequent occurrence in bromeliads in 
different areas of the state suggests its preference for phytotelmata microhabitats.
RAMOS ET Al.62 • Phytotaxa 346 (1) © 2018 Magnolia Press
FIguRES 1–25. Closterium cornu var. minus LM (1, 2); Cosmarium amoenum var. jiboensis: LM (3–6); SEM (7–8), ventral view (3–4, 
7), lateral view (44) apical view (46–48) and detail of cell wall with granules surrounded by pores; C. bahianum: LM (9–12); SEM (13), 
ventral view (9–10, 13), lateral view (11) apical view (12); C. elegantissimum: LM (14–16); SEM (19), ventral view (14–15, 17–19), 
apical view (16); C. scrobiculosum: LM (20–23); SEM (24–25), ventral view (20–21, 25), lateral view (22) apical view (23–24). Scale bar 
10 μm (1–7, 9–24), 5 μm (8, 25).
BROMELIAD PHyTOTELMATA IN BRAZIL Phytotaxa 346 (1) © 2018 Magnolia Press • 63
*Cosmarium elegantissimum Lundell (1871:53) (Figs 14–19)
Cell dimensions: 80–100 µm length, 25–40 µm width, 22–34 µm isthmus.
 Material examined: BRAZIL, Bahia, Santa Teresinha, Serra da Jiboia. 14-I-2015, G.J.P. Ramos et al. s/n (HUEFS 
224656), 18-VII-2015, G.J.P. Ramos et al. s/n (HUEFS 224682), 11-IX-2015, G.J.P. Ramos et al. s/n (HUEFS 224698), 
20-XI-2015, G.J.P. Ramos et al. s/n (HUEFS 224716)
 Habitat: bromeliad tanks (Alcantarea nahoumii); T 27.3 (± 3.0 ºC). pH 6.0 (± 0.5). C 0.25 (± 0.16 mS.cm-1). TDS 
0.13 (± 0.10 ppt). DO 6.9 (± 2.8 mg.L-1).
 Notes: Morphologically, Cosmarium elegantissimum resembles C. pseudamoenum Wille; the latter species differs, 
however, by having irregularly arranged granules, cells subcircular-elliptic in apical view, and smaller cell dimensions. 
During our analyses we observed some morphological variations in which the semicells were less elongated and 
had finer granules (Figs 17–18) similar to Cosmarium simplicius (West & G.S.West) Grönblad (1931: 7), which had 
previously been considered a variety of Cosmarium elegantissimum by West & West (1908). That morphotype was 
often associated with C. elegantissimum in samples from Serra da Jiboia, showing intermediate features between them. 
We therefore consider that variation as an example of the phenotypic plasticity of C. elegantissimum in populations 
from Serra da Jiboia. Further studies will be needed to verify the relationship between C. elegantissimum and C. 
simplicius.
Cosmarium majae Strøm (1922:131) (Figs 26–33)
Cell dimensions: 7.5–9 µm length, 5–7 µm width, 3.5–5 µm isthmus, 11–13 µm zygospore.
 Material examined: BRAZIL, Bahia, Santa Teresinha, Serra da Jiboia. 14-I-2015, G.J.P. Ramos et al. s/n (HUEFS 
224655), 18-VII-2015, G.J.P. Ramos et al. s/n (HUEFS 224673), 11-IX-2015, G.J.P. Ramos et al. s/n (HUEFS 224691), 
20-XI-2015, G.J.P. Ramos et al. s/n (HUEFS 224719).
 Habitat: bromeliad tanks (Alcantarea nahoumii); T 25.2 (± 2.9 ºC). pH 5.8 (± 0.5). C 0.17 (± 0.15 mS.cm-1). TDS 
0.08 (± 0.07 ppt). DO 6.9 (± 2.3 mg.L-1).
 Notes: This species is known from only three states in Brazil: Bahia (Oliveira et al. 2016), São Paulo (Araújo 
& Bicudo 2006), and Amazonas (Förster 1974). It was observed in the phytotelmata of bromeliad specimens at Serra 
da Jiboia with a variable sinus, ranging from V-shaped to U-shaped, the former being the most common. As those 
variations occurred within the same population, and included cells with intermediate characteristics,we consider them 
as representing the phenotypic plasticity of a single taxon.
 Those variations could lead to taxonomic confusion, however, as there are several species of Cosmarium with 
similar morphologies. According to Coesel & Krienitz (2008), a more precise identification of these tiny Cosmarium 
will require finding zygospores, which can be very different from one species to another. Due to the presence of 
spherical zygospores with smooth walls, and in light of the morphological features of the cells, we considered that 
taxon to be Cosmarium majae
Cosmarium oliveirae G.J.P.Ramos, C.E.M.Bicudo & C.W.N.Moura sp. nov. (Figs. 34–38)
Cells 1.2–1.6 times longer than wide, elliptical contour, deep median constriction, closed linear sinus, dilated proximally 
closed; semicells subtrapeziform, basal margins retuse, subquadratic angles, lateral margins concave, rounded angles, 
subtruncated apex; cell wall decorated with small conical granules in rows, near the margin; semicells oblong in apical 
view, subelliptic in lateral view; 2-furcated chloroplast; pyrenoid 2 per semicell; cells 42–46,5 µm length, 35–28 µm 
width, 10–17 µm isthmus.
 Holotype: material numbered (HUEFS 224668!) deposited at Herbário da Universidade Estadual de Feira de 
Santana, population partially illustrated here in LM (Figs 34–37) and SEM (Fig. 38).
 Paratypes (here designated): HUEFS 224682, HUEFS 224707, HUEFS 224723 (Herbário da Universidade 
Estadual de Feira de Santana, Feira de Santana, Brazil)
 Type locality and habitat: Collected at “Serra da Jiboia” Santa Teresinha, Bahia state, Brazil. 12° 51’ 7.5” S, 
39°28’ 35.7” W. Sample collected by G.J.P.Ramos, C.W.N. Moura and M.A. Santos (HUEFS 224668) on 14 January 
2015. Cosmarium oliveirae occurred in samples collected in bromeliad tanks (Alcantarea nahoumii); T 27.8 (± 2.5 ºC). 
pH 6.2 (± 0.48). C 0.28 (± 0.15 mS.cm-1). TDS 0.14 (± 0.07 ppt). DO 7.2 (± 3.1 mg.L-1).
 Etymology:—the new name of the present species is given in honor of Brazilian desmidiologist Dr. Ivania Batista 
de Oliveira for her contribution to knowledge of desmids in Bahia.
RAMOS ET Al.64 • Phytotaxa 346 (1) © 2018 Magnolia Press
FIguRES 26–48. Cosmarium majae: LM (26–33); SEM (33), ventral view (27, 29–31, 33), apical view (26), lateral view (28) and 
zygospore (32); C. oliveirae: LM (34–37); SEM (38), ventral view (34–35, 38), apical view (36), lateral view (37); C. pachydermum var. 
aethiopicum : LM (39–42); SEM (43), ventral view (39–40, 43), lateral view (41), apical view (42); C. pseudoconnatum: LM (44–47); 
SEM (48), ventral view (44–45, 48), apical view (46–47). Scale bar 10 μm (26–32, 34–38, 44–47), 5 μm (33), 20 μm (39–43).
BROMELIAD PHyTOTELMATA IN BRAZIL Phytotaxa 346 (1) © 2018 Magnolia Press • 65
 Notes:—The newly described species is morphological similar to Cosmarium obtusatum (Schmidle) Schmidle 
(1898: 38), however this latter differ by having margins faintly crenate (flattened verrucae) and elliptical semicells in 
apical view. Due this similarity, this taxon was identified previously for this area as C. obtusatum (Ramos et al. 2011). 
Now, after a detailed reanalysis of this taxon, especially in SEM, the ornamentation of the granules is clearly different 
from pattern of C. obtusatum.
Cosmarium pachydermum var. aethiopicum (West & G.S.West) West & G.S.West (1905: 140) (Figs 39–43)
Cell dimensions: 110–120 µm length, 87–90 µm width, 30–37 µm isthmus.
 Material examined: BRAZIL, Bahia, Santa Teresinha, Serra da Jiboia. 14-I-2015, G.J.P. Ramos et al. s/n 
(HUEFS 224653), 18-VII-2015, G.J.P. Ramos et al. s/n (HUEFS 224673), 20-XI-2015, G.J.P. Ramos et al. s/n (HUEFS 
224711).
 Habitat: bromeliad tanks (Alcantarea nahoumii); T 26.8 (± 3.4 ºC). pH 6.2 (± 0.7). C 0.26 (± 0.14 mS.cm-1). TDS 
0.13 (± 0.06 ppt). DO 6.0 (± 1.5 mg.L-1).
 Notes: The cells of Cosmarium pachydermum var. aethiopicum have smooth outlines and are broadly elliptic 
rather than broadly oval, with thinner cell walls than in the nominate variety (Coesel & Meesters 2007). This taxon was 
previously identified from bromeliads at Serra da Jiboia as C. ralfsii (=Micrasterias ralfisii (Brebisson ex Ralfs) Skaloud 
et al.) (Ramos et al. 2011), however after reanalysis of the material we have identified it now as C. pachydermum var. 
aethiopicum. Confusion between those two taxa has been very common in floristic studies in Brazil, and although 
part of the literature considers the size of the isthmus as the main diagnostic feature, we consider the apical view to 
be a more relevant morphological feature for differentiating them. In apical view, C. pachydermum var. aethiopicum 
presents an ellipsoidal cell, whereas C. ralfsii (=M. ralfisii) appears fusiform.
 There are only three records of C. pachydermum var. aethiopicum in Brazil, two from São Paulo (Taniguchi et al. 
2003, Araujo 2006) and one from Mato Grosso do Sul (Santos 2008). The present record is the first from northeastern 
Brazil.
*Cosmarium pseudoconnatum Nordstedt (1870: 214) (Figs 44–48)
Cell dimensions: 44–48 µm length, 33–35µm width, 31–33 µm isthmus.
 Material examined: BRAZIL, Bahia, Santa Teresinha, Serra da Jiboia. 14-I-2015, G.J.P. Ramos et al. s/n 
(HUEFS 224658), 18-VII-2015, G.J.P. Ramos et al. s/n (HUEFS 224682), 11-IX-2015, G.J.P. Ramos et al. s/n (HUEFS 
2246703), 20-XI-2015, G.J.P. Ramos et al. s/n (HUEFS 224720).
 Habitat: bromeliad tanks (Alcantarea nahoumii); T 28.1 (± 2.0 ºC). pH 6.2 (± 0.4). C 0.30 (± 0.15 mS.cm-1). TDS 
0.17 (± 0.12 ppt). DO 7.4 (± 3.5 mg.L-1).
 Notes: During our analysis, we noted variations in the chloroplasts of this species, from 4-radial (most common) 
to 5-radial. Morphologically, C. pseudoconnatum can be easily confused with Cosmarium connatum Brébisson ex 
Ralfs. The most important difference is the shape of the chloroplast in apical view: 2-radial in C. connatum, at least 
4-radial in C. pseudoconnatum.
Cosmarium scrobiculosum Borge (1903:87) (Figs 20–25)
Cells 1.7 times longer than wide, deep median constriction, closed sinus linear, slightly dilated next to isthmus,; 
semicells subreniform, with rounded basal angles, convex lateral margins, rounded apical angles, margins granular-
undulate; cell wall scrobiculate with granules arranged in vertical series between scrobicules, 1 pore in each scrobicule 
(SEM view); in lateral view semicells subcircular, in apical broadly elliptic; chloroplast with 2 pyrenoids; cells (65–) 
70–85μm long, 45–50 μm wide, 45–50 μm isthmus.
 Material examined: BRAZIL, Bahia, Santa Teresinha, Serra da Jiboia. 14-I-2015, G.J.P. Ramos et al. s/n (HUEFS 
224666), 18-VII-2015, G.J.P. Ramos et al. s/n (HUEFS 224678), 11-IX-2015, G.J.P. Ramos et al. s/n (HUEFS 224699), 
20-XI-2015, G.J.P. Ramos et al. s/n (HUEFS 224718)
 Habitat: bromeliad tanks (Alcantarea nahoumii); T 27.3 (± 2.7 ºC). pH 5.9 (± 0.5). C 0.24 (± 0.17 mS.cm-1). TDS 
0.12 (± 0.08 ppt). DO 7.9 (± 4.0 mg.L-1).
 Notes: This taxon was previously known for this area as Cosmarium logiense Bissett (1884:194) (Ramos et al. 
2011), however after a detailed reanalysis of the material including SEM observations, we decided to reidentify it as 
C. scrobiculosum. The confusion between these taxa is common in some floristic studies in Brazil. The specimens 
RAMOS ET Al.66 • Phytotaxa 346 (1) © 2018 Magnolia Press
FIguRES 49–72. Docidium baculum: LM (49–50); SEM (51), ventral view (49) and the isthmus details (50–51); Euastrum quadriceps: 
LM (52–56); SEM (57), lateral view (52), ventral view (53, 57), apical view (54–56); Euastrum luetkemuelleri var. carniolicum: LM (58–
65) SEM (66–67), ventral view (58–60, 66), diagonal view (67), lateral view (61), apical view (62–63), zygospores (64–65); Actinotaenium 
mooreanum: LM (68–69, 71–72) SEM (70), ventral view (68–670) and zygospores (71–72). Scale bars 10 μm (Figs 42–59, 62–66), 5 μm 
(Figs 60–61). Scale bar 10 μm (49–65, 68–72), 5 μm (66–67).
BROMELIAD PHyTOTELMATA IN BRAZIL Phytotaxa 346 (1) © 2018 Magnolia Press • 67
identified as C. logiense by Oliveira(2011) and Biolo (2016), probably, refer to C. scrobiculosum. In Brazil, there are 
records of this species in north region (Grönblad 1945, Thomasson 1977), southeast (Borge 1918) and midwest (Borge 
1903).
Docidium baculum Brébisson ex Ralfs (1848: 158) (Figs 49–51)
Cell dimensions: 185–200 µm length, 9.5–10 µm width, 7–8 µm isthmus.
 Material examined: BRAZIL, Bahia, Santa Teresinha, Serra da Jiboia. 14-I-2015, G.J.P. Ramos et al. s/n (HUEFS 
224657), 18-VII-2015, G.J.P. Ramos et al. s/n (HUEFS 224676), 11-IX-2015, G.J.P. Ramos et al. s/n (HUEFS 224702), 
20-XI-2015, G.J.P. Ramos et al. s/n (HUEFS 224718).
 Habitat: bromeliad tanks (Alcantarea nahoumii); T 26.9 (± 3.2 ºC). pH 5.9 (± 0.6). C 0.23 (± 0.17 mS.cm-1). TDS 
0.13 (± 0.11 ppt). DO 7.2 (± 2.7 mg.L-1).
 Notes: This species is relatively easy to identify, as the base of the semicell is ornamented by a set of plications. 
Nonetheless, certain care was needed in the analysis of the populations from Serra da Jiboia because that species was 
often found in the same samples as Pleurotaenium trabecula. Although the latter species differs by having a semicell 
base without plications and larger cell dimensions, the populations of those taxa, when occurring together, give the 
false impression of morphological variation in a single species. This is the first report of the genus Docidium occurring 
in a phytotelmata habitat.
Euastrum luetkemuelleri var. carniolicum (Lütkemüller) Willi Krieger (1937: 561). (Figs 58–67)
Cell dimensions: 22–27 μm length, 12–16 μm width, 4.5–6 μm isthmus; Zygospore 19–26 μm diameter.
 Material examined: BRAZIL, Bahia, Santa Teresinha, Serra da Jiboia. 14-I-2015, G.J.P. Ramos et al. s/n (HUEFS 
224662), 18-VII-2015, G.J.P. Ramos et al. s/n (HUEFS 224676), 11-IX-2015, G.J.P. Ramos et al. s/n (HUEFS 224698), 
20-XI-2015, G.J.P. Ramos et al. s/n (HUEFS 224718).
 Habitat: bromeliad tanks (Alcantarea nahoumii); T 27.2 (± 3.2 ºC). pH 5.8 (± 0.4). C 0.24 (± 0.18 mS.cm-1). TDS 
0.11 (± 0.09 ppt). DO 7.4 (± 3.2 mg.L-1).
 Notes: Euastrum luetkemuelleri var. carniolicum was previously identified in the area as Euastrum sublobatum 
var. sublobatum f. elongatum (Ramos et al. 2011). A more detailed analysis of the material (including other cell views 
and SEM analysis), however, allowed us to state that it is a different taxon. Euastrum sublobatum shows a median 
intumescence in lateral view, a characteristic absent in E. luetkemuelleri.
 Some morphological variation was observed in the outline of the basal lobe, which could be retuse or slightly 
concave. Variation was also observed in the shape of the zygospores, ranging from ellipsoidal to spherical (the most 
common), always with spines.
*Euastrum quadriceps Nordstedt (1870: 216) (Figs 52–57)
Cell dimensions: 120–135 µm length, 59–66 µm width, 10–20 µm isthmus.
 Material examined: BRAZIL, Bahia, Santa Teresinha, Serra da Jiboia. 18-VII-2015, G.J.P. Ramos et al. s/n 
(HUEFS 224687), 20-XI-2015, G.J.P. Ramos et al. s/n (HUEFS 220176).
 Habitat: bromeliad tanks (Alcantarea nahoumii); T 25.2 (± 3.2 ºC). pH 5.6 (± 0.5). C 0.29 (± 0.55 mS.cm-1). TDS 
0.14 (± 0.15 ppt). DO 8.4 (± 2.9 mg.L-1).
 Notes: Euastrum. quadriceps occurred in only a few bromeliads at Serra da Jiboia, although, when found, it 
generally occurred in large populations. Low desmid diversity was commonly observed in those bromeliads, but among 
the few associated species were Cosmarium scrobiculosum, Pleurotaenium trabecula, and Micrasterias radians.
Micrasterias radians W.B.Turner (1893: 91) (Figs 73–92).
Cell dimensions: 130–190 µm length, 103–150 µm width, 16–25 µm isthmus.
 Material examined: BRAZIL, Bahia, Santa Teresinha, Serra da Jiboia. 14-I-2015, G.J.P. Ramos et al. s/n (HUEFS 
224657), 18-VII-2015, G.J.P. Ramos et al. s/n (HUEFS 224676), 11-IX-2015, G.J.P. Ramos et al. s/n (HUEFS 224704), 
20-XI-2015, G.J.P. Ramos et al. s/n (HUEFS 224718).
 Habitat: bromeliad tanks (Alcantarea nahoumii); T 27.5 (± 2.8 ºC). pH 6.0 (± 0.5). C 0.24 (± 0.16 mS.cm-1). TDS 
0.13 (± 0.10 ppt). DO 7.8 (± 4.2 mg.L-1).
RAMOS ET Al.68 • Phytotaxa 346 (1) © 2018 Magnolia Press
FIguRES 73–92. Micrasterias radians: LM (73–89); SEM (90–92), ventral view (73–76, 77–79, 82–92), apical view (80), lateral view 
(81) and anomalous forms (82–89). Scale bar 30 μm (73–89), 40 μm (90–92).
BROMELIAD PHyTOTELMATA IN BRAZIL Phytotaxa 346 (1) © 2018 Magnolia Press • 69
 Notes: In a previous study (Ramos et al. 2011), this taxon was identified as two distinct species: M. furcata 
C.Agardh ex Ralfs (1848: 73) and M. crux-melitensis Ralfs (1848: 73). When we reevaluated the material examined by 
Ramos et al. (2011), together with new collections from the area, we were able to identify those taxa as Micrasterias 
radians. That desmid showed the highest degree of polymorphism in all taxa studied, often presenting anomalous 
forms. The phenotypic plasticity of that species generated forms very similar to taxa such as M. furcata C.Agardh 
ex Ralfs (1848: 73) (with elongated lateral lobes) and M. crux-melitensis Ralfs (1848: 73) (with short lateral lobes). 
Another variable feature was the base of the polar lobe, which ranged from straight to intumescent, and the shape of 
the apex: slightly curved or straight.
 Authors such as Neustupa et al. (2010) and Coesel & Van Geest (2014) have discussed taxonomic problems in 
the Micrasterias radians complex, noting frequent confusion with the taxa mentioned above. Among the varieties of 
M. radians reported by those authors, the ones that are closest to the morphotypes from the Brazilian phytotelmata are 
M. radians var. radians and M. radians var. brasiliense (Grönblad) Willi Krieger & A.M.Scott. However, since their 
variation can be very wide, including with the presence of semicells with intermediary characteristics (in addition to 
other varieties of M. radians), we considered it better to identify all of the variation as belonging to a single taxon: 
Micrasterias radians. We therefore recommend further studies, preferably using a polyphase approach, to evaluate 
whether the phenotypic diversity of the Micrasterias radians complex occurring in bromeliads from Serra da Jiboia 
corresponds to the phenotypic expression of a single species or to different taxa.
Netrium digitus (Brébisson ex Ralfs) Itzigsohn & Rothe in Rabenhorst (1856:508) (Figs 93–96)
Cell dimensions: 160–260 µm length, 35–50 µm width
 Material examined: BRAZIL, Bahia, Santa Teresinha, Serra da Jiboia. 14-I-2015, G.J.P. Ramos et al. s/n (HUEFS 
224657), 18-VII-2015, G.J.P. Ramos et al. s/n (HUEFS 224672), 11-IX-2015, G.J.P. Ramos et al. s/n (HUEFS 224697), 
20-XI-2015, G.J.P. Ramos et al. s/n (HUEFS 224718).
 Habitat: bromeliad tanks (Alcantarea nahoumii); T 27.4 (± 2.8 ºC). pH 5.9 (± 0.4). C 0.23 (± 0.16 mS.cm-1). TDS 
0.13 (± 0.11 ppt). DO 8.3 (± 4.4 mg.L-1).
 Notes, Netrium digitus was the desmid most parasitized by chytrid fungi in bromeliads at Serra da Jiboia, with 
2–4 zoosporangia commonly found inside the cell. This pattern was also recorded in specimens from Holland (Coesel 
& Meesters 2007), which suggests that this relationship may be specific and more widely distributed than previously 
thought.
 Netrium digitus was one of the most predominant desmids, showing large populations in some bromeliads, often 
forming floating masses visible to the naked eye in bromeliad tanks. This is the first record of genus Netrium in a 
phytotelmata habitat.
*Pleurotaenium trabecula Nägeli (1849:104) (Figs 97–100)
Cell dimensions: 450–550 µm length, 30–45 µm width, 18–25 µm isthmus.
 Material examined: BRAZIL, Bahia, Santa Teresinha, Serra da Jiboia. 14-I-2015, G.J.P. Ramos et al. s/n (HUEFS 
224657), 18-VII-2015, G.J.P. Ramos et al. s/n (HUEFS 224676), 11-IX-2015, G.J.P. Ramos et al. s/n (HUEFS 224702), 
20-XI-2015, G.J.P. Ramos et al. s/n (HUEFS 224718).
 Habitat: bromeliad tanks (Alcantarea nahoumii); T 26.8 (± 2.9 ºC). pH 5.8 (± 0.6). C 0.22 (± 0.16 mS.cm-1). TDS 
0.12 (± 0.10 ppt). DO 7.7 (± 3.5 mg.L-1).
 Notes: Variousmorphological variations were observed in P. trabecula, principally semicells that were curved, 
but also narrow specimens that could be confused with P. ehrenbergii (Ralfs) De Bary. The latter species, however, 
usually shows small papillae at its apices. Although Škaloud et al. (2012) considered the culture strains identified in 
Europe as P. ehrenbergii and P. trabecula to be polyphyletic, and are of opinion that the presence/absence of granules 
at the apex does not support their mutual separation, we prefer to identify our taxon under discussion as P. trabecula 
(absence of papillae at the apices—traditional taxonomy) until more detailed studies can clarify their true identity. 
Pleurotaenium trabecula has also been reported in bromeliad phytotelmata in Rio de Janeiro State (Sophia et al. 
2004).
RAMOS ET Al.70 • Phytotaxa 346 (1) © 2018 Magnolia Press
FIguRES 93–112. Netrium digitus: LM (93–95); SEM (96); Pleurotaenium trabecula: LM (97–98); SEM (99–100); Staurastrum 
pseudosebaldi var. compactum LM (101–104), ventral view (101–103), apical view (104); Staurastrum pseudoteliferum sp. nov.: LM 
(105–106, 109–110); SEM (107–108), ventral view (105–107), apical view (108–110); Xanthidium mamillosum var. borgei: LM (111–
112); ventral view (111), apical view (112). Scale bar 50 μm (93–95), 60 μm (92–94), 40 μm (97), 30 μm (104–105, 111–112), 20 μm (96, 
99–100), 10 μm (98, 101–108)
BROMELIAD PHyTOTELMATA IN BRAZIL Phytotaxa 346 (1) © 2018 Magnolia Press • 71
Staurastrum pseudosebaldi var. compactum A.M.Scott & Grönblad (1957: 44) (Figs 101–104)
Cells 3–radiate, 0.9–1.1 times wider than long, median constriction shallow, sinus open, processes generally short, 
with 3 short spines at their terminations; semicells subtrapeziform, triangular in apical view, with an intramarginal 
series of warts, usually curves into direction to the angles, margins between the processes almost straight, slightly 
crenate; chloroplast axial. Cells 43.5–46 μm long, 50–52 μm wide, 12.5–15 μm isthmus. 
 Material examined: BRAZIL, Bahia, Santa Teresinha, Serra da Jiboia. 14-I-2015, G.J.P. Ramos et al. s/n 
(HUEFS 224663), 18-VII-2015, G.J.P. Ramos et al. s/n (HUEFS 224683), 11-IX-2015, G.J.P. Ramos et al. s/n (HUEFS 
224693).
 Habitat: bromeliad tanks (Alcantarea nahoumii); T 27.2 (± 3.6 ºC). pH 5.7 (± 0.5). C 0.13 (± 0.08 mS.cm-1). TDS 
0.27 (± 0.31 ppt). DO 5.0 (± 1.5 mg.L-1).
 Notes: This variety has apical processes shorter than the typical variety; however, we encountered individuals 
with both semicells having either long or short processes (with the latter pattern being predominant). This variety was 
one of the rarest of all of the desmids identified in phytotelmata at Serra da Jiboia (Table 1). The desmid Staurastrum 
pseudosebaldi var. compactum was previously known for the United States (Scott & Grönblad 1957) and China (Wei 
2014). This is the third record of the variety, and the first for South America.
Staurastrum pseudoteliferum G.J.P.Ramos, C.E.M.Bicudo & C.W.N.Moura sp. nov. (Figs 105–110)
Cells 3–radiate, 1.3 times longer than wide, median constriction deep, sinus open; semicells broadly elliptic, 
subtriangular in apical view, angles widely rounded, margins straight between processes, provided with stout spines 
usually arranged linearly or slightly curved in 2 rows of 4 spines at angles, with approximately 9 tiny spines in the 
margins (3 on each side). Cells 31–32.5 μm long (without spines), 23.5–25 μm wide, 10–11.5 μm isthmus.
 Holotype: material numbered (HUEFS 224653!) deposited at Herbário da Universidade Estadual de Feira de 
Santana, population partially illustrated here in LM (Figs 105–106, 109–110) and SEM (Fig. 107–108).
 Paratypes (here designated): HUEFS 224684, HUEFS 224705, HUEFS 224716 (Herbário da Universidade 
Estadual de Feira de Santana, Feira de Santana, Brazil)
 Type locality and habitat: Collected at “Serra da Jiboia” Santa Teresinha, Bahia state, Brazil. 12° 51’ 7” S, 
39°28’ 35.4” W. Sample collected by G.J.P.Ramos, C.W.N. Moura and M.A. Santos (HUEFS 224653) on 14 January 
2015. Staurastrum pseudoteliferum was observed in samples collected in bromeliad tanks (Alcantarea nahoumii); T 
26.7 (± 3.1 ºC). pH 6.0 (± 0.5). C 0.22 (± 0.17 mS.cm-1). TDS 0.13 (± 0.11 ppt). DO 7.5 (± 4.3 mg.L-1).
 Etymology:—Due to morphological similarly with Staurastrum teliferum 
 Notes:—Staurastrum pseudoteliferum differs from Staurastrum teliferum Ralfs (1848: 128) by having two rows 
of 4 spines at the angles in the ventral view, apical view showing approximately 9 small spines located near the margins, 
3 on each side. Additionally, S. teliferum has, in apical view, triangular semicells with slightly concave margins, with 
5 tiny spines on each side. Another taxon similar is St. teliferum var. groenbladii Förster (1964), however, this variety 
differs by having semicircular-slightly elliptic cells (in front view), semicells ovoid (to subsphaerical) in apical view, 
and thicker spines. 
Xanthidium mamillosum var. borgei K.Förster (1981: 247): (Figs. 111–112)
Cells 1.1 times wider than long; semicells about hexagonal, medial constriction deep, convex basal margins, angle with 
1 pair of long, straight spines, with almost mamiliform base, arranged side by side, lateral margins slightly convex; 
apical margin slightly concave, angles with 1 pair of spines similar to those of the lower angles; Cell length without 
spines 53 µm, with spines 84 µm; cell width without spines 58 µm, with spines 90 µm, 23–24µm isthmus.
 Material examined: BRAZIL, Bahia, Itaberaba, Fazenda Itaberaba. 21-I-2015, G.J.P. Ramos et al. s/n (HUEFS 
215777).
 Habitat: bromeliad tanks (Aechmea cf. lingulatoides); T 29.9 ºC. pH 5.3. C 0.35 mS.cm-1. TDS 0.15 ppt. DO 3.6 
mg.L-1.
 Notes: Xanthidium mamillosum var. borgei was the only desmid encountered in Aechmea cf. lingulatoides tanks 
in the region of Itaberaba, Bahia State. Those bromeliads are found in the semiarid Caatinga domain and tend to 
accumulate very little water; they can accumulate small amounts of moisture in the basal regions of their leaves or 
dry out completely. Although we only found that desmid in a single bromeliad, its mucilage sheath appears to be an 
important structure that allows it to survive (possibly only temporarily) in the phytotelmata by minimizing the effects 
RAMOS ET Al.72 • Phytotaxa 346 (1) © 2018 Magnolia Press
of desiccation. A few specimens were observed in the sample, always showing that same morphological pattern. In 
Brazil, this variety occurs in central-west region (Borge 1925) and in Amazonian region (Förster 1969, Souza & Melo 
2011). This is the first record of this variety for northeast Brazil.
FIguRES 113–116. Bromeliads from Serra da Jiboia before the huge fire (November 2015) (113–114); Bromeliads from Serra da Jiboia 
after the huge fire (April 2016) (115–116).
Final remarks 
Desmids are one of the main groups of freshwater microalgae that occur in bromeliad phytotelmata in Brazil, 
microhabitats with oligotrophic conditions in acidic water with low conductivity. The analyses of the environmental 
data at Serra da Jiboia, indicated that the bromeliads sampled had the greatest diversity of desmids (11 taxa) in 
November 2015 when the water in their phytotelmata had a temperature near 27 ºC, pH 6.5, a C of 0.45 mS.cm-1, and 
a DO level of 7 mg.L-1. The lowest diversity was observed in July/2015, with only one desmid species (P. trabecula), 
when the water temperature was 25.8 ºC, pH 4.8, C of 0.09 mS.cm-1 and DO level of 13.7 mg.L-1. Large populations 
of phytoflagellates (Euglenophyceae) such as Heterenoma sp. and Phacus polytrophos Pochmann (1942: 128) were 
observed inhabiting those bromeliads which had low desmid diversity.
 Remarkably, all of the 80 bromeliads sampled at Serra da Jiboia had at least one desmid species, demonstrating 
the adaptability and wide distribution of those microalgae. That was due, among other factors, to local environmental 
conditions that allow the maintenance of waterin bromeliad tanks even during extended dry periods. 
BROMELIAD PHyTOTELMATA IN BRAZIL Phytotaxa 346 (1) © 2018 Magnolia Press • 73
 The only desmid species recorded at the Itaberaba site was Xanthidium mamillosum var. borgei, and it occurred in 
only one bromeliad out of a total of 80 sampled. The bromeliads in that area are subjected to conditions totally different 
from Serra da Jiboia, as Itaberaba is in a semiarid area (Caatinga domain) under a high temperature regime, and very 
little water is available in their tanks. Nonetheless, it is possible that other desmids are likewise adapted to the adverse 
conditions of the phytotelmata (with little water available) in other areas of the Caatinga. Lyra (1976), although focused 
on diatoms, reported the occurrence of desmids in several bromeliads from semiarid areas in Pernambuco State in 
northeastern Brazil, including Micrasterias sp, Euastrum sp., Closterium sp., and Cosmarium sp. (the most frequent), 
occurring in acidic conditions (pH 5.5-6) and high temperatures (up to 29 ºC).
 The little that is known about the desmids from bromeliad phytotelmata in Brazil has mainly been reported from 
“restinga” (nearshore, sandy) areas in Rio de Janeiro State (Sophia 1999, Sophia et al. 2004). Comparisons of the 
desmids identified in Bahia with those from Rio de Janeiro indicated Pleurotaenium trabecula as only one species in 
common, which was the most frequent desmid in Serra da Jiboia (Table 1). 
 There are approximately 44 genera and 1340 species of bromeliads known from several ecosystems in Brazil 
(Forzza et al. 2015), most of them with possible phytotelmata, although only 7 genera have been reported with 
desmids in their tanks. Table 2 lists all of the desmid taxa recorded in bromeliad phytotelmata in Brazil, citing the 
vegetation formations where they were encountered and their genera. The genus Cosmarium predominates, with 18 
representatives.
 There have been only two reports of the occurrence of desmids in bromeliad tanks outside of Brazil: Euastrum 
bipartitum Krieger (1932: 211) in Mexico (Hernández-Rodríguez et al. 2014) and Cosmarium wittrockii Lundell 
(1871: 31) in French Guiana (Carrias et al. 2014). Further studies will therefore the needed to increase our knowledge 
of desmids in those overlooked environments in countries throughout the Neotropical region.
 It is important to stress the necessity of protecting bromeliads in both of the areas studied here, especially 
Alcantarea nahoummi at Serra da Jiboia—where they are constantly threatened by burning and harvesting (to be sold 
locally for ornamental purposes) (Versieux & Wanderley 2010, Forzza et al. 2015). A couple of months after our last 
sampling which was in November/2015 (Figs. 113–114), a huge fire (arson) devastated most of the bromeliads in that 
area (Figs. 115–116). There are many rare or even endemic microalgae species inhabiting those bromeliads, and if their 
habitats are threatened those microalgae will likewise be directly impacted. Thus, the conservation of the bromeliads 
at Serra da Jiboia is absolutely necessary for the maintenance of their phytotelmata communities.
TABLE 1. List of all desmids in bromeliad phytotelmata from Serra da Jiboia (Santa Teresinha) and Fazenda Itaberaba 
(Itaberaba), Bahia State, Brazil with their indicative notations. Acidity, (aci) acidic, (neu) neutral, (alk) alkalic; Frequency of 
occurrence, (1) very frequent; (2) frequent; (3) occasional; (4) rare; (5) very rare. 
 Serra da Fazenda
Jiboia Itaberaba
Desmids Acidity Frequency Acidity Frequency
Actinotaenium mooreanum acid 5 - -
Closterium cornu var. minus acid-neu 3 - -
Cosmarium amoenum var. jiboensis acid-neu 3 - -
Cosmarium bahianum acid 3 - -
Cosmarium elegantissimum acid-neu 2 - -
Cosmarium majae acid-neu 3 - -
Cosmarium oliveirae acid-neu 4 - -
Cosmarium pachydermum var. aethiopicum acid-neu 4 - -
Cosmarium pseudoconnatum acid 3 - -
Cosmarium scrobiculosum acid-neu 2 - -
Docidium baculum acid-neu 2 - -
Euastrum luetkemuelleri var. carniolicum acid 3 - -
Euastrum quadriceps acid 5 - -
Micrasterias radians acid-neu 2 - -
Netrium digitus acid 3 - -
Pleurotaenium trabecula acid-neu 1 - -
Staurastrum pseudosebaldi var. compactum acid 5 - -
Staurastrum pseudoteliferum acid-neu 3 - -
Xanthidium mamillosum var. borgei - - acid 5
RAMOS ET Al.74 • Phytotaxa 346 (1) © 2018 Magnolia Press
TABLE 2. Desmid taxa recorded in Brazilian bromeliad phytotelmata (BA—Bahia, RJ—Rio de Janeiro, PE—Pernambuco): 
1—present study, 2—Ramos et al. (2011), 3—Ramos et al. (2017a), 4—Ramos et al. (2017b), 5—Sophia (1999), 6—Sophia 
et al. (2004), 7—Lyra (1976), 8—Kolkwitz (1933). Bromeliad genera: A = Alcantarea (E.Morren ex Mez) Harms P = 
Portea Brongniart & K.Koch, H = Hohenbergia Schultes. & Schultes.f., Ae = Aechmea Ruiz & Pavón. B = Bromelia L., N 
= Neoregelia L.B.Smith., Ni = Nidularium Lemaire. Misidentification: ■ Cosmarium logiense, □ C. obtusatum, ○ C. ralfsii, 
● Euastrum sublobatum f. elongatum, ♦ Micrasterias furcata and M. crux-melitensis.
Taxa Vegetation Formation Bromeliad Brazilian studies
BA RJ PE
Actinotaenium mooreanum (W.Archer) Teiling Granite outcrops A 1
A. wollei (West & G.S.West) Teiling Granite outcrops A 2
Closterium cornu var. minus Irénée-Marie Granite outcrops A 1
Closterium sp. Caatinga H, P 7
Cosmarium amoenum var. jiboensis G.J.P.Ramos, C.E.M.Bicudo & 
C.W.N.Moura
Granite outcrops A 1
C. bahianum G.J.P.Ramos, C.E.M.Bicudo & C.W.N.Moura Granite outcrops, Restinga A, H 1, 3
C. blyttii Wille Granite outcrops A 2
C. botrytis Meneghini ex Ralfs Restinga Ae, B, N 5
C. elegantissimum P.Lundell Granite outcrops A 1, 2
C. excavatum Nordstedt Granite outcrops A 2
C. furcatum Förster Restinga Ae, B, N 5
C. majae Strøm Granite outcrops A 1
C. obsoletum (Hantzsch) Reinsch Granite outcrops A 2
C. oliveirae G.J.P.Ramos, C.E.M.Bicudo & C.W.N.Moura. Granite outcrops A 1, 2□
C. pseudoconnatum Nordstedt Granite outcrops A 1, 2
C. pachydermum var. aethiopicum (West & G.S.West) West & 
G.S.West
Granite outcrops A 1, 2○
C. scrobiculosum Borge Granite outcrops A 1, 2■
C. speciosum var. simplex Nordstedt Restinga Ae, B, N 5
C. subarctoum var. punctatum (West & G.S.West) Willi Krieger & 
Gerloff
Restinga Ae, N 6
C. undulatum var. circulare Irénée-Marie Granite outcrops A 2
Cosmarium sp1. Restinga Ni 8
Cosmarium sp2. Caatinga H, P 7
Desmidium grevillei (Kützing ex Ralfs) De Bary Restinga Ae, B, N 5
D. cylindricum Greville Restinga Ae, N 6
D. laticeps var. quadrangulare Nordstedt Restinga Ae, N 6
D. quadratum Nordstedt Restinga Ae, B, N 5
Docidium baculum Brébisson ex Ralfs Granite outcrops A 1
Euastrum crameri Raciborski Restinga Ae, N 6
E. denticulatum F.Gay Restinga Ae, B, N 5
E. luetkemuelleri var. carniolicum (Lütkemüller) Willi Krieger Granite outcrops A 1, 2●
E. quadriceps Nordstedt Granite outcrops A 1, 2
Euastrum sp. Caatinga H, P 7
Micrasterias radians W.B.Turner Granite outcrops A 1, 2♦ 
M. rotata Ralfs Restinga Ae, N 6
Micrasterias sp. Caatinga H, P 7
Netrium digitus (Brébisson ex Ralfs) Itzigsohn & Rothe Granite outcrops A 1
...continued on the next page
BROMELIAD PHyTOTELMATA IN BRAZIL Phytotaxa 346 (1) © 2018 Magnolia Press • 75
TABLE 2. (Continued)
Taxa Vegetation Formation Bromeliad Brazilian studies
BA RJ PE
P. ehrenbergii (Ralfs) De Bary Restinga Ae, B, N 5, 6
P. trabecula Nägeli Restinga A, Ae, N 1, 2 6
P. trabecula var. elongatum Cedergren Restinga Ae, B, N 5
Pleurotaenium sp. Restinga Ni 8
Spirotaenia closteridia (Kützing) Rabenhorst Granite outcrops A 4
S. endospira W.Archer Granite outcrops A 4
S. filiformis G.J.P.Ramos, C.E.M.Bicudo & C.W.N.Moura Granite outcrops A 4
Staurastrum anatinum var. parvum (West) Prescott Restinga Ae, B, N 5
S. chaetoceras (Schröder) G.M.Smith Restinga Ae, B, N 5
S. hirsutum Ehrenberg ex Ralfs Restinga Ae, B, N 5, 6
S. leptocladum var. coronatum A.M.Scott & Grönblad Granite outcrops A 2
S. pseudosebaldi var. compactum A.M.Scott & Grönblad Granite outcrops A 1
S. pseudoteliferumG.J.P.Ramos, C.E.M.Bicudo & C.W.N.Moura. Granite outcrops A 1
Staurodesmus convergens var. laportei Teiling Restinga Ae, N 6
S. triangularis var. subparallelus (G.M.Smith) Teiling Restinga Ae, B, N 5
Xanthidium. concinnum W.Archer Restinga Ae, B, N 5, 6
X. mamillosum var. borgei K.Förster Caatinga Ae 1
Acknowledgments
Authors are deeply grateful to Peter F.M. Coesel for their valuable comments leading to an improvement of the 
manuscript, to CNPq, Conselho Nacional de Desenvolvimento Científico e Tecnológico and FAPESB, Fundação de 
Amparo à Pesquisa do Estado da Bahia (Project “Flora da Bahia”, 483909/2012) for financial support; and to the 
“Serviço de Microscopia Eletrônica” of CPqGM of the technological facilities of FIOCRUZ for SEM use. GJPR 
thanks FAPESB for doctoral fellowship (Nº BOL0513/2014).
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