Comparative pollen morphology and taxonomic ... - CNCFlora

Comparative pollen morphology and taxonomic considerations in Eriocaulaceae 

Ricardo Landim Bormann de Borges ⁎, Francisco de Assis Ribeiro dos Santos, Ana Maria Giulietti 

Programa de Pós-Graduação em Botânica, Universidade Estadual de Feira de Santana (UEFS), Av. Transnordestina s.n. Novo Horizonte 44036-900 Feira de Santana-BA, Brazil 

article info 

Article history: 

Received 27 June 2008 

Received in revised form 2 December 2008 

Accepted 14 December 2008 

Available online 24 December 2008 

Keywords: 

Palynology 

Poales 

Spiraperturate 

1. Introduction 

abstract 

Eriocaulaceae is a monocot group that has very small, unisexual 

flowers grouped into a dense “capitulum”, a 3–2 locular superior 

ovary, one ovule per locule, and spiraperturate pollen (Giulietti et al., 

2000). The family is included in the Poales (APG II, 2003), and is 

considered a monophyletic group (Giulietti et al., 2000; Linder and 

Rudall, 2005). 

Eriocaulaceae includes about 1200 species in 11 genera and shows 

a pantropical distribution, where this is especially true for Eriocaulon. 

Mesanthemum is an endemic African genus and all others are 

American genera. Paepalanthus and Syngonanthus also have a small 

number of species in the African continent. A great number of species 

occurs specially in mountainous neotropical regions as part of the 

savanna and “campos rupestres” types of vegetation. 

Economically, the species of Eriocaulaceae are responsible for being 

a source of income for the municipal districts of the mountain range of 

Espinhaço (Minas Gerais and Bahia), Serra Geral (Goiás) and the 

cerrados of Planalto Central (Giulietti et al., 1988). Nowadays, in the 

Tocantins State, Syngonanthus nitens (Bong.) Ruhland (golden grass) 

has been the most important economic species. Residents of this area 

have received a large boost to their income because of the handicrafts, 

which they produce, made with the yellowish floral scape of that 

⁎ Corresponding author. 

E-mail addresses: rlbborges@gmail.com (R.L.B. de Borges), fasantos@uefs.br 

(F. de Assis Ribeiro dos Santos), anagiulietti@hotmail.com (A.M. Giulietti). 

0034-6667/$ – see front matter © 2009 Elsevier B.V. All rights reserved. 

doi:10.1016/j.revpalbo.2008.12.008 

Review of Palaeobotany and Palynology 154 (2009) 91–105 

Contents lists available at ScienceDirect 

Review of Palaeobotany and Palynology 

journal homepage: www.elsevier.com/locate/revpalbo 

Pollen morphology of 55 species from 10 of the 11 currently recognized genera of Eriocaulaceae 

(Actinocephlaus, Blastocaulon, Eriocaulon, Lachnocaulon, Leiothrix, Paepalanthus, Philodice, Rondonanthus, 

Syngonanthus and Tonina) has been studied using light and scanning electron microscopy. Species presented 

pollen grains in monads, which were small (Blastocaulon, Lachnocaulon, Philodice and Tonina) to medium 

sized (Eriocaulon, Leiothrix and Rondonathus), while the other genera presented pollen grains in both size 

classes. Apertures were sulci, mainly forming spirals, in one, two or three units in different features. The 

exine is thin, less than 3 µm in thickness; there are two types of supratectal processes, granulose and 

spinulose. The spinulose processes can be acute or obtuse, with side walls being straight, concave or convex. 

The pollen characteristics do not support the segregation of the genera Actinocephalus and Paepalanthus. The 

Syngonanthus species are the only ones to present two apertural patterns: spiraperturate and 2-zonasulcate. 

The spiraperturate species of Syngonanthus also possess similar characteristics to the genera Actinocephalus 

and Paepalanthus, so much so that it is not possible to distinguish them. The pollen morphology data support 

taxonomic arrangements at different levels: the separation of the family into two subfamilies, Eriocauloideae 

and Paepalanthoideae, the separation from Syngonanthus of sect. Eulepis by its 2-zonasulcate pollen grains 

and the segregation of Tonina fluviatilis, due to its grooved spines, into its own genus. 

© 2009 Elsevier B.V. All rights reserved. 

species, which is known for its gracefulness and color and is used to 

make earrings, bags, and others adornments (Schimidt et al., 2007). 

Although a significant number of studies on the Eriocaulaceae have 

been published in the last 100 years – the majority within the realm of 

taxonomy – the most complete taxonomic study of the family was 

undertaken by Ruhland (1903). Among the most important publications 

are those of Melchior (1964), Stützel (1985), Hensold (1991), 

Hensold and Giulietti (1991), Phillips (1997), Giulietti et al. (2000), 

and Sano (2004). 

Very few of the taxonomic studies outlined above have focused on 

the pollen morphology of the Eriocaulaceae. The pollen morphology of 

representatives of the family has been little studied, and most of the 

published papers have been restricted to Old World taxa (Table 1). 

Nevertheless, some important pollen studies have been published. 

The most complete and systematic study, however, was undertaken by 

Thanikaimoni (1965); who described, by light microscopy, the pollen 

grains of 54 species distributed among several genera of Eriocaulaceae 

(Table 1). His study did not include species of the genera Actinocephalus 

(Wikstr.) Sano, Blastocaulon Ruhland, Mesanthemum Körn nor 

Rondonanthus Herzog. 

Another important work on pollen grains of the Eriocaulaceae was 

published by Santos et al. (2000), who studied species of the genus 

Paepalanthus from the Brazilian flora (eight of which are currently 

assigned to the genus Actinocephalus). These authors indicated the 

necessity of still more detailed studies using scanning electron 

microscopy to examine more closely the pollen characteristics of the 

species.

92 R.L.B. de Borges et al. / Review of Palaeobotany and Palynology 154 (2009) 91–105 

Table 1 

Previous morphological pollen data of representatives of the Eriocaulaceae family 

Authors Taxa Method Subject 

Erdtman (1943) Eriocaulon septangulare Withering LM Pollen morphology and plant taxonomy - Angiosperms 

Kuprianova (1948) Eriocaulon benthamii Kunth and Paepalanthus blepharocnemis Mart. LM Pollen morphology of monocotyledons 

Erdtman (1952) Syngonanthus wahlbergii (Wikstr.) Ruhl and Tonina fluviatilis Aubl. LM Pollen morphology and plant taxonomy - Angiosperms 

Ikuse (1956) Eriocaulon miquelianum Körn., E. nipponicum Maxim., E. parvum Körn, 

E. senile Honda, and E. sielboldianum Sieb. et Zucc. (= E. cinereum R. Br.) 

LM Pollen flora of Japan 

Thanikaimoni (1965) Eriocaulon L. (46 spp.), Lachnocaulon Kunth (2 spp.), 

Leiothrix Ruhland (1 sp.), Paepalanthus Mart. (1 sp.), 

Philodice Mart. (1 sp.), Syngonanthus Ruhland (2 spp.), 

and Tonina Aubl. (1 sp.) 

LM Pollen morphology of the Eriocaulaceae family 

Huang (1972) Eriocaulon cinerum R. Br. Var. sieboldianum (Sieb. & Zucc.) T. Kayama, 

E. merrillii Ruhl., E. nantoense Hay., E. trisectum Satake, 

E. truncatum Buch.-Ham. 

LM Pollen flora of Taiwan 

Ybert (1979) Mesanthemum prescottianum (Bong.) Körn. LM Pollen flora of the Ivory Coast 

Furness (1988) Eriocaulon aquaticum (Hill) Druce. LM, SEM Eriocaulaceae in the Northwest European pollen flora 

Santos et al. (2000) Actinocephalus (8 spp.) and Paepalanthus (7 spp.) LM Pollen morphology of some Brazilian Eriocaulaceae 

Rull (2003) Paepalanthus perplexans Mold. LM Illustrated pollen key of some Venezuelan species 

Coan et al. (2007a) Syngonanthus caulescens (Poir.) Ruhland SEM Embryological study of the species, only SEM pictures 

of pollen grains were presented 

Coan et al. (2007b) Leiothrix fluitans (Mart.) Ruhland SEM Embryological study of the species, only SEM pictures 

of pollen grains were presented 

Method: LM — light microscopy, SEM — scanning electron microscopy. 

The first study of the Eriocaulaceae using scanning electron 

microscopy was undertaken by Furness (1988), who described the 

pollen grains of Eriocaulon aquaticum (Hill) Druce. Coan et al. (2007a,b) 

published electro-micrographs of the pollen grains of Syngonanthus 

caulescens (Poir.) Ruhland and Leiothrix fluitans (Mart.) Ruhland as a 

part of their study of the embryology of these taxa. 

Considering all the pollen morphological data related to the 

Eriocaulaceae to date, only 87 taxa have been described, of which 60 are 

species belonging to the genus Eriocaulon. Furthermore, the principal 

pollen study of the family (Thanikaimoni, 1965) was published 43 years 

ago and employed only light microscopy. For these reasons, the objective 

of the present work was to provide a more complete study of the family 

and to clarify the patterns of the pollen apertures exhibited by the different 

taxonomic groups, using both light and scanning electron microscopy. 

2. Materials and methods 

In accordance with Ruhland's (1903) classification system for 

Eriocaulaceae, we studied the pollen grains of 55 species from the 

genera Actinocephalus, Blastocaulon, Eriocaulon, Lachnocaulon, Leiothrix, 

Paepalanthus, Philodice, Rondonanthus, Syngonanthus, andTonina. The 

genus Mesanthemum was not included in this work due to the lack of 

study material, but both of the subfamilies cited by Ruhland (1903), 

Eriocauloideae and Paepalanthoideae, were considered. Taxonomic 

arrangement of the group is: 

Order Poales 

Fam. Eriocaulaceae 

Subfam. Eriocauloideae 

Eriocaulon 400/4 1 

Mesanthemum 12/0 

Paepalanthoideae 

Actinocephalus 29/6 

Blastocaulon 5/2 

Lachnocaulon 7/2 

Leiothrix 37/9 

Paepalanthus 485/15 

Philodice 2/1 

Rondonanthus 6/1 

Syngonanthus 200/14 

Tonina 1/1 

1 Total number of species/number of species included in present study. 

Pollen samples (the capitula) for our analyses were obtained from 

herbarium specimens that had been securely identified and incorporated 

into the Herbário da Universidade Estadual de Feira de Santana — 

HUEFS, Herbarium of the Miami University — MU, and Herbário do 

Departamento de Botânica, Instituto de Biociências (Universidad de São 

Paulo) — SPF. All of the specimens examined are listed in the Appendix A. 

The acetolysis method (Erdtman, 1960) was used for the light 

microscope (LM) analysis of the pollen grains of 36 species. For each 

species, at least 20 pollen grains were used (except when not possible) 

in the morphometric analyses of the principal pollen parameters: 

diameter, width of the interapertural exine strip, thickness of the 

exine layers, and length of the spines. 

For the scanning electron microscopy (SEM) studies, 41 species of the 

above-mentioned genera were examined. Due to the fragile nature of 

the pollen grains and their low production, the anthers were removed 

from the flowers and opened directly over a stub covered by doublefaced 

self-adhesive carbon tape. The material was subsequently coated 

with gold and analyzed (LEO 1430 VP — Carl Zeiss). 

Photographs of the pollen characteristics were obtained under 

both LM and SEM techniques. The palynological nomenclature used 

follows Punt et al. (2007), and that of the type of aperture follows 

Pozhidaev (2000). 

All of the prepared material is deposited in the palynotheca of the 

Laboratory of Plant Micromorphology (LAMIV) at the Universidade 

Estadual de Feira de Santana, Bahia State, Brazil. 

3. Results 

3.1. General pollen morphology 

All species of Eriocaulaceae studied here had pollen grains that 

were monads, generally spherical, and varied in size from small (up to 

25 µm) to medium (N25 µm) (Table 2). The smallest pollen grains are 

observed in Blastocaulon albidum, Lachnocaulon minus, Paepalanthus 

tortilis, and P. subtilis, while the largest grains are seen in Paepalanthus 

comans, P. planifolius (both P. subg. Platycaulon) and Rondonanthus 

duidae. Variations among the sizes of the pollen grains of the same 

specimen are relatively large: up to 14 µm as recorded for Syngonanthus 

schwackei (T. B. Cavalcanti et al. 2906) and 15 µm for Paepalanthus 

pulvinatus (M. J. G. Andrade 106). This fact may be due to 

rupturing of the aperture during chemical treatment (acetolysis), 

which increases the size of the pollen grains. 

Constant features among the pollen grains of the species of 

Eriocaulaceae that were analyzed are the spiral apertures and that the

Table 2 

Morphometric data and apertural type of pollen grains of studied Eriocaulaceae species through light microscopy 

Species Diameter⁎ Interapertural 

strips⁎ 

surface of the exine is composed of (micro)spinose and granulose 

supratectal processes (Tables 2–3, Plates I–VII). 

3.2. Apertures 

The pollen grains of Eriocaulaceae are spirapeturate. They have 

one or more spiral apertures. Apertures isolate strips of exine 

between the spirals (Plate I, 3;Plate IV, 9;Plate VI, 1;Plate VII, 5). 

Here, these strips are considered to be “interapertural exine”. The 

R.L.B. de Borges et al. / Review of Palaeobotany and Palynology 154 (2009) 91–105 

Exine 

thickness⁎ 

Apertural type 

Actinocephalus 

Actinocephalus bongardii (A. St.-Hill.) Sano (21) 24 (26) (7) 9 (12) (1) 1 (1) Spiraperturate (helicoidal) 

Actinocephalus denudatus (Körn.) Sano (25) 28 (31) (9) 11 (14) (1) 1 (2) Spiraperturate (helicoidal) 

Actinocephalus divaricatus (Bong.) Sano (24) 27 (33) (8) 10 (15) (1) 1 (1) Spiraperturate (helicoidal, other arrangements) 

Actinocephalus ramosus (Wikstr.) Sano (24) 27 (33) (9) 10 (13) (1) 1 (1) Spiraperturate (helicoidal, other arrangements) 

Blastocaulon 

Blastocaulon albidum Ruhland (19) 22 (25) (6) 7 (9) (1) 1 (1) Spiraperturate (helicoidal) 

Eriocaulon 

Eriocaulon elichrysoides Bong. (30) 33 (38) (6) 8 (11) (2) 2 (2) Spiraperturate (interapertural exine in a crosshatch pattern, other arrangements) 

Eriocaulon humboldtii Kunth (28) 32 (38) (6) 8 (10) (2) 2 (2) Spiraperturate (interapertural exine in a crosshatch pattern, other arrangements) 

Eriocaulon ligulatum (Vell.) L. B. Sm. (25) 29 (34) (5) 8 (9) (2) 2 (3) Spiraperturate (interapertural exine in a crosshatch pattern, other arrangements) 

Lachnocaulon 

Lachnocaulon minus Small (19) 22 (29) (6) 9 (13) (1) 1 (2) Spiraperturate (as the seams of a tennis ball) 

Leiothrix 

Leiothrix subgen. Leiothrix 

Leiothrix angustifolia (Körn.) Ruhland (25) 26 (31) (9) 11 (13) (1) 1 (1) Spiraperturate (helicoidal, other arrangements) 

Leiothrix distichoclada Herzog (25) 26 (30) (8) 11 (13) (1) 1 (2) Spiraperturate (helicoidal) 

Leiothrix flavescens (Bong.) Ruhland (25) 30 (34) (8) 12 (15) (1) 1 (2) Spiraperturate (as the seams of a tennis ball) 

Leiothrix hirsuta (Wikstr.) Ruhland (24) 26 (29) (8) 9 (11) (1) 1 (1) Spiraperturate (as the seams of a tennis ball) 

Leiothrix plantago (Mart.) Giul. (29) 32 (35) (9) 11 (13) (1) 1 (2) Spiraperturate (many arrangements) 

Leiothrix subgen. Rheocaulon 

Leiothrix fluitans (Mart.) Ruhland (28) 33 (39) (13) 15 (16) (1) 1 (2) Spiraperturate (as the seams of a tennis ball) 

Paepalanthus 

Paepalanthus subgen. Paepalanthus 

Paepalanthus applanatus Ruhland (29) 34 (40) (12) 13 (16) (1) 1 (2) Spiraperturate (helicoidal, other arrangements) 

Paepalanthus giganteus Sano (26) 29 (34) (10) 11 (13) (1) 1 (2) Spiraperturate (many arrangements) 

Paepalanthus pulvinathus N. E. Br. (20) 26 (35) (8) 11 (14) (1) 1 (1) Spiraperturate (helicoidal) 

Paepalanthus subtilis Miq. (19) 21 (24) (6) 7 (9) (1) 1 (1) Spiraperturate (helicoidal) 

Paepalanthus tortilis (Bong.) Mart. (19) 22 (25) (6) 9 (13) (1) 1 (1) Spiraperturate (helicoidal) 

Paepalanthus subgen. Platycaulon 

Paepalanthus bromelioides Silveira (26) 32 (39) (8) 12 (15) (1) 1 (1) Spiraperturate (helicoidal) 

Paepalanthus planifolius (Bong.) Körn. (30) 35 (40) (11) 13 (16) (1) 1 (2) Spiraperturate (many arrangements) 

Paepalanthus subgen. Xeractis 

Paepalanthus comans Silveira (31) 37 (43) (10) 13 (18) (1) 1 (2) Spiraperturate (many arrangements) 

Philodice 

Philodice hoffmannseggii Mart. (20) 25 (29) (6) 8 (13) (1) 1 (2) Spiraperturate (different arrangements) 

Rondonanthus 

Rondonanthus duidae (Gleason) Hensold & Giul. (31) 36 (41) (11) 13 (15) (2) 2 (2) Spiraperturate (helicoidal) 

Syngonanthus 

Syngonanthus sect. Syngonanthus 

Syngonanthus fuscescens Ruhland (25) 28 (31) (10) 13 (18) (1) 1 (2) Spiraperturate (many arrangements) 

Syngonanthus schwackei Ruhland 

Syngonanthus sect. Carphocephalus 

(26) 32 (40) (9) 11 (15) (1) 1 (2) Spiraperturate (many arrangements) 

Syngonanthus caulescens (Poir.) Ruhland 

Syngonanthus sect. Eulepis 


Syngonanthus bisulcatus (Körn.) Ruhland (19) 24 (31) (9) 12 (15) (1) 1 (2) 2-zonasulcate 

Syngonanthus elegans var. elegans (Körn.) Ruhland (25) 27 (33) (10) 12 (14) (1) 1 (1) 2-zonasulcate 

Syngonanthus euschemus Ruhland (25) 28 (30) (11) 13 (15) (1) 1 (2) 2-zonasulcate 

Syngonanthus giuliettiae L. R. Parra (23) 25 (29) (11) 13 (18) (1) 1 (2) 2-zonasulcate 

Syngonanthus mucugensis Giul. 

Syngonanthus sect. Thysanocephalus 

(25) 32 (39) (13) 15 (18) (1) 1 (2) 2-zonasulcate 

Syngonanthus cipoensis Ruhland (29) 32 (38) (9) 12 (14) (1) 1 (2) Spiraperturate (many arrangements) 

Syngonanthus imbricatus (Körn.) Ruhland (26) 30 (35) (13) 17 (23) (1) 1 (2) 2-zonasulcate 

Syngonanthus vernonioides (Kunth) Ruhland 

⁎Average (range), in µm. 


spiral pattern of the apertures varies according to the group of 

species being examined (Plate I,1;Plate II, 9;Plate IV, 7). 

There are pollen grains with only one spiraperture, as seen 

in Actinocephalus (Plate I, 7–9), Blastocaulon (Plate II, 7–9), 

Lachnocaulon (Plate III, 1–3), Leiothrix (Plate IV, 4–6), and Rondonanthus 

(Plate VI, 1–3). On occasion, the apertures have an 

arrangement reminiscent of the pattern on a tennis ball, such as 

that observed in Lachnocaulon (Plate III, 1–3) and Leiothrix 

(Plate III, 7–9, 12). 

93


Table 3 

Characterization of the supratectal processes on the surface of pollen grains of the studied Eriocaulaceae species 

Species (Micro)Spines Granules diam. (µm) 

Shape Wall Size (µm) 

Actinocaphalus 

Actinocephalus bongardii (A. St.-Hil.) Sano Acute and obtuse Straight and concave 0.38–0.96 0.33–0.36 

Actinocephalus cabralensis (Silveira) Sano Acute and obtuse Straight and concave 0.62–0.91 0.40–0.41 

Actinocephalus polyanthus (Bong.) Sano Acute and obtuse Straight and concave 0.37–1.05 0.39–0.44 

Actinocephalus ramosus (Wikstr.) Sano Acute Straight and concave 0.52–1.09 0.29–0.40 

Blastocaulon 

Blastocaulon albidum Ruhland Acute and obtuse Straight and concave 0.32–0.99 0.21–0.32 

Blastocaulon scirpeum (Mart.) Giul. Acute Straight 0.76–0.81 – 

Eriocaulon 

Eriocaulon elichrysoides Bong. Obtuse Straight 0.3–0.87 – 

Eriocaulon humboldtii Kunth Obtuse Straight 0.2–0.7 – 

Eriocaulon sellowianum Kunth Obtuse Straight 0.2–0.9 – 

Lachnocaulon 

Lachnocaulon engleri Ruhland Acute and obtuse Straight 0.53–0.82 0.31–0.39 

Lachnocaulon minus Small Acute and obtuse Straight 0.27–0.91 0.27–0.35 

Leiothrix 

Leiothrix subgen. Leiothrix 

Leiothrix angustifolia (Körn.) Ruhland Acute Straight 0.38–1.35 – 

Leiothrix hirsuta (Wikstr.) Ruhland Acute Straight 0.78–1.61 – 

Leiothrix plantago (Mart.) Giul. Acute Straight 0.41–1.07 – 

Leiothrix rufula (A. St. Hil.) Ruhland Base width and height equal Straight 0.47–1.23 – 

Leiothrix subgen. Stephanophyllum 

Leiothrix spiralis (Körn.) Ruhland Acute and obtuse Straight 0.35–0.82 0.22–0.32 

Leiothrix vivpara (Bong.) Ruhland Acute Straight 0.38–0.85 – 

Paepalanthus 

Paepalanthus subgen. Paepalanthus 

Paepalanthus myocephalus Mart. ex Körn. Acute and obtuse Straight 0.51–1.02 – 

Paepalanthus pulchelus Herzog Acute and obtuse Straight 0.62–1.36 0.34–0.35 

Paepalanthus pulvinathus N. E. Br. Acute Straight and convex 0.38–1.45 – 

Paepalanthus sphaerocephalus Ruhland Acute and obtuse Straight and convex 0.36–1.12 0.24–0.29 

Paepalanthus subtilis Miq. Acute and obtuse Straight 0.35–0.73 0.31–0.34 

Paepalanthus subgen. Platycaulon 

Paepalanthus planifolius (Bong.) Körn. Acute and obtuse Straight 0.35–2.09 – 

Paepalanthus subgen. Thelxinöe 

Paepalanthus leucocephalus Ruhland Acute Straight 0.88–0.96 – 

Paepalanthus scleranthus Ruhland Acute Straight 0.78–0.89 0.22–0.29 

Paepalanthus subgen. Xeractis 

Paepalanthus comans Silveira Acute and obtuse Straight and convex 0.40–1.32 0.39–0.5 

Paepalanthus chrysolepis Silveira Acute and obtuse Straight 0.56–1.97 0.38–0.42 

Paepalanthus superbus Ruhland Acute and obtuse Straight and convex 0.45–1.56 0.38–0.49 

Philodice 

Philodice hoffmannseggii Mart. Acute Straight 0.65–0.9 – 

Rondonanthus 

Rondonanthus duidae (Gleason) Hensold & Giulietti Acute and obtuse Straight 0.56–1.15 – 

Syngonanthus 


Syngonanthus arenarius (Gardn.) Ruhland Acute and obtuse Straight 0.36–1.2 – 

Syngonanthus schwackei Ruhland Acute and obtuse Straight and convex 0.43–0.89 – 

Syngonanthus verticillatus (Bong.) Ruhland Acute and obtuse Straight and convex 0.56–1.22 – 

Syngonanthus sect. Carphocephalus 

Syngonanthus caulescens (Poir.) Ruhland Acute and obtuse Straight 0.55–0.71 – 

Syngonanthus sect. Eulepis 

Syngonanthus bisulcatus (Körn.) Ruhland Acute Straight 0.63–1.47 0.38–0.44 

Syngonanthus brasiliana Giul. Acute and obtuse Straight and convex 0.32–1.05 – 

Syngonanthus giuliettiae L. R. Parra Acute and obtuse Straight 0.41–1.42 – 

Syngonanthus mucugensis Giul. Acute and obtuse Straight and convex 0.45–1.79 0.41–0.47 


Syngonanthus cipoensis Ruhland Acute and obtuse Straight 0.48–1.26 – 

Syngonanthus vernonioides (Kunth.) Ruhland Acute and obtuse Straight and convex 0.53–1.74 0.61–0.64 

Tonina 

Tonina fluviatilis Aubl. Acute and obtuse Straight (longitudinally grooved) 0.57–1.37 – 

In some species of Actinocephalus and Paepalanthus, andinthe 

genus Eriocaulon, the pollen grains can have more than one furrow 

and demonstrate different spiral patterns. Species of the genus 

Eriocaulon have pollen grains with 2–3 apertures (Plate I, 1–2), 

arranged in a characteristic pattern. In these pollen grains, interapertural 

exines are generally visible under LM, arranged in a two-

y-two formation in different plans as in a crosshatch pattern 

(Plate I, 1–2). 

In the genus Actinocephalus, A. bongardii, and A. denudatus, have 

pollen grains with a furrow in a single spiral pattern, but A. divaricatus 

and A. ramosus have two spirals in varied patterns (Plate I,10–12). 

The pollen grains of Paepalanthus have apertures that generally 

trace a single spiral (Plate IV,7–9), but do show significant variation in 

the number of spirals, while the genus Philodice has grains with 

various types of apertures in spiral form but no one predominant type 

(Plate V, 2–10). Among the species of Syngonanthus studied, pollen 

grains could be 2-zonasulcate (Plate VI, 7–9) or could have spiral 

apertures in various arrangements (Plate VII, 2–4). 

The interapertural exines are of varying widths, with the 

narrowest seen in Blastocaulon (~7 µm) and in Eriocaulon (~8 µm), 

while the widest strips were noted in Syngonanthus (up to 17 µm). 

The greater number of studied taxa has unornamented apertural 

membranes, except for the pollen grains of Eriocaulon and Syngonanthus 

arenarius. Eriocaulon are ornamented with obtuse microspines 

on this membrane (Plate I, 5), whereas the pollen grains of 

Syngonanthus arenarius have spines and microspines on the furrow 

membrane (Plate VII, 1). At the apertures of the pollen grains of 

Rondonanthus, the apertural membranes are not ornamented (Plate 

VI, 6), as is the case in Actinocephalus (Plate II, 5), Blastocaulon (Plate II, 

11), Lachnocaulon (Plate III, 4), Leiothrix (Plate IV, 3), Paepalanthus 

(Plate V, 1) and Philodice (Plate V, 11). 

3.3. Exine — structure and ectexine ornamentation 

The exine is usually thin, varying in thickness from 1 to 3 µm. The 

thickest exine was observed in specimens of the genera Eriocaulon 

and Rondonanthus, whose average exine thickness is 2 µm; in the 

other genera, the average exine thickness is 1 µm (Table 2). 

The layer stratification of the exine is discernible, with the nexine 

being thicker than the sexine in Eriocaulon and some species of Syngonanthus. 

The exine is undifferentiated in Leiothrix spp., showing no 

individualization of the layers under LM, with the exception of 

L. flavescens and L. plantago, whose sexine and nexine are quite 

distinct. The same is observed in Blastocaulon albidum. In Paepalanthus, 

the exine stratification can be either undifferentiated or 

differentiated into sexine and nexine and, in the latter case, the 

analyses show that the exine can have a thicker sexine, or that the 

sexine and nexine could be of equal thickness. 

Due to the thinness of the exine in the pollen grains of the majority 

of the species analysed, the exine layers (sexine and nexine) were not 

measured individually, although this does not prevent a qualitative 

analysis of the relative thicknesses of the two layers (Table 4). 

The ectexine surface is irregular (Plate III, 10–11), sometimes 

presenting areas with microperforations (Plate II, 2;Plate IV, 3)or 

being slightly microrugulate (Plate III,5–6,10; Plate V,1;Plate VI,11–12). 

The ectexine surface has two morphological types of supratectal 

processes: granulose and (micro)spinose (the latter have tapered 

apices). Due to their small size, both types are more easily visualized 

under SEM. In spite of the heterogeneity in their size, the granules 

appear to be more uniform than the microspines. The granules vary in 

diameter from 0.22 to 0.64 µm and are present in the majority of the 

genera studied. They are generally distributed among the microspines 

and along the edge of the interaperture exine strip in Actinocephalus 

(Plate II, 6),Lachnocaulon (Plate III, 5),Paepalanthus and Syngonanthus 

(Plate VII, 6). They were not observed in Eriocaulon, Philodice or Tonina. 

The (micro)spines show a wide range of variation in size, being 

between 0.2 and 2.09 µm in length, with a majority being acute (Plate II, 

5; Plate III, 11;Plate V, 12) while others are more obtuse (Plate I, 5; 

Plate V,1;Plate VII, 1). The acute or obtuse shape is related to the size of 

the supratectal processes, with the larger processes (spines) generally 

being acute (Plate II, 5; Plate III, 11), while the smaller processes 

(microspines) are generally obtuse (although some are occasionally 


acute) (Plate I,5;Plate VI,12;Plate VII, 1). The largest spinose processes 

are observed in Paepalanthus chrysolepis and P. planifolius (Table 3). 

4. Discussion 

4.1. Pollen morphological considerations 

The pollen grains in Eriocaulaceae present few variations with 

regard to size or shape, being small to medium and mainly spheroidal. 

Large variations in the size of the pollen grains at the intra-specific 

level were seen among the specimens studied, which could be 

attributed to the different degrees of maturation of the pollen grains 

derived from different flowers on the same inflorescence. Giulietti 

(1996), for example, reported that the maturation of the flowers in the 

inflorescences of the genus Leiothrix is centripetal, a condition that 

has been seen in the entire family. Coan et al. (2007a) pointed out that 

flowers of many different ages occur on the same inflorescence in 

Syngonanthus caulescens. 

The sulcate feature, colpate according to some authors (Furness, 1988; 

Rull, 2003), the typical apertural pattern of the family, was observed in all 

species included here. Thanikaimoni (1965) described several apertural 

arrangements for pollen grains of the Eriocaulon species, and the patterns 

seen here are very similar to those described by that author. 

Apertures having a design resembling the pattern on a tennis ball 

are quite unique, and are described here for the genera Leiothrix and 

Lachnocaulon, although Thanikaimoni (1965) also observed this 

apertural type in two different genera, Syngonanthus and Tonina, as 

well as a very similar pattern in the pollen grains of Lachnocaulon. The 

genera Actinocephalus, Blastocaulon, and Paepalanthus have very 

similar aperture patterns, with apertures in a single spiral (helicoidal) 

or tracing various designs. However, the pollen4 grains of all species of 

Syngonanthus sect. Eulepis and of S. imbricatus [S. sect. Thysanocephalus 

after Ruhland (1903) and S. sect Eulepis after Lazzari (2000)] 

have a 2-zonasulcate apertural pattern — the first time this pattern has 

been reported for this family. 

On many occasions, even under SEM, it is difficult to distinguish 

between supratectal processes to determine which are microspines 

and which are, in fact, granules (Plate I,6;Plate II,2;Plate III,6;Plate IV, 

12). The following species have pollen grains that were entirely 

microspinose: Actinocephalus cabralensis, Blastocaulon scirpeum, 

Lachnocaulon engleri, Paepalanthus leucocephalus and P. subtilis. All of 

the other taxa have exines that varied from microspinose to spinose, 

with acute spines and acute to obtuse microspines, with the exception 

of the species of Leiothrix that had both acute spines and microspines. 

The morphology of the grooved spines of Tonina fluviatilis, unique 

in the family, was very similar to those described by Halbritter et al. 

(2007) for Thymelaea passerina (L.) Coss. & Germ. (Thymelaeaceae). 

The representatives of Eriocaulon, the basal group of the family 

according to M. J. G. Andrade (unpubl. data.), generally have three 

apertures and a large apertural area. The number of apertures 

diminishes and the interapertural exine strips are wider among the 

genera of the subfamily Paepalanthoideae, which indicates a decrease in 

the apertural area. In the genus Syngonanthus, a significant decrease in 

the apertural area has been noted, mainly in the species of S. sect. Eulepis, 

and in S. imbricatus, which have zonasulcate apertures and, 

consequently, have smaller apertural areas than the others. 

4.2. Taxonomic considerations 

The contribution of pollen morphological characteristics to taxonomic 

studies of the Eriocaulaceae can be seen at many levels. Although 

the family can be considered stenopalynous, as recorded by Erdtman 

(1943, 1952), Thanikaimoni (1965), andSantos et al. (2000), the pollen 

characteristics presented here suggest that they have taxonomic value. 

The pollen characteristics of the Eriocaulaceae are sufficient to 

distinguish them from the other families of the order Poales. 

95


Spiraperturate pollen grains with a spinulose exine surface are 

especially characteristic of the family among those from the same 

order (Kuprianova, 1948; Erdtman, 1952; Raj and Saxena, 1966; 

Sharma, 1967; Maley, 1970; Melhem and Abreu, 1981; Zavada, 1983; 

Salgado-Labouriau and Rinaldi, 1990; Wanderley and Melhem, 1991; 

Harley and Zavada, 2000; Linder and Rudall, 2005). 

According to Ruhland (1903), the family includes two subfamilies: 

Eriocauloideae (Eriocaulon and Mesanthemum) with four or six 

stamens and glandular petals and Paepalanthoideae (the remaining 

genera) with two or three stamens and non-glandular petals. This 

proposal, although not corroborated by morphological analyses 

(Giulietti et al., 2000), was supported by molecular analysis (M. J. G. 

Andrade, unpubl. data) and now by our pollen data. 

Eriocaulon differs from the other genera by having interapertural 

exines arranged in a two-by-two pattern and in displaying an exine 

with obtuse microspines and an ornamented apertural membrane. 

Mesanthemum, the other genus of the subfamily Eriocauloideae, 

Plate I. Pollen grains of Eriocaulaceae. 

1–5. Eriocaulon elichrysoides Bong., 1 — high focus, 2 — optical section, 3 and 4 — general view (SEM), 5 — surface (SEM). 

6. Eriocaulon humboldtii Kunth., surface (SEM). 

7–9. Actinocephalus bongardii (A. St.-Hil.) Sano, 7 — high focus, 8 — optical section, 9 — low focus. 

10–12. Actinocephalus divaricatus (Bong.) Sano, 10 — high focus, 11 — optical section, 12 — low focus. All bars=2 µm. 

Plate II. Pollen grains of Eriocaulaceae. (see on page 98) 

shows medium-sized pollen grains (ca. 35 µm), echinate exine, and 

apertures in spiral form (see Ybert, 1979). 

Tonina and Syngonanthus stand out from the other genera within the 

subfamily Paepalanthoideae. Tonina fluviatilis, however, presented a 

unique character (grooved spines) within the Eriocaulaceae, confirming 

its position as a monotypic genus, which is now sustained by pollen data. 

The pollen grains of Syngonanthus have two differential aperture 

patterns. In S. sect. Syngonanthus (S. fuscescens and S. schwackei), S. sect. 

Carpocephalus (S. caulescens), and S. sect. Thysanocephalus (S. cipoensis 

and S. vernonioides) these are spiraperturate, while in the species of 

Syngonanthus sect. Eulepis and in S. imbricatus [S. sect. Thysanocephalus, 

which was recently proposed to be included in S. sectEulepis by 

Lazzari (2000)] they are 2-zonasulcate. This latter type of aperture, here 

discovered in Syngonanthus, is especially interesting because the genus 

forms two monophyletic groups based on morphological, molecular 

(M. J. G. Andrade, unpubl. data.) and flavonoid data (Ricci et al., 1996): 

Group 1 — Syngonanthus sect. Syngonanthus together with S. sect. 

1–2. Actinocephalus cabralensis (Silveira) Sano, 1 — general view (SEM), 2 — surface (SEM). 

3–6. Actinocephalus ramosus (Wikstr.) Sano, 3 — general view (SEM), 4–6 — surface (SEM). 7–11 — Blastocaulon albidum Ruhland,; 7 — high focus, 8 — optical section, 9 — 

low focus, 10 — general view (SEM), 11 — surface (SEM). 

12. Blastocaulon scirpeum (Mart.) Giul., surface (SEM). All bars =2 µm. 

Plate III. Pollen grains of Eriocaulaceae. (see on page 99) 

1–6 — Lachnocaulon minus Small, 1 — high focus, 2 — optical section, 3 — low focus. 

4 — general view (SEM), 5–6; surface (SEM). 

7–11. Leiothrix hirsuta (Wikstr.) Ruhland, 7 — high focus, 8 — optical section, 9 — low focus, 10 — general view (SEM), 11 — surface (SEM). 

12. Leiothrix plantago (Mart.) Giul. general view (SEM). All bars=2 µm. 

Plate IV. Pollen grains of Eriocaulaceae. (see on page 100) 

1–3. Leiothrix plantago (Mart.) Giul., 1 — surface (SEM), 2 — general view (SEM), 3 — surface (SEM). 

4–6. Leiothrix curvifolia (Bong.) Ruhland, 4 — high focus, 5 — optical section, 6 — low focus. 

7–9. Paepalanthus applanatus Ruhland, 7 — high focus, 8 — optical section, 9 — low focus; 

10. Paepalanthus pulvinathus N. E. Br., surface (SEM). 

11–12. Paepalanthus scleranthus Ruhland, 11 — general view (SEM), 12 — surface (SEM). All bars =2 µm. 

Plate V. Pollen grains of Eriocaulaceae. (see on page 101) 

1. Paepalanthus subtilis Miq., surface (SEM). 

2–12. Philodice hoffmanseggii Mart., 2 — high focus, 3 — optical section, 4 — low focus; 5 — high focus, 6 — optical section, 7 — low focus; 8 — high focus, 9 — optical section, 10 — 

low focus; 11 — general view (SEM), 12 — surface (SEM). All bars=3 µm. 

Plate VI. Pollen grains of Eriocaulaceae. (see on page 102) 

1–6. Rondonanthus duidae (Gleason) Hensold & Giul., 1 — high focus, 2 — optical section, 3 — low focus, 4 — general view (SEM), 5–6 — surface (SEM). 

7–9. Syngonanthus mucugensis Giul., 7 — high focus, 8 — optical section, 9 — low focus. 

10–11. Syngonanthus giuliettiae L. R. Parra, 10 — general view (SEM), 11 — surface (SEM). 

12. Syngonanthus schwackei Ruhland, surface (SEM). All bars =3 µm. 

Plate VII. Pollen grains of Eriocaulaceae. (see on page 103) 

1 — Syngonanthus arenarius (Gardn.) Ruhland, surface (SEM). 

2–6. Syngonanthus vernonioides (Kunth.) Ruhland, 2 — high focus, 3 — optical section, 4 — low focus, 5 — general view (SEM), 6 — surface (SEM). 

7–12. Tonina fluviatilis Aubl., 7 — high focus, 8 — optical section, 9 — low focus, 10 — general view (SEM), 11 — surface (SEM), 12 — spine (SEM). Arrow = microspine on apertural 

membrane. All bars=3 µm except Fig. 12=1 µm.


Plate I. 

97


Plate II (caption on page 96).


Plate III (caption on page 96). 

99


Plate IV (caption on page 96).


Plate V (caption on page 96). 

101


Plate VI (caption on page 96).


Plate VII (caption on page 96 ). 

103


Table 4 

Summary of pollen characteristics of Eriocaulaceae genera 

Taxa No. of 

studied 

spp. 

Carpocephalus, andGroup2— S. sect. Eulepis together with S. sect. 

Thysanocephalus. 

Thus, pollen data allied with others, raise the possibility of 

separating Syngonanthus into two distinct genera, as proposed by 

M. J. G. Andrade (unpubl. data.). The homogeneity of Syngonanthus 

sect Eulepis with 2-zonasulcate pollen grains is strong evidence 

supporting the new genus, but the position of S. sect. Thysanocephalus 

with spiraperturate pollen grains needs more study. 

According to Hensold and Giulietti (1991), the genus Rondonanthus 

appears to combine characteristics from both the Eriocauloideae and 

Paepalanthoideae subfamilies. Rondonanthus was the only representative 

of Paepalanthoideae having a thick exine, a characteristic found 

in the species of Eriocaulon. However, in this latter genus, the nexine is 

thicker than the sexine, whereas the sexine and nexine are of equal 

thickness in Rondonanthus. 

The genera Paepalanthus and Syngonanthus demonstrated the greatest 

variation in pollen characteristics, but only the zonasulcate species of 

Syngonanthus sect Eulepis can be differentiated from the others. All of 

the characteristics of Actinocephalus fit with those of Paepalanthus, as 

recorded by Santos et al. (2000) in studying species of both genera. A more 

detailed study of these pollen grains under transmission electron 

microscopy could provide important data for the taxonomy of the 

Eriocaulaceae, especially in terms of a better diagnosis of their apertures. 

5. Conclusion 

Size 

class 

Subfam. Eriocauloideae 

Eriocaulon 4 Medium Spiraperturate, 2–3 apertures; 

interapertural exine in a 

crosshatch pattern 

Subfam. Paepalanthoideae 

Actinocephalus 6 Small to 

medium 

Apertural type Exine layers under LM Spines Granules 

Spiraperturate, only one spiral 

(helicoidal) or two spirals in 

Blastocaulon 2 Small 

varied arrangement 

Spiraperturate, only one spiral 

(helicoidal) 

Lachnocaulon 2 Small Spiraperturate, only one spiral in 

tennis ball pattern 

Leiothrix 9 Medium Spiraperturate, perfect spiral 

(helicoidal), or in tennis ball 

pattern 

Paepalanthus 15 Small to 

medium 

Spiraperturate, many 

arrangements 

The stenopalynous condition stated to Eriocaulaceae family is not 

well-supported here, because pollen characteristics are usefull to the 

characterization of some taxonomic groups. This study of the diversity of 

Eriocaulaceae pollen brought to light characteristics previously unrecorded: 

the grooved spines of Tonina fluviatilis and the 2-zonasulcate 

grains of pollen of Syngonanthus sect. Eulepis. Furthermore the pollen 

grains of 26 species are described here for the first time and 41 species 

have been subjected to more detailed examination through SEM. 

SbN Obtuse microspines, regularly 

distributed 

S=N but SNN inA. denudatus Acute spines and acute and obtuse 

microspines with irregular 

distribution 

Undifferentiated Acute spines with acute and obtuse 

microspines disperse among they 

SNN Acute spines with acute (larger) and 

obtuse (smaller) microspines 

disperse among they 

Undifferentiated but S=N inL. flavescens 

and L. plantago 

Undifferentiated but SNN inP. applanatus 

and P. comans and S=N inP. bromelioides 

and P. giganteus 

Acknowledgments 

Acute (micro)spines with irregular 

distribution 

Acute spines and acute and obtuse 


distribution 

The authors wish to express their thanks to: Coordenação de 

Aperfeiçoamento de Pessoal de Nível Superior (CAPES), Conselho 

Nacional de Desenvolvimento Científico e Tecnológico (CNPq) and 

Ministério de Ciência e Tecnologia (MCT) for financial support and 

grants; IMSEAR for permitting use of the SEM; directive board of the 

herbaria HUEFS, MU, SP and SPF for providing pollen samples; 

Dr. Raymond Mervin Harley (RBG, Kew) for critical reading of the text; 

Mrs. Gisele Pinto Rocha (UEFS) for technical assistance with SEM; 

Mrs. Neuza Magalhães Moraes Santos (SISBI/UEFS) for providing copies 

of many of articles cited on this paper; Drs. Maria José Gomes de Andrade 

(UNEB), Paulo Takeo Sano (USP) e Vera Lucia Scatena (UNESP) for giving 

some specimens. Authors are deeply thankful to three anonymous 

referees of RPP for their suggestions and corrections. 

Appendix A. Specimens investigated 

Absent 

On the edge of exine 

strips and irregularly 

scattered 

Few, distributed close 

to the spines 

On the edge of exine 

strips and between 

the spines 

Absent, except for L. 

plantago (Mart.) Giul. 

Irregularly distributed 

and on the edge of 

exine strips 

Philodice 1 Small Spiraperturate, one aperture in 

different arrangements 

S=N Acute microspines Absent 

Rondonanthus 1 Medium Spiraperturate, one aperture in S=N Acute spines and acute and obtuse Rare 

varied arrangement 


distribution 

Syngonanthus 14 Small to Zonasulcate (2 apertures), S=N but undifferentiated in S. caulescens, S. Acute spines with convex walls and Irregular distributed 

medium spirapertures in many 

cipoensis, S. schwackei and SbN inS. acute and obtuse microspines and on the edge of 

arrangements 

imbricatus, S. vernonioides 

exine strips 

Tonina 1 Small⁎ Spiraperturate – Acute and obtuse microspines with 

straight walls; acute grooved spines 

Absent 

Sizes: small (≤25 µm), medium (N25 µm); S = sexine; N = nexine. ⁎Based on SEM measurements (av. 20.51 µm). 

Actinocephalus bongardii (A. St.-Hill.) Sano, A. M. Giulietti et al. 932- 

80, SPF; Actinocephalus cabralensis (Silveira) Sano, CFCR 15363, SPF; 

Actinocephalus denudatus (Körn.) Sano, CFCR 2428, SPF; Actinocephalus 

divaricatus (Bong.) Sano, CFCR 2258, SPF; Actinocephalus polyanthus 

(Bong.) Sano, CFCR 6349, SPF; Actinocephalus ramosus (Wikstr.) 

Sano, A. M. Carvalho & J. Saunders 3140, SPF. 

Blastocaulon albidum Ruhland, N. Hensold 234, HUEFS; Blastocaulon 

scirpeum (Mart.) Giul., CFCR, SPF 3006. 

Eriocaulon elichrysoides Bong., W. Monteiro 9, HUEFS; Eriocaulon 

humboldtii Kunth, Almeida & Cordeiro552, HUEFS; Eriocaulon ligulatum 

(Vell.) L. B. Sm., CFCR 1522, SPF; Eriocaulon sellowianum Kunth, 

Farias et al. 97/127, HUEFS. 

Lachnocaulon engleri Ruhland, Unwin 237, MU; Lachnocaulon 

minus Small, Unwin 230, MU. 

Leiothrix angustifolia (Körn.) Ruhland, CFCR 1427, SPF; Leiothrix 

distichoclada Herzog, W. Ganev 1964, HUEFS; Leiothrix flavescens (Bong.)

Ruhland, W. Ganev 2735, HUEFS; Leiothrix fluitans (Mart.) Ruhland, CFCR 

8317, SPF; Leiothrix hirsuta (Wikstr.) Ruhland, W. Ganev 830, HUEFS; 

Leiothrix plantago (Mart.) Giul., J. Semir et al. 2753, HUEFS; Leiothrix rufula 

(A. St. Hil.) Ruhland, CFCR 1895, SPF; Leiothrix spiralis (Körn.) Ruhland, 

CFSC 3903, SPF; Leiothrix vivipara (Bong.) Ruhland, CFCR 3802, SPF. 

Paepalanthus applanatus Ruhland, N. Hensold 600, SPF; Paepalanthus 

bromelioides Silveira, CFSC 9068, SPF; Paepalanthus chrysolepis 

Silveira, M. J. G. Andrade 531, HUEFS; Paepalanthus comans Silveira, N. 

Hensold 510, SPF; Paepalanthus giganteus Sano, CFCR 3592, SPF; Paepalanthus 

leucocephalus Ruhland, M. J. G. Andrade et al. 549, HUEFS; 

Paepalanthus myocephalus Mart. ex Körn., M. J. G. Andrade 613, 

HUEFS; Paepalanthus planifolius (Bong.) Körn., CFSC 12292, SPF; Paepalanthus 

pulchelus Herzog, M. J. G. Andrade 350, HUEFS; Paepalanthus 

pulvinathus N. E. Br., M. J. G. Andrade 106, HUEFS; Paepalanthus 

scleranthus Ruhland, M. J. G. Andrade 537, HUEFS; Paepalanthus 

sphaerocephalus Ruhland, W. Ganev 3430, HUEFS; Paepalanthus 

subtilis Miq., F. Juchum 16, HUEFS; Paepalanthus superbus Ruhland, 

A. M. Giulietti 2504, HUEFS; Paepalanthus tortilis (Bong.) Mart., HUEFS 

40087, HUEFS. 

Philodice hoffmannseggii Mart., A. M. Giulietti 122, HUEFS. 

Rondonanthus duidae (Gleason) Hensold & Giul., R. Liesner 25223, SPF. 

Syngonanthus arenarius (Gardn.) Ruhland, M. J. G. Andrade 493, 

HUEFS; Syngonanthus bisulcatus (Körn.) Ruhland, M. J. G. Andrade 494, 

HUEFS; Syngonanthus brasiliana Giul., CFCR 10199, SPF; Syngonanthus 

caulescens (Poir.) Ruhland, W. Ganev 2142, SPF; Syngonanthus 

cipoensis Ruhland, CFSC 10467, SPF; Syngonanthus elegans var. elegans 

(Körn.) Ruhland, Parra et al. 86, SPF; Syngonanthus euschemus 

Ruhland, Hatschback 51849, HUEFS; Syngonanthus fuscescens Ruhland, 

Hatschback 49593, SPF; Syngonanthus giuliettiae L. R. Parra, I. S. Santos 

03, HUEFS; Syngonanthus imbricatus (Körn.) Ruhland, Andrade Lima s. 

n, SPF; Syngonanthus mucugensis Giul., CFCR 1459, SPF; Syngonanthus 

schwackei Ruhland, T. B. Cavalcanti et al. 2906, HUEFS; Syngonanthus 

vernonioides (Kunth) Ruhland, Arbo & Becker 1640, SPF; Syngonanthus 

verticillatus (Bong.) Ruhland, Arbo et al. 4614, SPF. 

Tonina fluviatilis Aubl., M. J. G. Andrade 616, HUEFS. 

References 

APG II, 2003. An update of the Angiosperm Phylogeny Group classification for the orders 

and families of flowering plants: APG II. Botanical Journal of the Linnean Society 

141, 399–436. 

Coan, A.I., Rosa, M.M., Scatena, V.L., 2007a. Embryology and seed development of 

Syngonanthus caulescens (Poir.) Ruhland (Eriocaulaceae-Poales). Aquatic Botany 

86, 148–156. 

Coan, A.I., Rosa, M.M., Scatena, V.L., 2007b. Contribution to the embryology of Leiothrix 

fluitans (Eriocaulaceae: Poales). Aquatic Botany 87, 155–160. 

Erdtman, G., 1943. An Introduction to Pollen Analysis. Chronica Botanica Company, 

Waltham. 

Erdtman, G., 1952. Pollen Morphology and Plant Taxonomy – Angiosperms. Almqvist 

and Wiksell, Stockholm. 

Erdtman, G., 1960. The acetolysis method: a revised description. Svensk Botanisk 

Tidskrift 39, 561–564. 

Furness, C.A., 1988. The Northwest European Pollen Flora (39): Eriocaulaceae. Review of 

Palaeobotany and Palynology 57, 27–32. 

Giulietti, A.M.,1996. Leiotrix Ruhland (Eriocaulaceae) no Estado da Bahia. Sitientibus 15, 61–81. 

Giulietti, N., Giulietti, A.M., Pirani, J.R., Menezes, N.L., 1988. Estudos em sempre-vivas: 

importância do extrativismo em Minas Gerais, Brasil. Acta Botanica Brasilica 1, 

179–193. 


Giulietti, A.M., Scatena, V.L., Sano, P.T., Parra, L.R., Queiroz, L.P., Harley, R.M., Menezes, N.L., 

Iseppon, A.M.B., Salatino, A., Salatino, M.L., Vilegas, W., Santos, L.C., Ricci, C.W., Bonfim, 

M.C.P., Miranda, E.B., 2000. Multidisciplinary studies on neotropical Eriocaulaceae. In: 

Wilson, K.L., Morrison, D.A. (Eds.), Monocots: Systematics and Evolution. CSIRO, 

Melbourne, pp. 580–589. 

Halbritter, H., Weber, M., Zetter, R., Frosh-Radivo, A., Buchner, R., Hesse, M., 2007. PalDat – 

Illustrated Handbook on Pollen Terminology. Society for the Promotion of Palynological 

Research in Austria, Vienna. 

Harley, M.M., Zavada, M.S., 2000. Pollen of the monocotyledons: selecting characters for 

cladistic analysis. In: Wilson, K.L., Morrison, D.A. (Eds.), Monocots: Systematics and 

Evolution. CSIRO, Melbourne, pp. 194–211. 

Hensold, N., 1991. Revisionary studies in the Eriocaulaceae of Venezuela. Annals of 

Missouri Botanical Garden 78, 424–440. 

Hensold, N., Giulietti, A.M., 1991. Revision and redefinition of the genus Rondonanthus 

Herzog (Eriocaulaceae). Annals of Missouri Botanical Garden 78, 441–459. 

Huang, T.C., 1972. Pollen flora of Taiwan. Botany Department Press, Taiwan. 

Ikuse, M., 1956. Pollen grains of Japan. Hirokawa Publishing Co., Tokyo. 

Kuprianova, L.A., 1948. Morphologie des pollens et phylogénie des Monocotilédones. 

Communications of Komarov Institute, Academy Sciences (URSS) 1 (7), 163–262. 

Lazzari, L.R.P. 2000. Redelimitação e Revisão de Syngonanthus sect. Eulepis (Bong. ex 

Koern.) Ruhland – Eriocaulaceae. Ph.D. Thesis, São Paulo Univ., Brazil. 

Linder, H.P., Rudall, P.J., 2005. Evolutionary History of Poales. Annual Review Ecological 

Evolution Systematics 36, 107–124. 

Maley, J., 1970. Contributions à l’étude du Bassin tchadien ?. Atlas de pollens du Tchad. 

Bulletin du Jardin Botanique National de Belgique 40, 29–48. 

Melchior, H., 1964. Commelinales. In: Melchior, H. (Ed.), A. Engler's Syllabus der 

Pflanzenfamilien. Gebrüder Borntraeger, Berlin, pp. 554–556. 

Melhem, T.S., Abreu, L.C.,1981. Grãos de Pólen de Angiospermas Aquáticas. Hoehnea 9, 23–40. 

Phillips, S., 1997. Flora of Tropical East Africa: Eriocaulaceae. Royal Botanic Garden, Kew. 

Pozhidaev, A.E., 2000. Pollen variety and aperture patterning. In: Harley, M.M., Morton, 

C.M., Blackmore, S. (Eds.), Pollen and Spores: Morphology and Biology. Royal 

Botanic Gardens, Kew, pp. 205–225. 

Punt, W., Hoen, P.P., Blackmore, S., Nilsson, S., Le Thomas, A., 2007. Glossary of pollen 

and spore terminology. Review of Palaeobotany and Palynology 143, 1–81. 

Raj, B., Saxena, M.R., 1966. Pollen morphology of aquatic angiosperms. Pollen et Spores 3 

(1), 49–55. 

Ricci, C.V., Patricio, M.C.B., Salatino, M.L.F., Salatino, A., Giulietti, A.M., 1996. Flavonoids 

of Syngonanthus Ruhl. (Eriocaulaceae): taxonomic implications. Biochemical 

Systematic and Ecology 24 (6), 577–583. 

Ruhland, W., 1903. Eriocaulaceae. In: Engler, A. (Ed.), Das Pflanzenreich. 13. Heft (IV.30). 

Engelman, Leipzig. 

Rull, V., 2003. An illustrated key for the identification of pollen from Pantepuy and Gran 

Sabana (Eastern Venezuelan Guayana). Palynology 27, 99–133. 

Salgado-Labouriau, M.L., Rinaldi, M., 1990. Palynolgy of the Gramineae of the 

Venezuelan Mountains. Grana 29, 119–128. 

Sano, P.T., 2004. Actinocephalus (Körn.) Sano (Paepalanthus sect. Actinocephalus), a new 

genus of Eriocaulaceae, and other taxonomic and nomenclatural changes involving 

Paepalanthus Mart. Taxon 53 (1), 99–107. 

Santos, F.A.R., Sano, P.T., Giulietti, A.M., 2000. Pollen Morphology of Brazilian species of 

Paepalanthus Mart. (Eriocaulaceae). Revista Universidade de Guarulhos-Geociências 

V(n° especial), pp. 216–218. 

Schmidt, I.B., Figueiredo, I.B., Scariot, A., 2007. Ethnobotany and effects of harvesting on 

the population ecology of Syngonanthus nitens (Bong.) Ruhland (Eriocaulaceae), a 

NTFP from Jalapão Region, Central Brazil. Economic Botany 61 (1), 73–85. 

Sharma, M., 1967. Pollen Morphology of Indian Monocotyledons. Journal of Palynology 

1–98 (Special Volume). 

Stützel, T., 1985. Die systematische Stellung der Gattung Wurdackia (Eriocaulaceae). 

Flora 177, 335–344. 

Thanikaimoni, G., 1965. Contribution to the pollen morphology of Eriocaulaceae. Pollen 

et Spores 7 (2), 181–191. 

Wanderley, M.G.L., Melhem, T.S., 1991. Flora polínica da Reserva do Parque Estadual das 

Fontes do Ipiranga (São Paulo, Brasil). Hoehnea 18 (1), 5–42. 

Ybert, J.P., 1979. Atlas de pollens de Côte D'Ivoire. O.R.S.T.O.M., Paris. 

Zavada, M.S., 1983. Comparative Morphology of Monocot Pollen and Evolutionary 

Trends of Apertures and Wall Structures. Botanical Review 49, 331–379. 

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