Comparative pollen morphology and taxonomic ... - CNCFlora
Comparative pollen morphology and taxonomic ... - CNCFlora
Comparative pollen morphology and taxonomic ... - CNCFlora
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104 R.L.B. de Borges et al. / Review of Palaeobotany <strong>and</strong> Palynology 154 (2009) 91–105<br />
Table 4<br />
Summary of <strong>pollen</strong> characteristics of Eriocaulaceae genera<br />
Taxa No. of<br />
studied<br />
spp.<br />
Carpocephalus, <strong>and</strong>Group2— S. sect. Eulepis together with S. sect.<br />
Thysanocephalus.<br />
Thus, <strong>pollen</strong> data allied with others, raise the possibility of<br />
separating Syngonanthus into two distinct genera, as proposed by<br />
M. J. G. Andrade (unpubl. data.). The homogeneity of Syngonanthus<br />
sect Eulepis with 2-zonasulcate <strong>pollen</strong> grains is strong evidence<br />
supporting the new genus, but the position of S. sect. Thysanocephalus<br />
with spiraperturate <strong>pollen</strong> grains needs more study.<br />
According to Hensold <strong>and</strong> Giulietti (1991), the genus Rondonanthus<br />
appears to combine characteristics from both the Eriocauloideae <strong>and</strong><br />
Paepalanthoideae subfamilies. Rondonanthus was the only representative<br />
of Paepalanthoideae having a thick exine, a characteristic found<br />
in the species of Eriocaulon. However, in this latter genus, the nexine is<br />
thicker than the sexine, whereas the sexine <strong>and</strong> nexine are of equal<br />
thickness in Rondonanthus.<br />
The genera Paepalanthus <strong>and</strong> Syngonanthus demonstrated the greatest<br />
variation in <strong>pollen</strong> characteristics, but only the zonasulcate species of<br />
Syngonanthus sect Eulepis can be differentiated from the others. All of<br />
the characteristics of Actinocephalus fit with those of Paepalanthus, as<br />
recorded by Santos et al. (2000) in studying species of both genera. A more<br />
detailed study of these <strong>pollen</strong> grains under transmission electron<br />
microscopy could provide important data for the taxonomy of the<br />
Eriocaulaceae, especially in terms of a better diagnosis of their apertures.<br />
5. Conclusion<br />
Size<br />
class<br />
Subfam. Eriocauloideae<br />
Eriocaulon 4 Medium Spiraperturate, 2–3 apertures;<br />
interapertural exine in a<br />
crosshatch pattern<br />
Subfam. Paepalanthoideae<br />
Actinocephalus 6 Small to<br />
medium<br />
Apertural type Exine layers under LM Spines Granules<br />
Spiraperturate, only one spiral<br />
(helicoidal) or two spirals in<br />
Blastocaulon 2 Small<br />
varied arrangement<br />
Spiraperturate, only one spiral<br />
(helicoidal)<br />
Lachnocaulon 2 Small Spiraperturate, only one spiral in<br />
tennis ball pattern<br />
Leiothrix 9 Medium Spiraperturate, perfect spiral<br />
(helicoidal), or in tennis ball<br />
pattern<br />
Paepalanthus 15 Small to<br />
medium<br />
Spiraperturate, many<br />
arrangements<br />
The stenopalynous condition stated to Eriocaulaceae family is not<br />
well-supported here, because <strong>pollen</strong> characteristics are usefull to the<br />
characterization of some <strong>taxonomic</strong> groups. This study of the diversity of<br />
Eriocaulaceae <strong>pollen</strong> brought to light characteristics previously unrecorded:<br />
the grooved spines of Tonina fluviatilis <strong>and</strong> the 2-zonasulcate<br />
grains of <strong>pollen</strong> of Syngonanthus sect. Eulepis. Furthermore the <strong>pollen</strong><br />
grains of 26 species are described here for the first time <strong>and</strong> 41 species<br />
have been subjected to more detailed examination through SEM.<br />
SbN Obtuse microspines, regularly<br />
distributed<br />
S=N but SNN inA. denudatus Acute spines <strong>and</strong> acute <strong>and</strong> obtuse<br />
microspines with irregular<br />
distribution<br />
Undifferentiated Acute spines with acute <strong>and</strong> obtuse<br />
microspines disperse among they<br />
SNN Acute spines with acute (larger) <strong>and</strong><br />
obtuse (smaller) microspines<br />
disperse among they<br />
Undifferentiated but S=N inL. flavescens<br />
<strong>and</strong> L. plantago<br />
Undifferentiated but SNN inP. applanatus<br />
<strong>and</strong> P. comans <strong>and</strong> S=N inP. bromelioides<br />
<strong>and</strong> P. giganteus<br />
Acknowledgments<br />
Acute (micro)spines with irregular<br />
distribution<br />
Acute spines <strong>and</strong> acute <strong>and</strong> obtuse<br />
microspines with irregular<br />
distribution<br />
The authors wish to express their thanks to: Coordenação de<br />
Aperfeiçoamento de Pessoal de Nível Superior (CAPES), Conselho<br />
Nacional de Desenvolvimento Científico e Tecnológico (CNPq) <strong>and</strong><br />
Ministério de Ciência e Tecnologia (MCT) for financial support <strong>and</strong><br />
grants; IMSEAR for permitting use of the SEM; directive board of the<br />
herbaria HUEFS, MU, SP <strong>and</strong> SPF for providing <strong>pollen</strong> samples;<br />
Dr. Raymond Mervin Harley (RBG, Kew) for critical reading of the text;<br />
Mrs. Gisele Pinto Rocha (UEFS) for technical assistance with SEM;<br />
Mrs. Neuza Magalhães Moraes Santos (SISBI/UEFS) for providing copies<br />
of many of articles cited on this paper; Drs. Maria José Gomes de Andrade<br />
(UNEB), Paulo Takeo Sano (USP) e Vera Lucia Scatena (UNESP) for giving<br />
some specimens. Authors are deeply thankful to three anonymous<br />
referees of RPP for their suggestions <strong>and</strong> corrections.<br />
Appendix A. Specimens investigated<br />
Absent<br />
On the edge of exine<br />
strips <strong>and</strong> irregularly<br />
scattered<br />
Few, distributed close<br />
to the spines<br />
On the edge of exine<br />
strips <strong>and</strong> between<br />
the spines<br />
Absent, except for L.<br />
plantago (Mart.) Giul.<br />
Irregularly distributed<br />
<strong>and</strong> on the edge of<br />
exine strips<br />
Philodice 1 Small Spiraperturate, one aperture in<br />
different arrangements<br />
S=N Acute microspines Absent<br />
Rondonanthus 1 Medium Spiraperturate, one aperture in S=N Acute spines <strong>and</strong> acute <strong>and</strong> obtuse Rare<br />
varied arrangement<br />
microspines with irregular<br />
distribution<br />
Syngonanthus 14 Small to Zonasulcate (2 apertures), S=N but undifferentiated in S. caulescens, S. Acute spines with convex walls <strong>and</strong> Irregular distributed<br />
medium spirapertures in many<br />
cipoensis, S. schwackei <strong>and</strong> SbN inS. acute <strong>and</strong> obtuse microspines <strong>and</strong> on the edge of<br />
arrangements<br />
imbricatus, S. vernonioides<br />
exine strips<br />
Tonina 1 Small⁎ Spiraperturate – Acute <strong>and</strong> obtuse microspines with<br />
straight walls; acute grooved spines<br />
Absent<br />
Sizes: small (≤25 µm), medium (N25 µm); S = sexine; N = nexine. ⁎Based on SEM measurements (av. 20.51 µm).<br />
Actinocephalus bongardii (A. St.-Hill.) Sano, A. M. Giulietti et al. 932-<br />
80, SPF; Actinocephalus cabralensis (Silveira) Sano, CFCR 15363, SPF;<br />
Actinocephalus denudatus (Körn.) Sano, CFCR 2428, SPF; Actinocephalus<br />
divaricatus (Bong.) Sano, CFCR 2258, SPF; Actinocephalus polyanthus<br />
(Bong.) Sano, CFCR 6349, SPF; Actinocephalus ramosus (Wikstr.)<br />
Sano, A. M. Carvalho & J. Saunders 3140, SPF.<br />
Blastocaulon albidum Ruhl<strong>and</strong>, N. Hensold 234, HUEFS; Blastocaulon<br />
scirpeum (Mart.) Giul., CFCR, SPF 3006.<br />
Eriocaulon elichrysoides Bong., W. Monteiro 9, HUEFS; Eriocaulon<br />
humboldtii Kunth, Almeida & Cordeiro552, HUEFS; Eriocaulon ligulatum<br />
(Vell.) L. B. Sm., CFCR 1522, SPF; Eriocaulon sellowianum Kunth,<br />
Farias et al. 97/127, HUEFS.<br />
Lachnocaulon engleri Ruhl<strong>and</strong>, Unwin 237, MU; Lachnocaulon<br />
minus Small, Unwin 230, MU.<br />
Leiothrix angustifolia (Körn.) Ruhl<strong>and</strong>, CFCR 1427, SPF; Leiothrix<br />
distichoclada Herzog, W. Ganev 1964, HUEFS; Leiothrix flavescens (Bong.)