ArticlePDF Available

Paepalanthus mellosilvae (Eriocaulaceae), a new species from the Mantiqueira Range in Minas Gerais, Brazil


Abstract and Figures

Paepalanthus mellosilvae is here described and illustrated as a new species of Eriocaulaceae from the Atlantic Forest domain in Minas Gerais state, Brazil. The new species is known from a single locality in the Mantiqueira Range, growing along trails through cloud forests. According to the IUCN criteria, it is suggested here as critically endangered. Paepalanthus mellosilvae is compared to P. harmsii, a morphologically similar species also growing in the Mantiqueira Range, being mainly distinguished by the presence of a rhizome, narrower leaves, more numerous scapes, narrower capitula, sparsely pilose involucral bracts, and obdeltate floral bracts. Additional comments on the taxonomy, morphology, and distribution of the species, along with illustrations are provided. The specific epithet is a tribute to Prof. Dr. Renato de Mello-Silva.
Content may be subject to copyright.
Phytotaxa 500 (3): 248–252
Copyright © 2021 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
248 Accepted by Pedro Fiaschi: 24 Apr. 2021; published: 13 May 2021
Paepalanthus mellosilvae (Eriocaulaceae), a new species from the Mantiqueira
Range in Minas Gerais, Brazil
1 Laboratório Integrado de Sistemática Vegetal, Departamento de Botânica, Universidade Federal do Rio de Janeiro, Av. Carlos Chagas
Filho 373, 21941−590, Rio de Janeiro, Brazil.;
Paepalanthus mellosilvae is here described and illustrated as a new species of Eriocaulaceae from the Atlantic Forest domain
in Minas Gerais state, Brazil. The new species is known from a single locality in the Mantiqueira Range, growing along trails
through cloud forests. According to the IUCN criteria, it is suggested here as critically endangered. Paepalanthus mellosil-
vae is compared to P. harmsii, a morphologically similar species also growing in the Mantiqueira Range, being mainly distin-
guished by the presence of a rhizome, narrower leaves, more numerous scapes, narrower capitula, sparsely pilose involucral
bracts, and obdeltate floral bracts. Additional comments on the taxonomy, morphology, and distribution of the species, along
with illustrations are provided. The specific epithet is a tribute to Prof. Dr. Renato de Mello-Silva.
Keywords: Atlantic Forest, Endemism, Poales, Taxonomy, Tribute
Paepalanthus Martius (1834: 28) nom. cons. (Eriocaulaceae) is a genus of Monocots widespread in the Neotropical
region with a few species occurring in Africa (Giulietti & Hensold 1990, Stützel 1998, Giulietti et al. 2012). Most of
its ca. 400 species occur in the Cerrado Domain, especially in elevated areas from Minas Gerais, Bahia, and Goiás
states (Giulietti & Hensold 1990, Stützel 1998, Giulietti et al. 2012), but a significant diversity is also reported for the
Atlantic Forest domain (Sano et al. 2009, BFG 2018). In the Atlantic Forest, such diversity is associated with coastal
sand dune habitats and especially with open fields in elevated areas, such as the Serra do Mar and the Mantiqueira
Range (Sano et al. 2009, Sano & Giulietti 2012, Trovó et al. 2015, BFG 2018).
The diversity of Paepalanthus from the Mantiqueira Range is poorly documented, being restricted to a few
checklists and local floras. Trovó et al. (2006) recorded four species of Eriocaulaceae from the Caparaó National Park
in the northernmost part of the Mantiqueira Range, all of them belonging to Paepalanthus. Records from the southern
part of the Mantiqueira Range are available from the Flora of São Paulo State, with eight species of Paepalanthus (Sano
& Giulietti 2012). Within the core of the Mantiqueira Range, two floristic treatments for Eriocaulaceae are available, for
the Ibitipoca State Park and for the Itatiaia National Park, recording 16 and five species of Paepalanthus, respectively
(Ferreira et al. 2011, Freitas & Trovó 2017). Trovó et al. (2015) provided a nomenclatural and taxonomic treatment
for the Eriocaulaceae from the core of the Mantiqueira Range between São Paulo and Rio de Janeiro states, recording
a total of 15 species of Paepalanthus and indicating the need of further studies for the entire Mantiqueira Range.
In this manuscript a new species of Paepalanthus from the Mantiqueira Range is described and illustrated. The
species is known from a single locality in the state of Minas Gerais, growing along trails through cloud forests. The
first collection of this species dates back to 1999, while photos of the species were recently published in social media.
The specimens first caught my attention when I was analyzing the Paepalanthus collection of the SPF herbarium and
discussing taxonomic issues with the curator and collector of the specimens, Prof. Dr. Renato de Mello-Silva, to whom
I dedicate this species.
A NEW SPECIES OF PAEPALANTHUS Phytotaxa 500 (3) © 2021 Magnolia Press 249
Materials and methods
The new species is here described according to the taxonomic species concept (Stuessy 1990), following the
recommendations of the current ICN (Turland et al. 2018). The species is described from herborized specimens
analyzed under a Leica EZ4 stereoscopic microscope with camera. The cited herbaria follow the acronyms provided by
Thiers (continuously updated) and the numbers between square brackets are herbarium barcodes. The morphological
concepts follow Radford (1974), Weberling (1989), and Stützel & Trovó (2013). A preliminary conservation status
following the IUCN (2019) criteria is suggested.
Taxonomic treatment
Paepalanthus mellosilvae Trovó, sp. nov. Fig. 1.
Paepalanthus mellosilvae differs from Paepalanthus harmsii Ruhland (1903: 216) by the presence of a rhizome, narrower leaves,
more numerous scapes, narrower capitula, sparsely pilose involucral bracts, and obdeltate floral bracts. The pilosity of the floral
parts are also rather different, being usually more densely pilose on the abaxial surface and margins of the petals and sepals of P.
Type:—BRAZIL. Minas Gerais: São Tomé das Letras, Pico do Gavião, contrafortes sudoeste. Mata Nebular. 21o37’S 44o55W. Elev.
1400m, 22 February 1999, R. Mello-Silva & al. 1620 (holotype, RB! [600436]; isotypes BHCB!, HUEFS! [83902], K!, NY!, SP!,
SPF! [135081]).
Herbs, solitary, 25.0−35.0 cm tall; rhizome present, shortly elongated, aerial stem short ca. 1.5 cm long. Leaves
persistent even when dry, chartaceous, lanceolate, multi-nerved, patent to recurved, rosulate, 12.0−16.0 × 0.5−1.0
cm, mostly glabrous with very sparse long trichomes, apex acute. Spathes 4.0−5.5 cm long, greenish, mostly glabrous
with very sparse long trichomes, apex acute. Scapes 80−130, arranged in subsequent terminal cohorts of 5−8 units,
20.0−27.0 cm long, mostly glabrous with very sparse trichomes on the distal part. Capitula 3.5−6.0 mm diam., usually
hemispherical, whitish to ochraceous; involucral bracts in 2−4 series, stramineous to light-castaneous, external just
slightly surpassing the floral disc, usually oblong with acute apex, internal deltate to obdeltate with obtuse apex, flat
to navicular, ca. 2.0 mm long, sparsely pilose with ciliate margin, glabrescent when older; receptacle flat with long
trichomes. Flowers trimerous, ca. 50 per capitulum; floral bracts obdeltate with obtuse to mucronate apex, flat to
slightly navicular, ca. 2.0 mm long, densely pilose distally, densely ciliate toward the margin. Staminate flowers ca.
3.5 mm long; pedicel ca. 0.5 mm long; sepals fused at the base, obovate, flat to navicular, castaneous, ca. 2.5 mm long,
pilose distally, margin ciliate toward the apex, apex obtuse to rounded; anthophore fleshy, elongated; corolla fused into
a tube, whitish, membranaceous, ca. 1.0 mm long; stamens ca. 1.5 mm long; carpellodes 3, papillose. Pistillate flower
ca. 2.0 mm long, sessile to sub sessile; sepals fused at the very base, narrowly oblong, castaneous, ca. 3.0 mm long,
pilose distally, margin ciliate toward the apex, apex obtuse to acute; petals linear to narrowly oblong, whitish, ca. 2.5
mm long, sparsely pilose distally, margin sparsely ciliate toward the apex, apex acute to obtuse; gynoecium ca. 3.5
mm long, stigmatic branches bifid at the apex, 3 × longer than the nectariferous branches. Fruits a loculicidal capsule.
Seeds elliptical, reddish.
Etymology:—The epithet mellosilvae is a tribute to Prof. Dr. Renato de Mello-Silva, who collected a very
complete set of specimens of this new species, which was selected as the type collection. Renato was a tenacious
botanist, an inspiring professor, and a good friend.
Distribution, Habitat, and Conservation:Paepalanthus mellosilvae is known from a single locality within the
Mantiqueira Range in Minas Gerais, Brazil. From the collection label, discussions with the first collector of the type
specimens, and photos posted on popular social media, it is clear that the species forms small, dense populations growing
along trails in shaded areas of cloud forests, sometimes exposed to sunlight. However, according to Prof. Dr. Renato de
Mello-Silva, the surrounding area of the populations from São Tomé das Letras municipality is under intense quartzitic
mining activity. Considering the few known populations, and the absence of populations in conservation units, the
species is suggested here as critically endangered according to the IUCN (2019) distribution criteria B1B2ab(i, ii, iii,
Comments:—Paepalanthus mellosilvae is provisionally placed in P. subg. Paepalanthus Ruhland (1903: 122) in
250 Phytotaxa 500 (3) © 2021 Magnolia Press
Ruhland’s (1903) classification system due to its unicapitulate scapes, involucral bracts adaxially glabrous, trimerous
flowers, and capsulate fruits. This placement is not definitive within the classification system and may need further
adjustments when a much-needed revised classification system is available (Andrade et al. 2010, Trovó et al. 2013,
Andrino et al. 2020). The species is placed along with P. harmsii, P. acuminatus Ruhland (1903: 217), and P. leiseringii
Ruhland (1903: 216), which were previously described in P. subg. Xeractis Koernicke (1863: 336) and later transferred
to P. subg. Paepalanthus (Hensold 1988). Paepalanthus harmsii and P. acuminatus emerged as sister species, nested
within a clade of species mostly occurring in the Mantiqueira Range (Andrino et al. 2020). Based on the morphological
similarities and the geographical distribution of these species, I suspect that P. mellosilvae may be related to this
FIGURE 1. Paepalanthus mellosilvae Trovó. A. Habit detail. B. Capitulum detail. C. Outer involucral bract, abaxial surface. D. Floral
bract, abaxial surface. E. Staminate flower. F. Staminate flower with opened corolla and sepals removed. G. Pistillate flower. H. Pistillate
flower with sepals removed, evidencing the gynoecium. (line drawing: Klei Sousa).
A NEW SPECIES OF PAEPALANTHUS Phytotaxa 500 (3) © 2021 Magnolia Press 251
The morphologically most similar species to Paepalanthus mellosilvae is P. harmsii, which also occurs in the
Mantiqueira Range, being restricted to Ibitipoca State Park and Serra Negra, two localities at approximately 100 km
far from the occurrence site of P. mellosilvae. Both species share the robust habit, the stramineous to light castaneous
involucral bracts just slightly surpassing the floral disc, and the shaded habitat. Paepalanthus mellosilvae is mainly
distinguished by its rhizome present (vs. usually absent), narrower leaves (0.5−1.0 vs. 0.7−2.7 cm wide), more
numerous scapes (80−130 vs. 15−50), narrower capitula (3.5−6.0 vs. 7.0−12.0 mm diam.), sparsely pilose involucral
bracts (vs. tomentose) and obdeltate floral bracts (vs. linear). The outer involucral bracts of P. harmsii are also useful
for distinction, as they are frequently greenish, while in P. mellosilvae they are stramineous to light-castaneous in all
series. Additionally, the pilosity of the floral parts are also rather different in these species, being usually more densely
pilose on the abaxial surface and margins of petals and sepals in P. mellosilvae when compared to P. harmsii.
Finally, Paepalanthus mellosilvae also resembles P. macaheensis Koernicke (1871: 311) and an unidentified
collection from Santa Rita do Jacutinga in Minas Gerais state. Paepalanthus macaheensis is a more gracile species
with usually elongated stem, being also easily differentiated by its densely packed capitula with castaneous involucral
bracts shorter than the floral disc, and gynoecium with stigmatic branches only slightly longer than the nectariferous
branches. At first glance, the unidentified specimens L. Krieger s.n. (CESJ 8900, RB 806590, SPF 151271) from Santa
Rita do Jacutinga (Minas Gerais) are morphologically similar to P. mellosilvae, being relatively smaller and more
glabrous. However, the brow and ovate, glabrous involucral bracts, and especially the linear floral bracts with sparse
trichomes are much too different to be included in the morphological variation of P. mellosilvae. Additional specimens
of this taxon would be worth analyzing to assure its identity.
I would like to thank the curators of the cited herbaria for access to the Eriocaulaceae collections. Financial support
was provided by the Alexander von Humboldt Foundation, FAPERJ (E-26/202.708/2019—JCNE) and CNPq (proc.
Andrade, M.J.G. de, Giulietti, A.M., Rapini, A., Queiroz, L.P. de, Conceição, A.d.S., Almeida, P.R.M. de & van den Berg, C. (2010)
A comprehensive phylogenetic analysis of Eriocaulaceae: Evidence from nuclear (ITS) and plastid (psbA-trnh and trnL-F) dNA
sequences. Taxon 59: 379–388.
Andrino, C.O., Sano, P.T., Inglis, P.W., Hensold, N., Costa, F.N. & Simon, M.F. (2020) Phylogenetics of Paepalanthus (Eriocaulaceae), a
diverse Neotropical monocot lineage. Botanical Journal of the Linnean Society 195 (1): 34–52.
Ferreira, C.S.A.M., Trovoì, M. & Forzza, R.C. (2011) A famiìlia Eriocaulaceae no Parque Estadual do Ibitipoca, Minas Gerais, Brasil.
Boletim de Botânica da Universidade de SaÞo Paulo 29: 19–35.
Freitas, S.N.S. & Trovoì, M. (2017) Eriocaulaceae no Parque Nacional do Itatiaia, Brasil. Rodriguésia 68: 1387–1395.
Giulietti, A.M. & Hensold, N. (1990) Padrões de distribuição geográfica dos gêneros de Eriocaulaceae. Acta Botanica Brasilica 4: 133–
Giulietti, A.M., Andrade, M.J.G., Scatena, V.L., Trovó, M., Coan, A.I., Sano, P.T., Santos, F.A.R., Borges, R.L.B. & van den Berg, C.
(2012) Molecular phylogeny, morphology and their implications for the taxonomy of Eriocaulaceae. Rodriguésia 63: 1–19.
Hensold, N. (1988) Morphology and Systematics of Paepalanthus Subgenus Xeractis (Eriocaulaceae). In: Anderson, C. (Ed.) Systematic
Botany Monographs 23. The American Society of Plant Taxonomists. Michigan, 150 pp.
IUCN (2019) Guidelines for Using the IUCN Red List Categories and Criteria. Version 14. Available from: (accessed
10 November 2020)
252 Phytotaxa 500 (3) © 2021 Magnolia Press
Koernicke, F. (1863) Eriocaulaceae. In: Martius, C.F.P. & Eichler, A.W. (Eds.) Flora brasiliensis 3 (1). Royal Typography, Munich, pp.
Koernicke, F. (1871) Symbolae ad floram Brasiliae centralis cognoscesdam. Videnskabelige Meddelelser fra den Naturhistoriske Forening
i Kjøbenhavn 20–22: 309–316.
Martius, K.F.P. (1834) Eriocaulaceae. Anales des Sciences Naturelles, Botanique 2: 25–43.
Radford, A.E., Dickson, W.C., Massey, J.R. & Bell, C.R. (1974) Vascular plant systematics. Harper & Row Pub, New York, 891 pp.
Ruhland, W. (1903) Eriocaulaceae. In: Engler, A. (Ed.) Das Pflanzenreich. Regni vegetabilis conspectus 4 heft 30. Wilhelm Engelmann,
Leipzig, 294 pp.
Sano, P.T., Echternacht, L.A., Trovó, M. & Giulietti, A.M. (2009) Eriocaulaceae. In: Stehmann, J.R., Forzza, R.C., Salino, M., Sobral,
M., Costa, D.P. & Kamino, L.H.Y. (Eds.) Plantas da Floresta Atlântica. Jardim Botânico do Rio de Janeiro, Rio de Janeiro, pp.
Sano, P.T. & Giulietti, A.M. (2012) Eriocaulaceae. In: Wanderley, M.GL., Shepherd, G.J., Melhem, T.S., Guilietti, A.M. & Martins, S.E.
(Eds.) Flora Fanerogâmica do Estado de São Paulo. Instituto de Botânica, São Paulo, pp. 173–200.
Stuessy, T.F. (1990) Plant taxonomy, the systematic evaluation of comparative data. Columbia University, Press New York, 562 pp.
Stützel, T. (1998) Eriocaulaceae. In: Kubitzki, K. (Ed.) The families and genera of vascular plants IV - flowering plants: Monocotyledons
- Alismatanae and Comelinanae (except Gramineae). Springer-Verlag, Berlin, pp. 197–207.
Stützel, T. & Trovó, M. (2013) Inflorescences in Eriocaulaceae: taxonomic relevance and practical implications. Annals of Botany 112:
Thiers, B. (2020 [continuously updated]) Index Herbariorum: A global directory of public herbaria and associated staff. New York
Botanical Garden’s Virtual Herbarium. Available from: (accessed 10 November 2020)
Trovó, M., Sano, P.T., Costa, F.N. & Giulietti, A.M. (2006) Flora Fanerogâmica do Parque Nacional do Caparaoì: Eriocaulaceae. Pabstia
17: 2–8.
Trovó, M., Andrade, M.J.G. de, Sano, P.T., Ribeiro, P.L. & van den Berg, C. (2013) Molecular phylogenetics and biogeography of
Neotropical Paepalanthoideae with emphasis on Brazilian Paepalanthus (Eriocaulaceae). Botanical Journal of the Linnean Society
171: 225–243.
Trovó, M., Echternacht, L.A., Costa, F.N., Giulietti, A.M. & Sano, P.T. (2015) Nomenclatural and taxonomic notes on Eriocaulaceae from
the Atlantic Forest, Brazil. Phytotaxa 205: 249–258.
Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, W., Hawksworth, D.L., Herendeen, P.S., Knapp, S, Kusber, W.H., Li, D.Z., Marhold,
K., May, T.W., McNeill, J., Monro, A.M., Prado, J., Price, M.J. & Smith, G.F. (2018) International Code of Nomenclature for
algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017.
Regnum Vegetabile 159. Koeltz Botanical Books, Glashütten, 254 pp.
Weberling, F. (1989) Morphology of flowers and inflorescences. Cambridge University Press, Cambridge, 423 pp.
ResearchGate has not been able to resolve any citations for this publication.
Full-text available
Paepalanthus is a diverse monocot genus with remarkable diversity distributed in the Neotropical highlands of South America. The genus comprises 410 species arranged in subgenera, sections, subsections and series. Added to this complex classification, Paepalanthus shows considerable morphological heterogeneity and includes three other genera in it, Actinocephalus (Körn.) Sano, Lachnocaulon Kunth and Tonina Aubl. A broadly sampled phylogenetic inference for the genus is still missing, precluding a better understanding of its delimitation and further studies in the group. Here we present the most comprehensive phylogenetic study for Paepalanthus to date, as well as morphological survey of characters that delimit the main lineages found. We assembled a morphologically and geographically representative sampling of Paepalanthus and associated genera comprising 356 accessions in a combined dataset of plastid (trnL-F, psbA-trnH) and nuclear (ITS, ETS) regions. Bayesian inference and maximum likelihood methods were used for phylogenetic reconstruction. We found that Paepalanthus and 16 of its 28 infrageneric categories are not monophyletic, as well as the closely related genus Actinocephalus. Thirty-six well-supported clades are recognized. Morphological characters show high levels of homoplasy, and concepts traditionally used in the classification of Paepalanthus were found to be inconsistent. We confirmed that Paepalanthus as currently circumscribed is not monophyletic and revealed several new relationships in Eriocaulaceae. To make Paepalanthus monophyletic, the genus must be re-circumscribed. These results also provide a foundation for future investigations of the diversification and evolution of flora of the Neotropical highlands of South America.
Full-text available
Resumo Os trabalhos florísticos em Eriocaulaceae estão concentrados na Cadeia do Espinhaço, o principal centro de diversidade da família. O presente trabalho teve como objetivo realizar o tratamento florístico de Eriocaulaceae no Parque Nacional do Itatiaia, contribuindo com o conhecimento da família na Serra da Mantiqueira, um importante centro de diversidade do grupo no Brasil. Com base em expedições de campo e análise de espécimes de herbário, foram encontrados quatro gêneros e nove espécies, das quais sete são endêmicas da Serra da Mantiqueira. Todas as espécies são terrestres e restritas às áreas mais secas das formações campestres da parte alta do parque, exceto Eriocaulon majusculum, que ocorre nos brejos de altitude. Fazem parte do manuscrito: uma chave de identificação para as espécies, além de descrições, comentários e documentação fotográfica. Abstract The Eriocaulaceae floristic surveys are concentrated in the Espinhaço Range, the family main center of diversity. This manuscript aimed to survey the Eriocaulaceae from the Itatiaia National Park, increasing the knowledge of the family in the Mantiqueira Range, a relevant Eriocaulaceae center diversity in Brazil. Based on field expeditions and herbarium analysis, four genera and nine species are recorded, being seven of them endemic to the Mantiqueira Range. All species are terrestrial and restricted to the open formations of the high parts of the park, except Eriocaulon majusculum, which occurs in the high altitude swamps. The manuscript contains an identification key for the species, along with descriptions, comments, and photos.
Full-text available
The diversity of Eriocaulaceae in the Neotropics is primarily concentrated in the Espinhaço Range in Brazil and in the Tepuis of Venezuela. Species richness outside of these areas, however, is significant but poorly known. Based on improved taxonomic knowledge about Eriocaulaceae in the Atlantic Forest domain in Brazil, we propose nomenclatural and taxonomic changes, and provide an identification key and an annotated list of the species of the core Serra da Mantiqueira. A total of 24 species were found, one described quite recently and two still known only from the type specimens. Lectotypes for the following names are designated: Eriocaulon majusculum, Paepalanthus exiguus, P. jordanensis, P. manicatus, and Syngonanthus caulescens.
Full-text available
Eriocaulaceae in the Ibitipoca State Park, Minas Gerais, Brazil). In Brazil, Eriocaulaceae comprise ca. 600 species and nine genera. Most of the taxonomic and morphological diversity center is concentrated in the campos rupestres from the Espinhaço Range. The Ibitipoca State Park (PEIB) is located within the Mantiqueira complex and is composed of different vegetation types. The present study is the floristic treatment of Eriocaulaceae species occurring in the PEIB. Twenty-two species were found, sixteen of Paepalanthus, two of Comanthera, two of Syngonanthus, one of Leiothrix, and one of Eriocaulon. These species occur in various habitats, but are frequently found in open grass dominated formations. An identification key, line drawings, descriptions, besides comments on morphological variation and geographical distribution are provided. Two new synonyms are proposed: Paepalanthus ibitipocencis Silveira = P. calvus Körn. e Paepalanthus orthoblepharus Silveira = P. henriquei Silveira & Ruhland.
Full-text available
Background and AimsInflorescences are thought to be of enormous taxonomic relevance; however, at the same time they are regarded as being notoriously difficult. This is partly due to the conflicting needs of floristics and evolutionary botany, but partly also due to the complicated and confusing terminology introduced by W. Troll and his school.Methods The branching patterns of representatives of the genera Eriocaulon, Syngonanthus and Paepalanthus have been studied in the field and from preserved material by scanning electron microscopy. Branching patterns and formation sequences have been analysed and documented in longitudinal schemes and diagrams. Repetitive units of different levels are detected and related to the body plans of other species of the family.Key ResultsThe repetition of very few different branching patterns on different levels of complexity may lead to highly complex inflorescences. However, terms are needed only for patterns; levels may be numbered consecutively. While complex inflorescences are often described as additions or aggregations of units, there is some evidence that complex inflorescences are often the result of fractionation of inflorescence meristems.Conclusions Precise descriptions of inflorescences useful for diagnostics and phylogenetics can be much simpler than they often are today. If complex inflorescences are the result of meristem fractionation, intermediate morphotypes cannot be expected. On the other hand, such intermediate morphotypes should occur if a complex inflorescence is formed following an aggregation pathway. Unless the repetitive patterns shown here are not correlated to complementary gene activities the inflorescences are not fully understood.
A monograph, based on field and herbarium studies, is provided for Paepalanthus subg. Xeractis, which is distinguished by involucral bracts usually exceeding the capitulum and by male corollas hairy within. Some of the species show markedly primitive characters, such as leaflike bracts, deeply divided corollas in male flowers, and sepals with stomata. On the basis of floral characters, six species are removed from the subgenus. Five species, four varieties, and one form are newly described: P. anamariae, P. clausenii, P. digitiformis, P. lanuginosus, P. revolutus, P. augustus var. picensis, P. mollis var. itambeensis, P. superbus var. gracilis, P. uncinatus var. rectus, and P. chlorocephalus f. parviflorus. In addition, three new combinations are proposed: P. argenteus var. elatus, P. calvulus, and P. superbus var. niveo-niger. Subgenus Xeractis comprises twenty-seven species, including eleven varieties and two forms. These are grouped into the newly described sections Chrysostegis, Gymnostegis, Pleurophyllon, and Xeractis, the last divided into series Albidi and Fuscati. The sections and series are distinguished on the basis of habit, leaf anatomy, involucral bract color, and floral pubescence and pigmentation. In addition, twelve putative hybrids were collected in the field or identified from herbarium material. Substantial introgression was found in one of these instances of hybridization. Subgenus Xeractis is wholly endemic to the Serra do Espinhaço of Minas Gerais, Brazil, a mountain range about 370 km long, which supports high diversity and endemism in many angiosperm taxa. Most species have a very small range, and the collective patterns of their distribution suggest a division of the Serra do Espinhaço into four floristic regions: the Diamantina plateau; the northern Serra do Cipó; the southern Serra do Cipó; and the southern Serra do Espinhaço. The species diversity is highest and the species distributions narrowest in the very poor quartzitic soils of the Serra do Cipó. Within the Serra do Cipó, the species occurring in seasonally wet habitats on shallow or poorly drained soils are the most problematic; they are characterized by reticulate variation patterns and otherwise show indications of recent diversification.
Subfamily Paepalanthoideae encompass the largest generic diversity in Eriocaulaceae. In the present study, the main goals were to infer the phylogeny of this subfamily focusing on Paepalanthus, to evaluate recent classifications and morphological characters in a phylogenetic context and to reconstruct the historical biogeography of the group. Sampling involved 94 ingroup species corresponding to all recognized genera and three outgroup species. Two molecular data sets, nuclear ribosomal internal transcribed spacer (nrITS) and plastid trnL-trnF, were analysed under parsimony and Bayesian methods. Rondonanthus is monophyletic and confirmed as sister to the remaining Paepalanthoideae. Leiothrix and Actinocephalus are each monophyletic, whereas Syngonanthus may be either monophyletic or paraphyletic with the recognition of Philodice. Four subgenera of Paepalanthus are monophyletic, but P. subgenus Paepalanthus is polyphyletic. Morphological characters used in previous classifications are assessed as putative synapomorphies for recognized genera. Some of the characters employed in defining Paepalanthus subcategories appear to have evolved multiple times, and many clades may be exclusively defined by molecular synapomorphies. Biogeographical reconstructions suggest that the current distribution patterns may be related to vicariance and a few long-distance dispersal events. Furthermore, some clades are restricted to narrow geographical areas, perhaps important as a means of conserving evolutionary processes.