Content uploaded by Paulo Labiak
Author content
All content in this area was uploaded by Paulo Labiak on Mar 13, 2017
Content may be subject to copyright.
BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research
libraries, and research funders in the common goal of maximizing access to critical research.
A New Species of Ceradenia (Polypodiaceae) from Southern Brazil
Author(s): Paulo H. Labiak and Jovani B. S. Pereira
Source: Systematic Botany, 41(4):902-905.
Published By: The American Society of Plant Taxonomists
URL: http://www.bioone.org/doi/full/10.1600/036364416X693991
BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and
environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published
by nonprofit societies, associations, museums, institutions, and presses.
Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of
BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use.
Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries
or rights and permissions requests should be directed to the individual publisher as copyright holder.
Systematic Botany (2016), 41(4): pp. 902–905
© Copyright 2016 by the American Society of Plant Taxonomists
DOI 10.1600/036364416X693991
Date of publication December 15, 2016
A New Species of Ceradenia (Polypodiaceae) from Southern Brazil
Paulo H. Labiak
1,3
and Jovani B. S. Pereira
2
1
Universidade Federal do Paraná, Depto. de Botânica, C. P. 19031, 81531-980, Curitiba, PR, Brazil.
2
Ruhr-Universität Bochum, Department of Evolution and Biodiversity of Plants, 44780 Bochum, Germany.
3
Author for correspondence (plabiak@ufpr.br)
Communicating Editor: Ricarda Riina
Abstract—A new species from the highlands of southern Brazil, Ceradenia maackii, is described and compared to its most similar species.
The new species is characterized by having small fronds, ciliate rhizome scales, petiole, axes, and laminar tissue without setae, and laminar
tissue bluish and densely covered by glandular hairs. Ceradenia maackii is epipetric and known only from “Pico do Paraná”State Park.
The new species is illustrated, and a key for Ceradenia from the Atlantic Rain Forest is also provided.
Keywords—Biodiversity, grammitid ferns, Neotropics, taxonomy.
Ceradenia L. E. Bishop is a genus of grammitid ferns with
about 70 species (Bishop 1988; Kessler and Smith 2008; Labiak
and Condack 2008; León-Parra 2012; Mostacero 2013; Parris
2005) occurring in the Neotropics, Africa, Madagascar, and
the Azores. Within the grammitid ferns, Ceradenia has been
recovered as monophyletic in several phylogenetic studies,
and sister to Zygophlebia L. E. Bishop + Enterosora Baker
(Ranker et al. 2004; Sundue 2010; Sundue et al. 2014). The
two subgenera, Ceradenia subgen. Ceradenia and Filicipecten,
originally proposed by Bishop (1988) on the basis of
morphology, are also supported as monophyletic (Sundue
2010; Sundue et al. 2010, 2014). Morphologically, Ceradenia is
easily distinguished from the other genera of grammitid ferns
by having glandular hairs that are 3-furcate, with rounded
glandular cells that produce an opaque, wax-like exudate
(Sundue et al. 2012). Another neotropical genus with
glandular hairs is Galactodenia Sundue & Labiak. However,
the glandular hairs in Galactodenia are simple, 2-celled, or
1-furcate and 3-celled (Sundue et al. 2012).
Ceradenia was established by Bishop (1988), who provided
a general description for the genus and the necessary combi-
nations. After the original publication of the genus, several
new species and new combinations have been proposed
(e.g. Bishop 1989; Kessler and Smith 2008; Labiak 2003;
Labiak and Condack 2008; León-Parra 2012; Mostacero 2013;
Parris 2002), but a comprehensive taxonomic revision is still
lacking. Nonetheless, a taxonomic treatment for the Brazilian
species was provided by Labiak and Prado (2003), and a pre-
liminary key to the Brazilian species by Labiak (2003).
The Atlantic Rain Forest of Brazil is considered one of the
major centers of diversity and endemism of neotropical ferns
and lycophytes (Tryon 1972; Prado et al. 2015). It harbors
approximately 880 species of ferns and lycophytes, 340
(∼38%) of which are endemic to this region (Prado et al.
2015). Recent collections carried out by us, along the
coastal mountains of southern Brazil, revealed a new species
that seems to be narrow endemic and represented by few
populations. Based on the presence of 3-furcate hairs, with
three rounded glandular cells that produce an opaque, wax-
like, exudate, we consider it to belong to Ceradenia. This new
species is described herein, and compared to the most simi-
lar species in the genus.
Ceradenia maackii Labiak & J. B. S. Pereira, sp. nov.—TYPE:
BRAZIL. Paraná: Município de Campina Grande do Sul,
Parque Estadual Pico Paraná, Pico Itapiroca, 25°14′S,
48°50′W, 1,600 m, 17 Feb 2008, Pereira, J. B. S. & Meyer,
M. 161 (holotype: UPCB; isotypes: B, NY).
Ceradenia maackii is characterized by having small fronds,
ciliate rhizome scales, setae absent from the petiole, axes and
laminar tissue, and laminar tissue bluish in appearance and
densely covered by glandular hairs.
Plants epipetric. Roots emerging from all sides. Rhizomes
erect, radial, symmetric, densely scaly; the scales 0.8–0.9 cm
long, golden brown, linear-lanceolate, with ciliate margins.
Fronds 6–13 × 1.8–3 cm, clustered, curved; stipes 1–3 × 0.04–
0.05 cm, dark brown, terete, conspicuously covered by whitish
hairs, lacking setae; laminae pinnatisect, elliptic to lanceo-
late, membranaceous, abruptly tapered at the base and
apex, the base with 1–2 proximal segments about the half
the length of the middle ones; rachises dark brown to dark
red, not covered by the lamina tissue and visible on both
sides; segments 8–18 × 2.5–3.5 mm, linear-deltate, oblique to
the rachises (45–70°), asymmetrical at the base, decurrent
basiscopically and cuneate acroscopically, the margins flat
or slighty revolute, apex acute, costae and veins obscure on
both surfaces, veins branched, 1-furcate, pale; indument of
whitish glandular hairs, 3-celled, on both sides of the rachises,
costae, veins, and laminar tissue. Sori round, arising at the
vein tips, not protruding beyond the lamina margins when
mature, with glandular paraphyses of enlarged global cells
similar to the hairs of the laminar tissue. Figures 1, 2, and 3.
Additional Specimens Examined —BRAZIL. Paraná: Campina Grande
do Sul. Parque Estadual Pico Paraná, Pico Itapiroca, 25°15′S, 48°50′W,
1,500–1,800 m, 09 Jul 2008, Labiak et al. 4788 (UPCB).
Etymology—The specific epithet honors the German geol-
ogist Reinhard Maack, for his valuable contributions to the
knowledge of the Natural History of Paraná State. He was
also the first naturalist to reach the summit of “Pico Paraná”
in 1942, the type locality of Ceradenia maackii.
Distribution and Ecology—Ceradenia maackii is epipetric,
growing among lichens and bryophytes in shaded and humid
boulders, above the tree line (Fig. 2). It is only known from two
populations in the type locality, at elevations about 1,600 m.
Comments—Ceradenia maackii is characterized by having
rhizome scales 0.8–0.9 cm long, with ciliate margins. The
fronds lack setae, and the indument is composed of whitish
glandular hairs on both laminar surfaces (Fig. 1B). Addition-
ally, the rachises are visible on both sides of the laminae. The
most similar species are C. albidula (Baker) L. E. Bishop and
C. itatiaiensis Labiak & Condack, which differ by having
entire rhizome scales (or with only short, capitate glands
902
Fig. 2. A–D. Habitat and habit of Ceradenia maackii. A. Overview of “Pico Paraná”State Park, in the Serra do Mar mountains. B. Cloud forest
and rock boulders, habitat of C. maackii. C. Population of C. maackii growing on rock with lichens and decomposing bryophytes. D. Abaxial surface of
the fronds, with sori (older leaves remain attached to the rhizome). (Photos by P. Labiak).
Fig. 1. A–E. Ceradenia maackii. A. Habit. B. Adaxial surface of the lamina showing the glandular hairs. C. Abaxial surface of the lamina showing the
sori and the glandular hairs. D. Detail of the glandular hair. E. Rhizome scale. (All from the holotype).
2016] LABIAK AND PEREIRA: CERADENIA FROM SOUTHERN BRAZIL 903
Fig. 3. Silhouettes of three species of Ceradenia subgen. Ceradenia from southeastern Brazil. A. C. albidula (Hirai & Prado 625, NY). B. C. itatiaiensis
(Condack & Cortines 308, RB). C. C. maackii (from the holotype).
904 SYSTEMATIC BOTANY [Volume 41
along the margins), and rachises that are covered by a green
laminar tissue on both sides.
Ceradenia maackii belongs to Ceradenia subg. Ceradenia, and
it is the sixth species in this subgenus reported from Brazil.
A key to distinguish it from the other species of Ceradenia
from the Atlantic Rain Forest is provided.
Conservation Status—Because Ceradenia maackii is cur-
rently known from two populations, and only from a few
individuals, it may deserve special attention concerning its
conservation status. However, based on our current knowl-
edge on this species, and according to IUCN red list criteria
(IUCN 2010), it is assessed here as data deficient (DD).
Key to the Species of CERADENIA from the Atlantic Rain Forest of Brazil
1. Rhizomes dorsiventral; lamina deltate, basal segments longer than the median ones . . . . . . . . . . . . . . . . . . C. spixiana (Mart. ex Mett.) L. E. Bishop
1. Rhizomes radial; lamina lanceolate to linear lanceolate, basal segments equal or shorter than the median ones . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Rachis blackish to reddish, visible on both sides of the lamina and not covered by the laminar tissue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Rhizome scales ca. 2 mm long; margins of the segments undulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. capillaris (Desv.) L. E. Bishop
3. Rhizome scales 8–9 mm long; margins of the segments straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. maackii Labiak & J. B. S. Pereira
2. Rachis green, not visible, covered by the laminar tissue on both sides of the lamina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Laminaupto5cmlongand1cmwide .................................................... C. pruinosa (Maxon) L. E. Bishop
4. Laminamorethan5cmlongand1cmwide ......................................................................... 5
5. Rachis with golden brown setae on the abaxial side; sinus “U”shaped, broader than
the segment width (except in the upper third) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. glaziovii (Baker) Labiak
5. Rachis without setae; sinus “V”shaped, equal or narrower than the segments width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Lamina strongly coriaceous, with crenate margins; sori protruding beyond lamina margins . . . . . . . C. warmingii (C. Chr.) Labiak
6. Lamina chartaceous, with entire margins; sori not protruding beyond lamina margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. Segments perpendicular to the rachis, ca. 80–90°; basal segments of mature fronds
strongly reduced to auricles, up to ⅛shorter than the median ones . . . . . . . . . . . . . . . . . . . . C. albidula (Baker) L. E. Bishop
7. Segments oblique to the rachis, ca. 45–60°; basal segments of mature fronds
reduced to ca. ½ to ⅓thelengthofthemedianones ............................... C. itatiaiensis Labiak & Condack
Acknowledgments. This research was partially funded by grants
from CNPq to PHL (304835/2013-7), and to JBSP (245951/2012-1). We
thank Diana Carneiro for preparing the line drawings, and two
anonymous reviewers for helpful comments on the manuscript.
Literature Cited
Bishop, L. E. 1988. Ceradenia, a new genus of Grammitidaceae. American
Fern Journal 78: 1–5.
Bishop, L. E. 1989. New species of Ceradenia subg. Ceradenia. American
Fern Journal 79: 14–25.
IUCN. 2010. IUCN red list categories and criteria. Version 8.1. <http://
intranet.iucn.org/webfiles/doc/SSC/RedList/RedListGuidelines
.pdf.>. Downloaded August 2015.
Kessler, M. and A. R. Smith. 2008. New species of grammitid ferns (Poly-
podiaceae, Polypodiopsida) from Bolivia. Organisms Diversity and
Evolution 8: 167e1–167e18.
Labiak, P. H. 2003. A new combination in the fern genus Ceradenia
(Grammitidaceae). Kew Bulletin 58: 991–994.
Labiak, P. H. and J. P. S. Condack. 2008. Ceradenia itatiaiensis, a new spe-
cies of Polypodiaceae from southeastern Brazil. Novon 18: 495–497.
Labiak, P. H. and J. Prado. 2003. Grammitidaceae (Pteridophyta) no
Brasil com ênfase nos gêneros Ceradenia, Cochlidium eGrammitis.
Hoehnea 30: 243–282.
León-Parra, L. 2012. Ceradenia (Polypodiaceae). Pp. 1–83 in Flora de
Colombia 29, eds. J. Betancourt, G. Galeano, J. Aguirre-C. and
G. Amat-G. Bogotá, D. C., Colombia: Universidad Nacional de
Colombia.
Mostacero, J. 2013. The identity of the type of Polypodium gamerianum
Vareschi (Polypodiaceae). American Fern Journal 103: 53–56.
Parris, B. S. 2002. New species and new combinations in African
Grammitidaceae (Filicales). Kew Bulletin 57: 423–434.
Parris, B. S. 2005. Grammitidaceae. Pp. 1–21 in Flora of Tropical East
Africa, ed. H. J. Beentje. Richmond, U. K.: Royal Botanic Gardens.
Prado, J., L. S. Sylvestre, P. H. Labiak, P. G. Windisch, A. Salino, I. C. L.
Barros, R. Y. Hirai, T. E. Almeida, A. C. P. Santiago, M. A. Kieling-
Rubio, A. F. N. Pereira, B. Ollgaard, C. G. V. Ramos, J. T. Mickel,
V. A. O. Dittrich, C. M. Mynssen, P. B. Schwartsburd, J. P. S.
Condack, J. B. S. Pereira, and F. B. Matos. 2015. Diversity of ferns
and lycophytes in Brazil. Rodriguesia 66: 1–11.
Ranker, T. A., A. R. Smith, B. S. Parris, J. M. O. Geiger, C. H. Haufler,
S. C. K. Straub, and H. Schneider. 2004. Phylogeny and evolution
of grammitid ferns (Grammitidaceae): A case of rampant morpho-
logical homoplasy. Taxon 53: 415–428.
Sundue, M. A. 2010. A morphological cladistic analysis of Terpsichore
(Polypodiaceae). Systematic Botany 35: 716–729.
Sundue, M. A., M. B. Islam, and T. A. Ranker. 2010. Systematics of
grammitid ferns (Polypodiaceae): using morphology and plastid
sequence data to resolve the circumscriptions of Melpomene and
the polyphyletic genera Lellingeria and Terpsichore. Systematic Botany
35: 701–715.
Sundue, M., P. H. Labiak, J. Mostacero, and A. R. Smith. 2012.
Galactodenia, a new genus of grammitid ferns segregated from
Terpsichore (Polypodiaceae). Systematic Botany 37: 339–346.
Sundue, M. A., B. S. Parris, T. A. Ranker, A. R. Smith, E. L. Fujimoto,
D. Zamora-Crosby, C. W. Morden, W.-L. Chiou, C.-W. Chen, G.
Rouhan, R. Y. Hirai, and J. Prado. 2014. Global phylogeny and
biogeography of grammitid ferns (Polypodiaceae). Molecular Phylo-
genetics and Evolution 81: 195–206.
Tryon, R. M. 1972. Endemic areas and speciation in tropical American
ferns. Biotropica 4: 76–84.
2016] LABIAK AND PEREIRA: CERADENIA FROM SOUTHERN BRAZIL 905