new aPProacheS in the fern genuS PteriS

Pteridaceae) from brazil

(

nuevoS enfoqueS en el género PteriS (Pteridaceae) de braSil

1

2

1

& Jeﬀerson Prado

Regina Y. Hirai * , Claudine M. Mynssen

Summary

Background and aims: Pteris is widely distributed with ca. 65 species in the Tropics.

In Brazil, 24 species were recognized and ﬁve of them are endemic to the Atlantic

Forest. The objective of the present paper is to re-evaluate the taxonomy of

Pteris occurring in Brazil, based on new morphological evidence of old and recent

specimens deposited in herbaria, analysis of types, and observations in the ﬁeld.

M&M: The new approaches are based on morphological data such as SEM images

of the spores and from the examination of herbarium collections mainly from SP

and RB Herbaria.

1

(

.Instituto de Pesquisas Ambientais

IPA), São Paulo, Brazil

. Instituto de Pesquisas Jardim

2

Botânico do Rio de Janeiro, Rio de

Janeiro, Brazil

*regina.hirai@gmail.com

Results: Four species are being recognized in the present paper (P. lata, P.

organensis, P. praealta, and P. sericea). For two of them, two new combinations

into Pteris are presented: P. organensis and P. praealta. A key to distinguishing the

newly recognized species and its relative taxa, as well as descriptions, illustrations,

and comments are presented in this paper.

Citar este artículo

HIRAI, R. Y., C. M. MYNSSEN & J.

PRADO. 2024. New approaches in

the fern genus Pteris (Pteridaceae)

from Brazil. Bol. Soc. Argent. Bot.

Conclusions: By reviewing herbarium materials, mainly observing the morphology of

the spores, and collecting more specimens in the ﬁeld, it was possible to recognize

three species (Pteris lata, P. organensis, and P. praealta) in Brazil and also verify

the correct application of the name Pteris sericea to the Brazilian plants. All these

names had been previously synonymized in previous studies. After this study, 27

species of Pteris are recognized in Brazil.

5

9: 259-272.

DOI: https://doi.

org/10.31055/1851.2372.v59.

n3.44498

Key wordS

Brazilian Atlantic Forest, endemic species, fern, morphology, palynology, taxonomy.

reSumen

Introducción y objetivos: Pteris está ampliamente distribuido con ca. 65 especies

en los trópicos. En Brasil fueron reconocidas 24 especies y cinco de ellas son

endémicas de la Mata Atlántica Brasileña. El objetivo del presente artículo es

reevaluar la taxonomía de Pteris presentes en Brasil, con base en nueva evidencia

morfológica obtenida de especímenes antiguos y recientes depositados en

herbarios, análisis de tipos y observaciones de campo.

M&M: Los nuevos enfoques se basan en datos morfológicos, como imágenes SEM

de las esporas, y en el examen de colecciones de herbarios, principalmente de SP

y RB Herbaria.

Resultados: En el presente artículo se reconocen cuatro especies (P. lata, P.

organensis, P. praealta y P. sericea). Para dos de ellos se presentan dos nuevas

combinaciones en Pteris: P. organensis y P. praealta. En este artículo se presenta

una clave para distinguir las especies recientemente reconocidas y sus taxones

relacionados, así como descripciones, ilustraciones y comentarios.

Conclusiones: Al revisar los materiales de los herbarios, observando principalmente

la morfología de las esporas y recolectando más especímenes en el campo, fue

posible reconocer tres especies (Pteris lata, P. organensis y P. praealta) en Brasil

y también veriﬁcar la correcta aplicación del nombre Pteris sericea a las plantas

brasileñas. Todos estos nombres habían sido previamente sinonimizados en

estudios anteriores. Después de este estudio, se reconocen 27 especies de Pteris

en Brasil.

Recibido: 8 Mar 2024

Aceptado: 1 Jul 2024

Publicado en línea: 30 Sep 2024

Publicado impreso: 30 Sep 2024

Editora: Olga G. Martínez

PalabraS clave

Especie endémica, helecho, Mata Atlántica Brasileña, morfología, palinología,

taxonomía.

ISSN versión impresa 0373-580X

ISSN versión on-line 1851-2372

259

Bol. Soc. Argent. Bot. 59 (3) 2024

introduction

and Mesoamerica, took into account the cingulum,

presence of a commissural ﬂange, types of macro-

The fern genus Pteris L. is monophyletic and ornamentation, ornamentation on distal and proximal

belongs to the Pteridaceae and subfamily Pteridoideae faces, and spore size.

(Prado et al., 2007; Schuettpelz et al., 2007; Zhang

Chao & Huang (2018) mapped seven Pteris

et al., 2015; PPG I, 2016). The genus is widely spore characters (equatorial ﬂange, laesural ridges,

distributed in the tropics but can reach subtropical and proximal ridges, distal ridges, tubercules on the distal

temperate regions. Some species are widely cultivated faces, coarse reticules on the distal faces, and a row

as ornamentals (e.g., P. multiﬁda Poir. and P. cretica of extervermiculi between the distal face and the

L.; Walker, 1970) and may occur as naturalized in equatorial ﬂange) onto a reconstructed phylogenetic

some places as Brazil (Prado & Hirai, 2024).

tree based on previous data published by Prado et al.

Pteris consists of ca. 250 species (PPG I, 2016) and (2007), Chao et al. (2014), and Zhang et al. (2015).

is characterized by having sporangia borne at lamina Chao & Huang (2018) studied 57 spores of Pteris

margins on a commissural vein connecting all veins and combined them with data from the literature,

and indusia strongly diﬀerentiated from the recurved totaling 100 species. In this study, there were some

margin (Prado & Windisch, 2000). The spores are species of Pteris from Brazil, whose sequences came

trilete, tetrahedral, or globose, with an equatorial from Prado et al. (2007), but the spores were not

ﬂange and with surface rugose, rarely granulate or observed, and this information appears as missing

reticulate (Tryon & Tryon, 1982; Tryon & Lugardon, data, except for P. denticulata whose spore’s data

1

990).

According to the previous studies involving Pteris results of Chao & Huang (2018) showed that most of

in Brazil, 24 species and two varieties were recognized these characters evolved independently several times

Prado & Windisch, 2000; Prado et al., 2015; Prado in diﬀerent lineages and, therefore, spore characters

Hirai, 2024) and ﬁve species were recognized as need to be combined with leaf characters to help in the

endemic to the Brazilian Atlantic Forest: P. angustata infrageneric taxonomy of Pteris.

Fée) C.V. Morton, P. congesta J. Prado, P. leptophylla Based on the above recent evidence of the

Sw., P. limae Brade, and P. schwackeana Christ (Prado importance of some morphological characters, the

Hirai, 2024). present paper proposes to re-evaluate the taxonomy

Chao et al. (2014) and Zhang et al. (2015) of Pteris occurring in Brazil. The new approaches

were used from Martínez & Morbelli (2009). The

(

&

(

&

conducted molecular phylogenetic studies showing here presented are based on new morphological data

relationships within Pteris. The ﬁrst carried out a plus the recent observations of specimens in the ﬁeld,

study with 135 species of Pteris (ca. 54% of the herbaria specimens, as well as on the type specimens

diversity of genus) from six continents, based on of the newly adopted names.

two cpDNA coding regions (rbcL and matK), to

reconstruct the phylogeny of Pteris plus the evolution

of the leaf morphology and ecological characters, also materialS and methodS

to understand the expansion and diversiﬁcation of this

cosmopolitan genus. On the other hand, Zhang et al. Taxonomic studies

(2015) studied six plastid loci of 119 species of Pteris

For this study, we analyzed specimens and types

and 18 related genera. The results obtained by Chao from the following herbaria: BHCB, BM, FCAB,

et al. (2014) with uncertain relationships between HB, HRCB, IPA, K, M, MBM, NY, P, PACA, R,

Actiniopteris Link, Onychium Kaulf., and Pteris were RB, S, SP, SPF, TAIF, UEC, UPCB, UPS, US,

clariﬁed by Zhang et al. (2015), and both found clades and VT. We also observed online collections from

well-supported within Pteris (11 clades by Chao et al., other herbaria, which are available at REFLORA

2014 versus 15 clades by Zhang et al., 2015).

(http://reflora.jbrj.gov.br/jabot/herbarioVirtual/

Recently, studies with Pteris spores have shown ConsultaPublico HVUC/ConsultaPublicoHVUC.

that they can help with the infrageneric taxonomy do), JStor Global Plants (https://plants.jstor.

of the group, e.g., Palacios-Rios et al. (2017) and org/), speciesLink (https://specieslink.net), and

Chao & Huang (2018). The ﬁrst study presented six Smithsonian National Museum of Natural History

spore patterns of 25 species of Pteris from Mexico (http://collections.nmnh.si.edu/search/botany/)

260

R. Y. Hirai et al. - New approaches in Pteris from Brazil

websites, especially types. The last author of this reSultS and diScuSSion

work studied almost all types and specimens from

many herbaria during his work published in 2000

Prado & Windisch, 2000).

Four species are being recognized in the present

study: Pteris lata, P. organensis, P. praealta, and P.

(

We personally studied living populations of Pteris sericea (Figs. 1-4). They are morphologically related

deﬂexa Link, P. lata Kaulf., P. organensis (Fée) J. mainly to Pteris deﬂexa, P. lechleri, and P. splendens.

Prado & R.Y. Hirai, P. sericea (Fée) Christ, and P. All ﬁrst three species recognized species are endemic

splendens Kaulf. Field trips were carried out from to the Brazilian Atlantic Forest and occur only along

2014 to 2016 in the following localities: Pernambuco the east coast of Brazil, except P. sericea, which can

state (November 2014); Serra da Bocaina, State of São also occur in northeastern Argentina.

Paulo (February 2015); Angra dos Reis, State of Rio

All these four taxa had been previously

de Janeiro (May 2015); several localities in Paraná and synonymized in the previous study by Prado

Santa Catarina States (June 2016); several places in the & Windisch (2000), and Prado & Hirai (2024)

States of Bahia, Minas Gerais, and Espírito Santo (July followed these concepts. With this update, 27

2016); and the Serra da Bocaina and Itatiaia Mountain, species of Pteris are now recognized in Brazil.

both in the State of Rio de Janeiro (November 2016).

Despite searches in Bahia State (Una region) and Rio

de Janeiro State (Angra dos Reis), it was not possible

to ﬁnd P. praealta in the ﬁeld.

Key to the segregated species (appear in bold)

of Pteris from Brazil and its relatives

The following additional material of Pteris were

illustrated and commented in the species discussion:

P. deﬂexa: BRAZIL. Paraná: Quatro Barras, Estrada

Velha da Graciosa, Oratório Anjo da Guarda, 25° 20’

1

. Venation completely areolate

2

. Rhizome long-creeping.

P. praealta

49.8” S, 48° 57’ 52” W, 860 m elevation, 6-VI-2016,

2

. Rhizome erect to suberect

Hirai & Prado 788 (SP); Idem: Coronel Vivida, Rod.

PR-367, 20-II-1971, Hatschbach 26371 (HB); P.

lechleri: BOLIVIA. La Paz: Abel Iturralde, Parque

Nacional Madidi, campamento de guardaparques

Sadiri, camino Sadiri-Tumupasa, por la senda a

las antenas de Entel, 14° 10’ S, 67° 53’ W, 990 m

elevation, 7-VII-2004, Jiménez & Huaylla 2659 (SP),

and P. splendens: BRAZIL. São Paulo: São Paulo,

Parque Estadual das Fontes do Ipiranga (PEFI), trilha

que margeia o Zoológico, 20-XII-2005, Prado & Silva

3

Sterile pinnae widely oblong (over 2.5 cm

wide).

P. splendens

. Sterile pinnae lanceolate.

P. organensis

1

. Venation partially areolate or venation free

4

. Venation free

5. Laminae pedate; (3-)4-pinnate-pinnatiﬁd at

base; rachises greenish or pale abaxially.

P. deﬂexa

1615 (SP); Idem; id., trilha próximo da Cursino, 20-

I-2015, Hirai et al. 760 (SP); Rio de Janeiro: Itatiaia,

Parque Nacional do Itatiaia, na Trilha da Cachoeira

Poranga, 22° 26’ 26.4” S, 44° 36’ 39.9” W, 905 m

elevation, 8-V-2015, Prado & Hirai 2395 (SP).

5. Laminae not pedate, plane; 1–3-pinnate-

pinnatifid at base; rachises dark brown

abaxially.

P. lata

4

. Venation partially areolate

The SEM images of the spores were obtained from

the examination of herbarium collections at SP and

RB Herbarium (Table 1). All images were prepared at

the Instituto de PesquisasAmbientais (IPA, previously

called Instituto de Botânica - IBt), except Pteris

deﬂexa were taken at the Kew Herbarium and P.

praealta at the Instituto de Pesquisas Jardim Botânico

do Rio de Janeiro. The spores were described based on

the morphology used by Tryon & Lugardon (1990),

Lorscheitter et al. (2001), Lellinger (2002), and

Palacios-Rios et al. (2017).

6

. Rhizome erect; rachises dark brown

abaxially; veins abaxially easily visible

and prominent; laminae densely pilose;

Andes.

P. lechleri

6

. Rhizome creeping; rachises pale abaxially;

veins abaxially inconspicuous and not

prominent; laminae moderately pilose or

just with few hairs spread; east Brazil and

northeastern Argentina.

P. sericea

261

Bol. Soc. Argent. Bot. 59 (3) 2024

Table 1. Spore characteristics of the studied species of Pteris from Brazil. The species names in bold

correspond to the species segregated ones (the names without bold correspond to the previous name

applied to the taxon).

Cingulum =

equatorial ﬂange

Commissural

Species

Distal view

Proximal view

Figure

ﬂange

P. deﬂexa

nearly continuous, well-deﬁned

coarsely tuberculate-

sparsely verrucose

Fig. 2A, B

entire

rugate

P. lata

nearly continuous, deﬁned

entire

sparsely tuberculate-

verrucate

laevigate with

scattered globules

Fig. 2C, D

Fig. 4E, F

Fig. 4A, B

P. splendens

continuous,

inconspicuous densely rugate-

rugate-tuberculate,

without perforations

undulate, ﬂattened

verrucate

P. organensis continuous,

slightly undulate

inconspicuous prominently rugate

low rugate-tuberculate

with globules,

with globules

without perforations

P. praealta

continuous,

inconspicuous coarsely rugate

low rugate with

Fig. 4C, D

entire, with a

some perforations

distal depression

P. lechleri

continuous, entire well-deﬁned

reticulate with free

laevigate with

Fig. 2E, F

tubercles in the lumens scattered globules

P. sericea

continuous, entire inconspicuous rugate slightly low tuberculate

Fig. 2G, H

Taxonomic treatment for the segregated species

the adaxial surface, dark brown abaxially; middle

. Pteris lata Kaulf., Link, Hort. Berol. 2: 28-29. pinnae 7-10 x 1.7-2.5 cm, pinnatiﬁd, diverging from

833. TYPE: BRAZIL. Material from Brazil, rachises at an acute angle; distal pinnae 4-8 x 0.8-

1

cultivated at Hortus Berolinensis, Berlin (Lectotype, 1 cm towards frond apex; apical pinna 7-10 x 2-3

designated by Prado 1996: 16, M! on 2 sheets [Herb. cm, deeply pinnatiﬁd; segments alternate, falcate or

J.G. Zuccarini Nr. 170602; Herb. M. Fürbringer Nr. deltate, or lanceolate, towards frond apex, pinnae

9

2574]). Figs. 1A-B; 2A-B.

or pinnules with margins entire, smooth or dentate,

or serrulate at apex of the segments, apices acute,

Plants terrestrial. Rhizomes 1-2 cm thick, short sometimes apiculate, costules awned at the base on

and compact, creeping to erect, woody, densely the adaxial surface and prominent on the abaxial

clothed at apex with lanceolate scales 0.2–0.5 cm face; apical segment tapering, sinuses between

long. Fronds 24-121 x 15-64 cm, monomorphic to the segments acute or roundish or sometimes

subdimorphic (fertile pinnae narrower than sterile biangulate; venation free, veins simple or furcate,

pinnae), erect; petioles 18-70 x 0.1-0.5 cm, 1-sulcate reaching margins of the segments, clavate at tip.

on the adaxial surface, light brown to dark brown at Sori interrupted in the sinuses and absent at apex

base, yellowish at median and distal portions, with of the segments; indusia pale, with entire margin;

scales at base, glabrous, surface smooth; laminae spores trilete; cingulum nearly continuous, entire,

chartaceous, not pedate, plane, tripartite, deltate, distal view sparsely tuberculate-verrucate, proximal

1

-3-pinnate-pinnatiﬁd at base, 1- or 2-pinnate- view laevigate with scattered globules, showing a

pinnatiﬁd at medium and distal portion, 21-51 x 12- deﬁned commissural ﬂange.

0 cm, with (4-)6-9 pairs of pinnae, these opposite,

subopposite to alternate, long-lanceolate or long-

2

Distribution: Endemic of the East coast Brazil

elliptic, sessile or petiolulate, bases asymmetric, (Rio de Janeiro and São Paulo States); it occurs at

cuneate, petiolules 1-sulcate on the adaxial surface forest margins; ca. 650-1600 m elevation.

and shortly winged, costae 1-sulcate on the adaxial

surface and prominent on the abaxial surface,

Selected specimens: BRAZIL. Rio de Janeiro:

deﬂexed, glabrous or with spreading whitish hairs; Itatiaia, Parque Nacional do Itatiaia, Hotel Simon

proximal pinnae 15–22 x 5.0-21 cm, 1- or 2-pinnate- abandonado, trilha dos Três Picos, 22° 26’ 8.8” S,

pinnatiﬁd; rachises 40-60 x 0.1-1 cm, 1-sulcate on 44° 36’43.7” W, 1116 m elevation, 9-V-2015, Prado

262

R. Y. Hirai et al. - New approaches in Pteris from Brazil

Fig. 1. A-B: Pteris lata. A: Habit. B: Detail of the rachis dark brown abaxially. C: Pteris deﬂexa; part of the

frond. D-E: Pteris organensis. D: Fertile frond. E: Detail of the veins and sori. F-G: Pteris splendens. F: Habit.

G: Detail of the veins and sori. H-I: Pteris sericea. H: Frond. I: Detail of the pinna abaxially (A: Prado et al.

2391, SP; B: Mynssen et al. 1435, SP; C: Hirai & Prado 788, SP; D-E: Hirai & Prado 793, SP; F: Hirai et al.

760, SP; G: Prado & Hirai 2395, SP; H-I: Prado & Hirai 2403, SP).

263

Bol. Soc. Argent. Bot. 59 (3) 2024

Fig. 2. Trilete spores of Pteris. A-B: P. lata. A. Distal view sparsely tuberculate-verrucate. B. Proximal

view laevigate with scattered globules, showing a deﬁned commissural ﬂange and entire cingulum. C-D: P.

deﬂexa. C: Distal view coarsely tuberculate-rugate. D: Proximal view sparsely verrucose, showing a well-

deﬁned commissural ﬂange. E-F: P. sericea. E: Distal view rugate. F: Proximal view slightly low tuberculate

showing inconspicuous commissural ﬂange and entire cingulum. G-H: P. lechleri. G: Distal view reticulate. H:

Proximal view showing a well-deﬁned commissural ﬂange and entire cingulum. Abbreviations= ci: cingulum;

cf: commissural ﬂange; gl: globules. Scale= 10 µm (A-B: Mynssen 1435, SP; C-D: Hatschbach 26371, HB;

E-F: Labiak 4264, SP; G-H: Jiménez & Huaylla 2659, SP).

264

R. Y. Hirai et al. - New approaches in Pteris from Brazil

&

Hirai 2404 (SP); idem, Trilha para a Cachoeira Martínez & Prado, also similar to P. deflexa.

Véu da Noiva, 22° 25’ 33.5” S, 44° 37’ 19.5” W, However, P. exigua has smaller indusia, 0.5-

200 m elevation, 7-XI-2016, Prado & Hirai 2454 5 mm long (versus larger 5-12 mm long) and

SP); Rio de Janeiro, Parque Nacional da Tijuca, whitish (versus paleaceous); costules sparsely scaly

trilha para o Bico do Papagaio, 22° 56’ 51” S, 43° abaxially (versus sparsely pilose), segment apex

7’ 29” W, 698 m elevation, 6-IX-2014, Mynssen obtuse (versus acute), and conspicuous dentate

1

(

1

et al. 1435 (NY, RB, SP); Terezópolis, Rua Paulo margins of the sterile segments (versus entire). The

Barreto, 22° 27’ S, 42° 57’ W, 935 m elevation, 14- spore of P. exigua (Fig. 3A-C from Martínez &

VII-1992, Prado et al. 1089 (NY, SP). São José do Prado, 2011) is more similar to P. lata (Fig. 2A-B)

Barreiro, subindo a trilha atrás da Pousada Campos than P. deﬂexa (Fig. 2C-D).

da Bocaina, 22° 43’ 12.3” S, 44° 37’ 8.7” W, 1590

m elevation, Prado et al. 2391 (SP); Idem, descendo

a mata à direita, em frente a Pousada Campos da 2. Pteris organensis (Fée) J. Prado & R.Y Hirai,

Bocaina, 22° 42’ 54.7” S, 44° 37’ 8.9” W, 1528 m comb. nov. Litobrochia organensis Fée, Cr. Vasc. Br.

elevation, 11-XI-2016, Prado & Hirai 2479 (SP); 1: 240. 1869. Pteris splendens Kaulf. var. organense

a

São Paulo: São Luiz do Paraitinga, Parque Estadual (Fée) Dutra, An. 1 Reun. Sul Amer. Bot. 2: 33. 1938.).

da Serra do Mar, Núcleo Santa Virgínia, trilha do TYPE. BRAZIL. Rio de Janeiro, Serra dos Órgãos,

Rio Paraitinga, 19-XI-1998, Kameyama et al. 125 VI-1869, A. Glaziou 3329 (Lectotype, ﬁrst-step

(SP); Ubatuba, Parque Estadual da Serra do Mar, designated by Prado, 1996: 12, P!; second-step here

Núcleo Cunha, 18-XII-1996, Salino 2950 (BHCB, designated P! barcode P00609078; isolectotypes, P!

SP).

barcode P00609077, RB! barcode RB00543343).

Figs. 1D-E; 3A-C; 4A-B.

The Brazilian specimens of Pteris lata were

treated together with the specimens of P. deﬂexa

Plants terrestrial. Rhizomes 1-4 cm thick, erect

by Prado & Windisch (2000) and Prado & Hirai or suberect, short, woody, densely clothed at the

2024). Whereas P. lata is restricted to the East apex with long-lanceolate scales 0.2-0.5 cm long.

(

coast of Brazil, P. deflexa occurs in Central Fronds 40-100 x 15-60 cm, subdimorphic (fertile

America, Antilles, and South America (Colombia pinnae narrower than sterile pinnae), erect; petioles

to Argentina); in Brazil from the South of Bahia and stout, 10-100 x 0.2-0.4 cm, deeply 1-sulcate on

Mato Grosso to Rio Grande do Sul states.

the adaxial surface, dark brown to reddish at base

Both species can be distinguished and light brown in the median and distal portions,

morphologically: The lamina of Pteris lata is plane with scales on the base, clothed with whitish

and 1-3-pinnate-pinnatiﬁd at base (Fig. 1A; versus to light brown hairs; laminae subcoriaceous,

pedate and 4-pinnate-pinnatiﬁd at base in P. deﬂexa, 1-pinnate-imparipinnate, oblong-lanceolate, 40-

Fig. 1C), the color of the rachises dark brown 120 x 15-60 cm, with (2-)6-10 pairs of pinnae, these

abaxially (Fig. 1B; versus pale or greenish). It is entire, lanceolate (the sterile ones) or narrowly

often diﬃcult to observe the division of the lamina oblong or lanceolate (the fertile ones), opposite or

in herbarium specimens because many of them only subopposite, petiolulate, short-petiolulate or sessile,

show part of the plant, especially in P. deﬂexa (with bases cuneate or asymmetric, costae sulcate on

fronds up to 2.5 m long), which are generally much the adaxial surface and prominent on the abaxial

larger plants than P. lata (up to ca. 1.2 m long). surface, margins entire to dentate; proximal pinnae

The pedate lamina division of P. deﬂexa is easily 10-25 x 1-2.5 cm, entire; rachises 10-60 x 0.1-

observed in plants in the ﬁeld but it is generally 0.3 cm, 1-sulcate on the adaxial surface; middle

absent in the herbarium specimens. Furthermore, pinnae 9-25 x 1-2 cm, diverging from the rachises

the spores in P. lata are morphologically distinct at an acute angle; distal pinnae 6.5-20 x 0.9-1.5

by the sparsely tuberculate-verrucate in distal view cm towards frond apex, base shortly decurrent on

(Fig. 2A-B; versus coarsely tuberculate-rugate in P. the rachises; apical pinna 8-15 x 1.5-2 cm, entire;

deﬂexa, Fig. 2C-D). venation areolate, visible and prominent on the

Martínez & Prado (2011) described a new abaxial surface of the pinnae, large areoles joined to

species from northwestern Argentina, Pteris exigua the costa, oblique, approximately isodiametric, free

265

Bol. Soc. Argent. Bot. 59 (3) 2024

Fig. 3. A-C: Pteris organensis. A: Habit with one sterile and one fertile frond. B: Detail of the venation in

a fertile pinna. C: Detail of the venation in a sterile pinna. D-E: Pteris praealta. D: Habit. E: Detail of the

venation in a sterile pinna. (A-C: Hirai & Prado 778B, SP; D-E: Paciencia 375, SP).

266

R. Y. Hirai et al. - New approaches in Pteris from Brazil

Fig. 4. Trilete spores of Pteris. A-B: P. organensis. A: Distal view prominently rugate. B: Proximal view

low rugate-tuberculate with inconspicuous commissural ﬂange. C-D: P. splendens. C: Distal view densely

rugate-verrucate. D: Proximal view rugate-tuberculate with inconspicuous commissural ﬂange, cingulum

undulate. E-F: P. praealta. E: Distal view coarsely rugate. F: Proximal view low rugate with some perforations,

cingulum continuous with a distal depression. Scale= 10 µm (A-B: Prado & Hirai 2482, SP; C-D: Prado &

Silva 1615, SP; E-F: Brade 14944, RB).

veins arising from the areoles near the margins of

Distribution: Endemic of the East coast Brazil

the pinnae, in the sterile frond the apex of the veins (Bahia, Minas Gerais, Espírito Santo, Rio de

clavate. Sori absent at base and at the apex of the Janeiro, São Paulo, and Paraná); it grows inside

pinnae; indusia pale, margin entire; spores trilete; forests at medium elevations; ca. 450-1700 m

cingulum continuous, slightly undulate, distal elevation.

view prominently rugate with globules, proximal

view low rugate-tuberculate with globules, without

perforations, and inconspicuous commissural Serra dos Sete Paus, 6 km de Almadina na estrada

ﬂange. para Ibitupã, 7 km N para a Comunidade de Sete

Selected specimens: BRAZIL. Bahia: Almadina,

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Bol. Soc. Argent. Bot. 59 (3) 2024

Paus, na nascente do Rio Almada, 14° 44’S, 39° 42’ (Fig. 3A; versus widely oblong in P. splendens, Fig.

W, 578 m elevation, 19-VII-2005, Matos et al. 708 1F) and by the fertile pinnae narrowly oblong or

(

SP). Minas Gerais: Araponga, Parque Estadual lanceolate (Figs. 1D, 3A; versus widely oblong in

da Serra do Brigadeiro, Serra das Cabeças, 1400 P. splendens). Furthermore, Pteris organensis has

m elevation, 30-I-2002, Valente et al. 868 (SP); prominently rugate spores in distal view (Fig. 4A),

Descobertos, Reserva Biológica do Grama, trilha whereas P. splendens has densely rugate-verrucate

principal, 15-VI-2005, Jascone et al. 432 (SP); spores (Fig. 4C).

Lima Duarte, Parque Estadual do Ibitipoca, 21° 42’

Pteris splendens occurs in Paraguay and Brazil

11” S, 43° 53’ 0.6” W, 10-VIII-2005, Mynssen et (from the state of Ceará to Rio Grande do Sul) and

al. 826 (RB, SP). Espírito Santo: Santa Teresa, Alto P. organensis is restricted to the East coast of Brazil.

de Santo Antônio, terreno do Bozza, 19° 54’ 31”

S, 40° 35’ 28” W, 760 m elevation, 12-VII-2007,

Labiak et al. 4043 (SP, UPCB); Idem, id., entrada 3. Pteris praealta (Fée) J. Prado & R.Y. Hirai,

à esquerda (Madeireira MADEFEL), 19° 54’ 36.3” comb. nov. Litobrochia praealta Fée, Cr. vasc.

S, 40° 35’ 30.2” W, 810 m elevation, 8-VII-2016, Br. 1: 46, t. 11, ﬁg. 2. 1869. TYPE. BRAZIL. Rio

Hirai & Prado 793 (SP). Rio de Janeiro: Angra dos de Janeiro, Jacuecanga, 16-VI-1868, A. Glaziou

Reis, Serra do Mar, descendo pela BR 494, entrada 2304 (Lectotype, designated by Prado, 1996: 13,

à direita, estacionamento da Capela, 22° 52’ 8” S, P! on 4 sheets barcodes P00609073, P00609074,

4

&

1

&

4° 15’ 12.1” W, 467 m elevation, 11-V-2015, Hirai P00609075, P00609076; isolectotypes, K! barcode

Prado 778B (SP), Idem, id., 22° 52’ 8” S, 44° K000589370, S05-9754 image!). Figs. 3D-E; 4E-F.

5’ 12.1” W, 467 m elevation, 4-XI-2016, Prado

Hirai 2444 (SP); Petrópolis, Vale das Videiras,

Plants terrestrial. Rhizomes 1-4 cm thick, long-

subida do Morro da Cuca, 13-VI-2000, Prado et creeping, woody, densely clothed at apex with long-

al. 1086 (NY, SP); Santa Maria Madalena, Pedra lanceolate scales 0.2-0.5 cm long. Fronds 40-200 x

Dubois, trilha para o alto da pedra, 800-1000 m, 20-60 cm, monomorphic to subdimorphic (fertile

1

8-VI-2004, Mello-Silva et al. 2649 (SP, SPF). pinnae narrower than sterile pinnae), erect; petioles

São Paulo: São José do Barreiro, subindo a trilha stout, 20-120 x 0.2-0.4 cm, deeply 1-sulcate on

atrás da Pousada Campos da Bocaina, 22° 43’12.3” the adaxial surface, dark brown to reddish at base,

S, 44° 37’ 8.7” W, 1590 m elevation, 23-II-2015, median and distal portions, with scales on the base;

Prado et al. 2387 (SP); Idem, Parque Nacional da laminae subcoriaceous, 1-pinnate-imparipinnate,

Serra da Bocaina, na trilha para a Cachoeira Santo oblong-lanceolate, 30-150 x 15-60 cm, with 4-8

Izidro, 22° 44’ 37.5” S, 44° 36’ 56.2” W, 1562 m pairs of pinnae, these entire, narrowly lanceolate to

elevation, 22-II-2015, Hirai et al. 766 (SP); Idem, oblong, opposite or subopposite, petiolulate, short-

trilha para o Condomínio, que começa em frente à petiolulate or sessile, bases cuneate or asymmetric,

Pousada do Lageado, 22° 42’ 34.8” S, 44° 37’30.5” costae sulcate on the adaxial surface and prominent

W, S-SE, 1710 m elevation, 12-XI-2016, Prado & on the abaxial surface, margins entire to dentate;

Hirai 2482 (SP). Paraná: Alexandra, PARNA Saint- proximal pinnae 10-30 x 1-2 cm, entire or furcate;

Hilaire/Lange (Serra da Prata), 1030 m elevation, rachises 10-40 x 0.1-0.2 cm, 1-sulcate on the

2

1-V-2005, Paciencia et al. 2195 (SP); Sengés, Vale adaxial surface, shortly winged in the distal region,

do Corisco, próximo ao Mirante, 24° 12’ 16” S, 49° dark brown abaxially; middle pinnae 9-20 x 1-2 cm,

2

1

1’ 6” W, 880 m elevation, 4-I-2008, Prado et al. diverging from the rachises at an acute angle; distal

801 (UPCB, SP).

pinnae 15-20 x 0.9-2 cm towards frond apex, base

shortly decurrent on the rachises; apical pinna 10-

This species was put in synonym of Pteris 20 x 1.5-2 cm, entire; venation areolate, visible and

splendens by Prado & Windisch (2000) and Prado prominent on the abaxial surface of the pinnae, large

Hirai (2024) followed this concept, but the re- areoles joined to the costa, oblique, approximately

&

analyses of more herbarium specimens, as well as isodiametric, free veins arising from the areoles

some populations of P. organensis in the ﬁeld, we near the margins of the pinnae, in the sterile frond

are now able to recognize it as a diﬀerent species. the apex of the veins clavate. Sori absent at base

It can be recognized by the sterile pinnae lanceolate and at the apex of the pinnae; indusia pale, margin

268

R. Y. Hirai et al. - New approaches in Pteris from Brazil

entire; spores trilete; cingulum continuous, entire, scales 0.3-0.5 cm long. Fronds 25-200 x 10-60

with a distal depression, distal view coarsely rugate, cm, monomorphic to subdimorphic (fertile pinnae

proximal view low rugate with some perforations, narrower than sterile pinnae), erect; petioles 20-

and inconspicuous commissural ﬂange.

70 x 0.2-0.5 cm, 2- or 3-sulcate on the adaxial

surface, light brown to dark brown (sometimes

Distribution: Endemic of the East coast Brazil reddish at base), with scales at base, clothed with

(Bahia and Rio de Janeiro); it grows inside forests a prominent sericeous indument, hairs whitish,

at low elevations near the coast; 0-600 m elevation. 1-3-celled, ca. 1 mm long, this indument covering

moderately and occurring on all parts of the

Selected specimens: BRAZIL. Bahia: Una, frond or just with few hairs spread; laminae

REBio de Una, 16-IV-1999, Paciencia 375 (SP). chartaceous, deltate, 1-pinnate-pinnatifid, 15-

Rio de Janeiro: Angra dos Reis, Serra do Mar, 600 90 x 10-60 cm, with 3-6 pairs of pinnae, these

m elevation, 29-VI-1935, Brade 14944 (RB).

opposite to subopposite, oblong-lanceolate to

elliptic, sessile to petiolulate (proximal pinnae),

Prado & Windisch (2000) treated this species bases of the basiscopic side shortly decurrent on

as a synonym of Pteris splendens and Prado & the rachises, costae sulcate on the adaxial surface

Hirai (2024) followed this concept. However, and prominent on the abaxial surface; pair of

the rhizome of Pteris praealta is long-creeping proximal pinnae furcate, pinnatiﬁd, basiscopic

(Fig. 3D), whereas in P. splendens is short, erect portion of the fork 9-20 x 2.5-7 cm, towards frond

to suberect. In 1935, Brade had already observed base, acroscopic portion of the fork 10-30 x 4-8

this morphological characteristic of the rhizome cm; rachises 7-70 x 0.1-0.4 cm, 2-sulcate on the

of P. praealta and he noted this information on his adaxial surface, pale abaxially; middle pinnae

herbarium specimen label (Brade 14944).

8-23 x 3-8 cm, diverging from the rachises at an

Additionally, Pteris praealta has pinnae narrower acute angle; distal pinnae 6-9 x 1.5-3 cm towards

up to 2 cm wide) than P. splendens (up to 4 cm frond apex; apical pinna 9-15 x 4-6 cm, deeply

(

wide). Distally the main rachis in P. praealta is dark pinnatiﬁd; proximal segments shorter than the

brown abaxially (versus light brown or yellowish in median segments, alternate, linear to lanceolate,

P. splendens). The spores in P. praealta are diﬀerent slightly falcate, apices acute or obtuse-roundish,

in both views: coarsely rugate in the distal view margins entire, slightly dentate at apex; apical

(Fig. 4E) and low rugate with some perforations segment short, lobate to pinnatifid, costules

in the proximal view (Fig. 4F), whereas the spores prominent on the abaxial surface, sinuses between

of P. splendens are densely rugate-verrucate in the the segments acute, roundish, or biangulate;

distal view (Fig. 4C) and rugate-tuberculate in the venation partially areolate, one large and elongate

proximal view (Fig. 4D).

areole, another small areole joined to the costa,

between two adjacent costules, the areolate joined

to the costule and with free veins arising from the

4

. Pteris sericea (Fée) Christ, Bull. Herb. areoles near the margins of the segments, veins

Boissier, sér. 2, 2: 551. 1902. Litobrochia abaxially inconspicuous and not prominent, apex

sericea Fée, Cr. Vasc. Br. 1: 118, t. 11, ﬁg 3. of the veins slightly clavate. Sori interrupted in

1

869. TYPE. BRAZIL. Rio de Janeiro, 1868, the sinuses and absent at apex of the segments;

A. Glaziou 2312 (Lectotype, designated by indusia pale or whitish, margin entire; spores

Prado, 1995[1996]: 40, P! on 4 sheets barcodes trilete; cingulum continuous, entire, distal view

P00609054, P00609055, P00609056, P00609057; rugate, proximal view slightly low tuberculate,

isolectotypes BR barcode BR0000013206406 and inconspicuous commissural ﬂange.

image!, BR barcode BR0000013206413 image!,

K! barcode K000589377, S05-9753 image!).

Figs. 1H-I; 2E-F.

Distribution: Southeastern Brazil (Espírito

Santo, Minas Gerais, Rio de Janeiro, São Paulo,

Paraná, Santa Catarina, and Rio Grande do Sul),

Plants terrestrial. Rhizomes 0.5-1 cm thick, and northeastern Argentina; it grows inside forests;

creeping, woody, clothed at apex with lanceolate ca. 600-1750 m elevation.

269

Bol. Soc. Argent. Bot. 59 (3) 2024

Selected specimens: BRAZIL. Espírito Santo: UPCB, VT); Ponta Grossa, Parque Estadual de

Venda Nova do Imigrante, Pedra do Rego, na trilha, Vila Velha, 25° 14’ S, 50° 0’ W, 1000 m elevation,

2

9

0° 18’ 21” S, 41° 10’ 25” W, 1230 m elevation, 10-XI-2003, Schwartsburd 1 (SP, UPCB); Serra

-IV-2019, Cordeiro & Brotto 6246 (MBM). Minas do Mar, Ypiranga, “in silva primaeva”, 830 m

Gerais: Caldas, 1873, Mosén 3088 (K, P, R, S, elevation, 16-I-1914, Dusén 14423a (BM, K, US).

UPS); Poços de Caldas, Morro do Ferro, 26-IV- Santa Catarina: Canoinhas, Rio da Areia, 21-IV-

1

968, Ana-Lima 68-102 (IPA). Rio de Janeiro: 1962, Reitz & Klein 12713 (HB, MBM); Fragoso,

ltatiaia, Parque Nacional do Itatiaia, Hotel Simon s.d., Hatschbach 2492 (MBM, RB); Marata,

abandonado, trilha dos Três Picos, 22° 26’ 8.8” S, Porto União, 19-I-1952, Reitz 4705 (HB, PACA);

4

&

4° 36’43.7” W, 1116 m elevation, 9-V-2015, Prado Hammonia, 11-X-1922, Luederwaldt s.n. (SP

Hirai 2403 (SP); Macaé, Glicério, trilha Água 21650); Porto União, Pinheiral near Porto União on

Fria, ca. 800 m elevation, 7-VI-2004, Mynssen & road to Santa Rosa, 750–800 m elevation, 18-XII-

Bovini 637 (RB, SP); Nova Friburgo, subida para 1956, Smith & Reitz 8749 (HB, K, R, US); Xanxerê,

o Pico da Caledônea, 22° 40’ S, 42° 35’ W, 1706 “Pinheiral and ruderal”, Faxinal dos Guedes, 700-

m elevation, 15-VI-2000, Prado et al. 1093 (NY, 900 m elevation, 3-I-1957, Smith & Reitz 9790 (R).

SP); Rio Funil, perto do Estado de São Paulo, 8-XI- Rio Grande do Sul: Santa Cruz, Herval do Paredão,

1

956, Handro 668 (SP, SPF); Teresópolis, Toca s.d., Juergens & Stier 190 (PACA).

dos Caçadores, 1300 m, s.d., Brade 9318 (BM,

R). São Paulo: Campos do Jordão, III-1946, Leite

Prado & Windisch (2000) and Prado & Hirai

1

56 (FCAB); Iporanga/Apiaí, Parque Estadual (2024) treated the Brazilian specimens of this

Turístico do Alto Ribeira (PETAR), 24° 32’ 22” S, species as Pteris lechleri Mett, but the available

8° 41’ 36.” W, 7-VII-2012, Mazziero & Albiero name to apply to this species from Brazil is P.

115 (SP, UPCB); Jundiaí, Serra do Japi, trilha indo sericea. Pteris lechleri occurs in Central America

4

1

para a Cachoeira Paraíso, 23° 14’ S, 46° 56’ W, 996 (Panama) and South America (from Colombia to

m elevation, 13-XI-2009, Prado et. al. 2061 (SP, Bolivia).

UEC); Paranapiacaba, Serra de Paranapiacaba-

Pteris sericea diﬀers from P. lechleri in having

Serra da Boa Vista, 700 m elevation, X-1925, rhizome creeping, laminae less pubescent, rachises

Brade 8404 (HB, R); São José dos Campos, pale abaxially, and geographically it is isolated

estrada São Francisco Xavier-Jardinópolis, Faz. in the Atlantic Forest (East coast of Brazil and

Santa Cruz, 14-IV-1981, Vieira 24 (HRCB); São Misiones in northeastern Argentina), whereas

Paulo, Parque Estadual das Fontes do Ipiranga, P. lechleri has its distribution restricted to the

2

3-IV-1979, Tosta Silva 303 (SP); Idem, Jaraguá, Andes, rhizome erect, laminae densely pubescent,

-II-1922, Luederwaldt s.n. (SP 21648). Paraná: and rachises dark brown. Tryon & Stolze (1989)

Adrianópolis, Parque Estadual das Lauráceas, 24° had already observed that P. lechleri has an erect

0’ S, 48° 32’ W, 12-XII-2006, Matos et al. 1296 rhizome in plants from Peru, where the type

SP, UPCB); Antônio Olinto, Água Amarela, 1-X- specimen of the name came from. However, this

969, Hatschbach 22304 (MBM, PACA, UPCB); characteristic may go unnoticed because many

4

(

1

Balsa Nova, Serra de São Luis, 16-VIII-1970, materials in herbaria are without rhizomes. The

Hatschbach 24479 (MBM, PACA); Campo Largo, ornamentation of the spores is also diﬀerent in both

Rodovia BR-277, junto ao viaduto de acesso a species: P. lechleri has reticulate spores with free

Campo Largo, 25° 26’ 3” S, 49° 30’ 31” W, 975 m tubercles in the lumens (Fig. 2G-H) and P. sericea

elevation, 3-IV-2011, Fiaschi et al. 3720 (SP, SPF); has rugate spores in distal view (Fig. 2E-F).

Campo Mourão, Mata do lado da Usina Mourão,

In the general aspect, Pteris sotae O. Martínez

4° 7’ 6” S, 52° 19’ 13” W, 600 m elevation, resembles P. sericea, but diﬀers by the entire and

4-XII-2007, Labiak et al. 4264 (SP, UPCB); long terminal segment of the pinnae, sometimes

2

Curitiba, Parque Iguaçú, 22-VIII-1984, Oliveira caudate (versus lobate to pinnatiﬁd and short),

07 (MBM); Guarapuava, Estrada Municipal rounded apex of the laminar hairs (versus acicular),

8

Benedito de Paula Louro, próximo a Cachoeira São and coarsely rugate spores on distal view (versus

Francisco, 25° 3’ S, 51° 12’ W, 1080 m elevation, rugate). Pteris sotae occurs only in the Tucuman-

1

5-X-2011, Prado & Hirai 2190 (SP, SPF, TAIF, Bolivia forests in Argentina (Martínez, 2016).

270

R. Y. Hirai et al. - New approaches in Pteris from Brazil

concluSionS

LELLINGER, D. B. 2002. A modern multilingual

glossary for taxonomic Pteridology. Pteridologia

3: 1-263.

The present study revised the current

circumscriptions of three species of Pteris: P. LORSCHEITTER, M. L., A. R. ASHRAF, P.

deﬂexa, P. lechleri, and P. splendens and based

on morphological evidence four new taxa were

recognized: P. lata segregated from P. deﬂexa; P.

G. WINDISCH & V. MOSBRUGGER. 2001.

Pteridophyte spores of Rio Grande do Sul ﬂora,

Brazil. Part III. Palaeontographica 260: 1-165.

organensis and P. praealta from P. splendens; and MARTÍNEZ, O. G. 2016. Pteris sotae (Pteridaceae),

P. sericea from P. lechleri. These new data revealed

that the Brazilian Atlantic Forest has more species

of ferns than previously recognized in the Flora of

Brazil by Prado et al. (2015).

a new endemic species for the Argentinean ﬂora.

Phytotaxa 267: 291-295.

http://dx.doi.org/10.11646/phytotaxa.267.4.6

MARTÍNEZ, O. G. & M. A. MORBELLI. 2009. The

spores of Pteris cretica complex (Pteridaceae-

Pteridophyta) in America. Grana 48: 193-204.

MARTÍNEZ, O. G. & J. PRADO. 2011. Pteris exigua

acKnowledgementS

(

Pteridaceae), a new endemic species from

This study was largely funded by a grant

to the ﬁrst author from Conselho Nacional de

Desenvolvimento Cientíﬁco e Tecnológico (CNPq,

Tucumano-Boliviano forests in northwestern

Argentina. Brittonia 63: 295-299.

http://dx.doi.org/10.1007/s12228-010-9166-9

grant no. 158895/2014-2) and the last author PALACIOS-RIOS, M., C. PRADA, J. M. G. GALÁN

from Fundação de Amparo à Pesquisa do Estado

de São Paulo (FAPESP, grants no. 2013/23702-

& J. NOA. 2017. Spore types in Mexican and

Mesoamerican species of Pteris L. (Pteridaceae).

Grana 56: 241-256.

0

, 2022/12597-0, 2023/13634-0). We thank Dra.

Luciana Benjamim Benatti (Instituto de Pesquisas

http://dx.doi.org/10.1080/00173134.2016.1217038

Ambientais-IPA, former Instituto de Botânica-IBt) PPG I. 2016. A community-derived classiﬁcation for

for preparing the SEM images of the spores. We

also thank the anonymous reviewers for suggestions

to improve this manuscript.

extant lycophytes and ferns. J. Syst. Evol. 54: 563-

603. https://doi.org/10.1111/jse.12229

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Hoehnea 22: 39-40.

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8

: 11-20.

RYH and JP contributed to the study conception PRADO, J. & R.Y. HIRAI. 2024. Pteris in Flora

and prepared the descriptions with taxonomical and

nomenclatural approaches. All authors, including

CMM, helped in the ﬁeldwork and prepared the

e Funga do Brasil. Jardim Botânico do Rio de

Janeiro. Available at: https://ﬂoradobrasil.jbrj.gov.

br/FB91978 [Accessed on: 14 February 2024].

material and images of the spores for SEM, as well PRADO, J. & P. G. WINDISCH. 2000. The genus Pteris

as writing this manuscript.

L. (Pteridaceae) in Brazil. Bol. Inst. Bot. (São Paulo)

1

3: 103-199.

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