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Articles • Hoehnea 52 • 2025 • https://doi.org/10.1590/2236-8906e022024 linkcopy

Open-access An updated checklist of lycophytes and ferns for Ceará State, Brazil1

Lista atualizada de licófitas e samambaias para o Estado do Ceará, Brasil

AuthorshipSCIMAGO INSTITUTIONS RANKINGS

ABSTRACT

As very few studies have focused on the ferns and lycophytes of Ceará State, Brazil, our research aims to examine and update the diversity and distribution of these taxa, as part of the “Flora do Ceará project: knowing to conserve.” The research was carried out through field trips, bibliographic surveys, and analyses of specimens in situ and by virtual herbaria. A total of 197 infrageneric taxa belonging to 76 genera and 28 families have been recorded for Ceará. Our investigation revealed 50 additional occurrences for the State compared to the Flora and Funga of Brazil project. The species occur predominantly in the Semideciduous Seasonal Forest (139 species, ~70%). The species were recorded in 13 legally constituted Conservation Areas, with the Serra de Baturité Environmental Protection Area standing out (61 species).

Keywords:
biodiversity; Northeastern Brazil; pteridophytes; richness; semiarid

RESUMO

Como poucos trabalhos anteriores estudaram licófitas e samambaias no Estado do Ceará, Brasil, este estudo foi concebido para examinar e atualizar a diversidade e as distribuições atuais dos táxons que ocorrem no Estado, como parte do “Projeto Flora do Ceará: conhecer para conservar”. A pesquisa foi realizada através de visitas a campo, levantamentos bibliográficos e análises de espécimes in situ e por herbários virtuais. Foram registrados 197 táxons infragenéricos pertencentes a 76 gêneros e 28 famílias para o Ceará. Esta pesquisa demonstrou 50 ocorrências adicionais para o Estado, comparado com o projeto Flora e Funga do Brasil. As espécies ocorrem predominantemente em Floresta Estacional Semidecidual (139 espécies, ~70%). Foram registradas ocorrências em pelo menos 13 Unidades de Conservação legalmente constituídas, com destaque para a Área de Proteção Ambiental da Serra de Baturité (61 espécies).

Palavras-chave:
biodiversidade; Nordeste do Brasil; pteridófitas; riqueza; semiárido

Introduction

The global diversity of lycophytes and ferns is approximately 11,916 species belonging to 337 genera, 51 families, 14 orders (PPG I 2016). Lycophytes and ferns constitute important components of the total flora of rainforests, generally comprising approximately 10% of the vascular plant species found there (Grayum & Churchill 1987).

In Brazil, this group of vascular plants includes 1,412 species, 162 genera, and 39 families (Ferns and Lycophytes 2024). The Southeastern region of Brazil is considered the third center of pteridophyte flora endemism in the Neotropical region, with approximately 15% of all endemic species being recorded there (Suissa & Sundue 2020). The Atlantic Forest phytogeographic domain has the highest species richness, harboring approximately 71% of the species occurring in Brazil (Prado et al. 2015).

The northeastern region of Brazil has approximately 554 species of lycophytes and ferns, corresponding to 39% of all species occurring in Brazil (Ferns and Lycophytes 2024). Several studies have been carried out in that region focusing in these taxa, including, e.g. Fernandes et al. (2022) - Maranhão State; Nascimento (2021) - Rio Grande do Norte State; Santiago et al. (2014) - Paraíba State; Pietrobom (2004) - Alagoas State; Paula-Zárate (2005) - Ceará State; Matos et al. (2010), - Bahia State; Barros et al. (2006) - Pernambuco State; and Andrade et al. (2022) - Sergipe State. No surveys have specifically covered Piauí State.

Studies on these taxa for the State of Ceará have been drawn up based on floristic lists such as those by Huber (1908), Brade (1940), Paula-Zárate (1993), Paula-Zárate et al. (2007), Pinheiro et al. (2022) with emphasis mainly on the species occurring in the Serra de Baturité, Dias Rocha (1908), including Serra de Maranguape and Serra de Baturité, Reinaldo et al. (2015), covering the species occurring in the municipalities of Ceará belonging to the Chapada do Araripe Environmental Protection Area in the south of the State; and Dias Rocha (1946), Braga (1951) and Paula-Zárate (2005) for the entire State of Ceará. With the aim of contributing to a broader understanding of the pteridophyte flora of Ceará, and to update the data on these taxa, an inventory of the current diversity and the distribution of lycophyte and fern species occurring in Ceará State was undertaken.

Material and Methods

In order to draw up the updated taxonomic list, we first reviewed the literature on the pteridophyte flora of Ceará: Brade (1940), Braga (1951), Paula-Zárate (1993), Paula-Zárate (2005), Paula-Zárate et al. (2007), Reinaldo et al. (2015) and Pinheiro et al. (2022). The names cited in these works have been revised to update their nomenclatural validity, and the synonyms are in accordance with IPNI (https://www.ipni.org/), Tropicos (https://www.tropicos.org/) and Ferns and Lycophytes (2024 - https://floradobrasil.jbrj.gov.br/FB128483), as well as their occurrence in Ceará and Brazilian territory (Ferns and Lycophytes 2024).

The collections of the EAC and HCDAL herbaria were analyzed in situ to confirm the occurrence of the species found in Ceará (herbarium acronyms according to Thiers 2024). Random field expeditions to obtain more specimens and photographic records were carried out in locations where the occurrence of the taxa was previously known, such as the municipalities of Ubajara, Ibiapina and Pacatuba. The material was collected according to the methodology proposed by Gadelha Neto et al. (2013) and deposited in the EAC Herbarium.

To complement the taxonomic list, the database Ferns and Lycophytes (2024) was consulted to check the taxa occurring in the State according to the search criteria: i) only species of native origin or naturalized, ii) species that occur in Ceará, iii) only accepted names. Based on the names obtained, they were compared with those previously listed to confirm their occurrence in the State, and for those not previously listed, searches were carried out in the literature cited in Ferns and Lycophytes (2024), e.g. Smith (1986), Xavier et al. (2012) and Schwartsburd et al. (2014, 2017), and in the databases previously cited to certify their occurrence.

In addition, a search was made for taxa registered for the State in the online databases of speciesLink (http://www.splink.org.br/), REFLORA - Reflora Virtual Herbarium (https://floradobrasil.jbrj.gov.br/reflora/herbarioVirtual/ConsultaPublicoHVUC/ConsultaPublicoHVUC.do) and SiBBr - Brazilian Biodiversity Information System (https://www.sibbr.gov.br/). From this search, the online collections of the herbaria ALCB, ASE, BHCB, CESJ, EAN, ESA, FCAB, F-PTERIDOPHYTES, HRSN, HSTM, HUEFS, HUESB, HVASF, IAC, IAN, IPA, JPB, MBM, MICH, MO, MOSS, NY, PACA-AGP, PEUFR, R, RBR, MNHN-P, SJRP-PTERIDOPHYTA, TEPB, UB, UFP, UPCB, VIC were analyzed. Based on the names found, the taxa were filtered, considering only those previously identified for the State and those identified by lycophyte and fern specialists listed in Ferns and Lycophytes (2024).

For the collected material and samples deposited in the EAC and HCDAL herbaria that were not identified or out of date, the previously cited studies were consulted, as well as the studies de Paula-Zárate (1993), Paula-Zárate (2005), Paula-Zárate et al. (2007) and Reinaldo et al. (2015). We also carried out a comparison with images available online in the aforementioned databases, using only materials identified by experts in accordance with Ferns and Lycophytes 2024. To confirm the identity of some specimens, some experts were consulted (see the Acknowledgments section). For the genus Selaginella, species cited by Alston et al. (1981) and deposited in the BM herbarium were also considered and for Serpocaulon, we adopted the species concepts used by Sanín et al. (2023). Except for Isoetes, we do not consider including material identified only at the genus level.

The vegetation types where the species were recorded were obtained from the specimen labels, and are based on (and modified from) Figueiredo (1997) and the Technical Manual of Brazilian Vegetation (IBGE 2012), Figure 1: Vegetation Complex of the Coastal Zone - VCCZ (Pioneer Psammophilous Vegetation, behind-dune Forests, and Seasonal Semideciduous Lowland Forest : Mata de Tabuleiro); Semideciduous Seasonal Forest - SSF (Dry Forest); Dense Ombrophilous Forest - DOF (Wet Forest); Neotropical Savanna - NS (Cerrado); Forest Savanna - FS (Cerradão); Steppic Savanna - SS (Caatinga/Carrasco); Vegetation under Fluvial and/or Lacustrine influences (Riparian Forest) - VFLCI; Carnaubal - CARN.

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Figure 1
Map of phytogeographic areas and geographic distribution map of ferns and lycophytes in Ceará State, Brazil. Modified from Figueiredo et al. (1997). 1-5 represent areas with high potential for the study of the pteridoflora of Ceará (see Results and Discussion section).

A table was prepared based on the information gathered, presenting the family classification according to PPG I (2016), the scientific name of the taxon, the vegetation type where it was collected, the name of the Conservation Area (CA) where it was encountered (if appropriate), the voucher reference (Table 1). Information concerning species occurrences in CAs in Ceará were based on the specimen labels. The online platform Canva was used to prepare photographic plates.

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Table 1
Diversity of lycophytes and ferns in Ceará State, Brazil, with indication of family, name of the species, type of vegetation, occurrence in Conservation Area (CA) and herbarium record. Vegetation: DOF: Dense Ombrophilous Forest; FS: Forested Savanna; SS: Steppic Savanna; SSF: Semideciduous Seasonal Forest; VCCZ: Vegetation Complex of the Coastal Zone; Conservation Areas: Aiuaba ES: Aiuaba Ecological Station; Chapada do Araripe EPA: Chapada do Araripe Environmental Protection Area; Cocó SP: Cocó State Park; Fazenda Trussú PNHR: Fazenda Trussú Private Natural Heritage Reserve; Francy Nunes Private Natural Heritage Reserve; Lagamar do Cauípe EPA: Lagamar do Cauípe Environmental Protection Area; Pecém ES: Pecém Ecological Station; Quixadá MNM: Quixadá Monolits Natural Monument; Serra das Almas PNHR: Serra das Almas Environmental Protection Area; Serra de Baturité EPA: Serra de Baturité Environmental Protection Area; Serra Ibiapaba EPA: Serra Ibiapina Environmental Protection Area; Ubajara NP: Ubajara Nacional Park; UECE ES: UECE Ecological Station. *: New occurrences for the State.

Results and Discussion

Lycophytes and ferns occurring in Ceará State include 197 infrageneric taxa, distributed among 76 genera and 28 families (Table 1; Figures 2-3). Lycophytes represent c. 9% of the pteridophyte flora of Ceará, and are represented by three families, five genera, and 17 species. Ferns represents 91% of the pteridophyte flora of Ceará, distributed among 25 families, 71 genera, and 176 species (plus type varieties), two varieties, and two subspecies. Pteridaceae (41 spp., 1 var.; 21%), Polypodiaceae (24 spp.; 13%), and Dryopteridaceae (19 spp.,1 var.; 10%) stand out among the families with the greatest taxonomic diversity. Equisetaceae, Isoetaceae, Marattiaceae, Oleandraceae, Psilotaceae, and Schizaeaceae (all with only one species each) constitute the families with the lowest species diversity in the study area.

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Figure 2
Species of ferns occurring in Ceará State, Brazil. a-b. Adiantum deflectens Mart. a. typical adult plant. b. abaxial surface of the pinna with sori along the margin. c. Adiantum diogoanum Glaz. ex Baker. abaxial surface of the pinnules, with sori along the margins. d. Adiantum humile Kunze typical adult plant. e. Adiantum petiolatum Desv. the apical pinna; f. Anemia dentata Gardner typical adult plant. g. Asplenium pumilum Sw. abaxial view of the fertile pinna. h-i. Blechnum occidentale L. h. underside of a pinna with sori. i. young leaf.

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Figure 3
Species of ferns occurring in Ceará State, Brazil. a. Cyathea phalerata Mart. leaf. b. Lygodium venustum Sw. the apical pinna. c. Pteris biaurita L. leaf. d. Pteris denticulata Sw. typical adult plant. e. Selaginella erythropus (Mart.) Spring typical adult plant. f-g. Goniopteris poiteana (Bory) C.Presl f. typical adult plant. g. abaxial view of the fertile pinna. h-i. Tectaria incisa Cav. h. apical pinna. i. abaxial view of the fertile pinna.

The data obtained in the present study indicate that the total number of species occurring in Ceará is significantly higher than those reported in previous studies, such as Paula-Zárate (2005) (with 143 species, 62 genera, and 23 families) and the Ferns and Lycophytes (2024) website (with 152 species, 65 genera, and 24 families). The greater recorded taxonomic diversity found since Paula-Zárate’s (2005) in situ study is likely due to increased sampling efforts in the State, as well as the availability of online resources for viewing more gatherings.

We excluded five species recorded in the Ferns and Lycophytes (2024) due to the absence of a voucher (Megalastrum canescens (Kunze ex Mett.) A.R.Sm. & R.C.Moran, Microgramma percussa (Cav.) de la Sota, Phlebodium decumanum (Willd.) J.Sm.) or discrepancies in species circumscription (Phlegmariurus flexibilis (Fée) B.Øllg., Serpocaulon caceresii (Sodiro) A.R.Sm.). So, considering a total of 152 species (plus type varieties) for the State, according to Ferns and Lycophytes (2024), a total of 50 additional occurrences were recorded for the State.

This increased species diversity of ferns and lycophytes as compared to the aforementioned studies can also be observed at the genus level. Our study registered (among the most representative genera in terms of their numbers of species) Adiantum L. (23 spp.; Pteridaceae), Asplenium L. (12 spp.; Aspleniaceae), Selaginella (9 spp.; Selaginellaceae), and Anemia Sw. (8 spp.; Anemiaceae). The study by Paula-Zárate (2005) also indicated those same genera, but with lower numbers of species for each: Adiantum (17 spp.), Selaginella (7 spp.), and Anemia (6 spp.), except by Asplenium (13 spp., 1 var.). The Ferns and Lycophytes (2024) indicated the diversities of genera as: Adiantum (18 spp.), Asplenium (12 spp.), Anemia (6 spp.), Selaginella, and Cyathea (5 spp.).

Regarding the vegetation types, the ferns and lycophytes species were recorded mainly in Vegetation Complex of the Coastal Zone (VCCZ), Semideciduous Seasonal Forest (SSF), Dense Ombrophilous Forest (DOF), Steppic Savanna (SS), Forest Savanna (FS), (Figure 1), but predominantly found in SSF (139 species, ~71%), and in DOF (89 species, ~45%) corroborating Paula-Zárate (2005), who noted that the greatest species diversity occurred in humid mountain forests (“Brejos de Altitude”). Pteridaceae and Salviniaceae are the only taxa that have species in all these phytogeographical units. The families that occur most in SSF are Pteridaceae (33 spp. - ~16% of the species occurring in the State), Polypodiaceae (18 sp. - ~9%), Dryopteridaceae (17 spp. - ~9%) and Hymenophyllaceae and Thelypteridaceae with 10 species each (5% of species each). In DOF, families with more than 10 species occurring in these areas are: Pteridaceae (22 spp. - ~11%), Polypodiaceae (13 spp. - ~7%) and Aspleniaceae (10 spp. - 5%).

Some families stand out for having representatives in Steppic Savanna (SS) such as Anemiaceae with four species of Anemia, Selaginellaceae with four species of Selaginella, Pteridaceae with three species belonging to Adiantum, Ceratopteris Brongn., and Doryopteris J. Sm., and Salviniaceae, with three species (one for Azolla Lam. and two for Salvinia Ség.). Most species of Anemia exhibit xeromorphic characteristics, which allow them to be found in drier environments, such as the Caatinga and Cerrado phytogeographic domains in Brazil (Windisch 1990; Mickel 2016). On the other hand, the aquatic ferns Azolla and Salvinia, which do not exhibit xeromorphic characteristics, manage to survive in semiarid regions because their sporocarp reproductive structures have thick walls that offer protection. According to Xavier et al. (2012), the following species can tolerate dry environments by curling their leaves in the dry season to avoid desiccation: Anemia oblongifolia (Cav.) Sw., Doryopteris concolor (Langsd. & Fisch.) J. Sm., and Selaginella convoluta (Arn.) Spring can tolerate dry environments by curling their leaves in the dry season to avoid desiccation.

The pteridoflora of Ceará occurs primarily in the phytogeographical units of Seasonal Semideciduous Forest (SSF) and Dense Ombrophilous Forest (DOF) in the northern and southernmost regions of the State, as shown in Figure 1. The species distribution map reveals that there are SSF areas in the central region (Figure 1 - areas 1, 2, and 3), in the southeast (area 4), and in the southwest (area 5) that show no records of species from these taxa. These regions are potential areas for studying the State’s pteridoflora, as it has been demonstrated that the species present in Ceará predominantly inhabit this type of vegetation. More sampling effort is needed to determine whether there is truly no occurrence of ferns and lycophytes in these areas, or if it is simply a matter of insufficient sampling. If the latter is the case, it is essential to explore the biodiversity of pteridophytes in these areas to complement the data presented here.

Among the 198 infrageneric taxa surveyed, 92 were recorded in Conservation Areas (CAs) in Ceará State. Sixty-one lycophyte and fern species occur in the Serra de Baturité EPA (EPA: Environmental Protection Area), followed by the Chapada do Araripe EPA (28 spp.) and the Ubajara NP (NP: Nacional Park) (27 spp.), see Table 1. The large number of species recorded for the Serra de Baturité EPA is most likely because that it was one of the focal study areas of Paula-Zárate (1993); additional studies were also developed there, including Paula-Zárate et al. (2007) and Pinheiro et al. (2022).

Although 37% (72 spp.) of the species have been recorded in at least one Conservation Area in the State, we cannot demonstrate its effectiveness in conserving the species. For the State of Ceará, there is no detailed data on the categories of extinction risk threat for the species. However, it is important to note that for Brazil, Stenogrammitis limula (Christ) Labiak (Polypodiaceae) - was classified as endangered (EN) by Labiak et al. (2012) and is classified as critically endangered (CR) according to CNCflora (http://cncflora.jbrj.gov.br/portal). Blechnum heringeri was considered vulnerable (VU), according to Dittrich et al. (2013). Based on the new records found after 2013, Santiago et al. (2019) classified this species as Least Concern (LC). It is necessary to adopt an approach that addresses the conservation of fern and lycophyte species in the State of Ceará, in order to better understand the conservation status of these species through the advances that have caused environmental disturbances in the areas where these taxa occur (Condack & Santiago 2023).

Conclusions

The update of the data presented here shows the need for continued investigation of regional biological diversity, as even with studies already carried out and the availability of online data through digital platforms, information on the occurrence of pteridophyte taxa in the State of Ceará remains outdated. The effort to compile these data requires extensive time for literature review and species identification. The results indicate that some species still require investigation for accurate identification, such as the only known record of Isoetes for the State. The results also demonstrate that investment in sampling effort contributes to a better knowledge of biodiversity, with a 25-30% increase in the number of species recorded quantitatively since 2005 (Paula-Zárate 2005), along with updated identification of specimens deposited in the studied herbaria. This number is expected to increase, as we did not have access to all possible materials collected for the State. Future studies on geographic distribution and local extinction risk assessment of species are encouraged, as these data highlight an important gap to be filled. Additionally, studies focused on taxonomic groups with identification keys and taxonomic descriptions are needed to aid in future specimen collections.

Data availability statement

The dataset for this article is available in the SciELO Dataverse of Hoehnea, at the following link: https://doi.org/10.1590/2236-8906e022024.

Acknowledgments

Igor Renan Bonfim de Souza thanks the Conselho Nacional de Desenvolvimento Científico e Tecnológico - CNPq, for the Scientific Initiation scholarship. Maria Iracema Bezerra Loiola thanks CNPq, for the Productivity Grant (Process Nº 306723/2023-4), and Fundação Cearense de Apoio ao Desenvolvimento Científico e Tecnológico (Process MLC-0191-00147.01.00/22), for financial support for field collections. We thank the experts Lana Sylvestre, David Sanin, Fernando Matos, Vinicius Dittrich, and Thaís Almeida, who helped with the identification of some species and contributed with suggestions about the occurrence of taxa in the State. Luana Mateus de Sousa is grateful to Coordenação de Aperfeiçoamento de Pessoal de Nível Superior, for the Scholarship (Financial Code 001) and Programa de Pós-graduação em Ecologia e Recursos Naturais da Universidade Federal do Ceará.

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  • 1
    Parte do Trabalho de Conclusão de Curso do primeiro Autor

Edited by

  • Associate Editor:
    Diego Vasques

Publication Dates

  • Publication in this collection
    16 Jan 2026
  • Date of issue
    2025

History

  • Received
    18 Jan 2024
  • Accepted
    27 Dec 2024
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