Ferns and Lycophytes of Pernambuco state, Brazil: Serpocaulon (Polypodiaceae)

Abstract

In order to contribute to the study of the fern flora of the state of Pernambuco, we provide a synopsis of the genus Serpocaulon (Polypodiaceae) for Pernambuco. The three main herbaria in the state (IPA, PEUFR, and UFP) and herbaria of national reference (BHCB, RB, and SP) were visited, and online databases were consulted. Field expeditions were carried out in the different phytogeographic zones of the state. Descriptions, diagnostic illustrations, a key to species identification, taxonomic comments, and the geographic distribution of the species in the state are presented. Five species were recorded: Serpocaulon catharinae, S. menisciifolium, S. polystichum, S. richardii, and S. triseriale. They occur from sea level up to 1100 m. All species were recorded in Atlantic Forest, and S. triseriale was also found in Caatinga. The occurrence of S. polystichum and S. richardii in Pernambuco are being confirmed in this study. Additionally, the northern distribution limit of the species S. catharinae and S. menisciifolium is recorded in this state.

Key words:
Northern distribution in the Atlantic Forest; Brejos de Altitude; unprotected pteridophytes; taxonomic synopsis

Resumo

Em adição aos estudos da flora de samambaias do estado de Pernambuco, esta contribuição tem o objetivo de apresentar a sinopse do gênero Serpocaulon (Polypodiaceae) para este estado. Foram visitados os três principais herbários de Pernambuco (IPA, PEUFR e UFP), além de herbários de referência nacional (BHCB, RB e SP) e fizemos consulta a bases de dados online. Também realizamos expedições de campo nas diferentes zonas fitogeográficas do estado. Apresentamos as descrições e ilustrações diagnósticas, uma chave para identificação das espécies, além de comentários taxonômicos e a distribuição geográfica das espécies no estado. Registramos cinco espécies (Serpocaulon catharinae, S. menisciifolium, S. polystichum, S. richardii e S. triseriale), as quais habitam ao nível do mar até 1100 m. Todas as espécies estão registradas em áreas de Floresta Atlântica, sendo S. triseriale também encontrada em áreas de Caatinga. Confirmamos a ocorrência de S. polystichum e de S. richardii para Pernambuco. Além disso, neste estado, registrou-se o limite norte da distribuição das espécies S. catharinae e S. menisciifolium.

Palavras-chave:
Brejos de Altitude; Distribuição Norte na Mata Atlântica; samambaias desprotegidas; sinopse taxonômica

Introduction

Biodiversity knowledge should be the main argument for land-use planning (Newbold et al. 2015; King et al. 2021), remarking that biodiversity is the result of complex historical processes that shape species interactions and ecosystem functions (Tilman et al. 2014). However, taxonomic, ecological, and biogeographic investigations are still necessary to understand the environmental offer (Dasgupta 2021). In this sense, an accurate description of the flora as a constitutive element of the landscape is fundamental (Ulloa-Ulloa et al. 2017).

Due to its geographic conformation, with a long and narrow shape and extending from the coast to the interior of the continent, the state of Pernambuco has an interesting vegetational diversity. The vegetation types present in the state include coastal formations (such as Mangroves and “Restingas”), areas of Atlantic Forest, and vegetation types of semi-arid regions (such as “Caatinga” and Savanna) (Andrade Lima 2007). Remnants of Atlantic Forest are the most favorable for the establishment of ferns. The Northeastern Atlantic Forest (NAF), Atlantic Forest located to the north of the São Francisco River, includes two biogeographic regions: 1.) The Pernambuco Endemism Center, comprising the coastal forests and 2) “Brejos Nordestinos” or “Brejos de Altitude”, which are high elevation forests in the Northeastern region of Brazil without connection with coastal forests (Silva & Casteleti 2003). The São Francisco River has been recognized as the limit of some species’ distributions: some species occur in the South and Southwest of Brazil but are absent in the NAF, and others occur in the Amazon but do not reach the south and southeast portions of the NAF. Examples of this limit in the distribution range among ferns can be found in several species of Blechnaceae (Santiago et al. 2014), making thus the São Francisco River a natural barrier. The Caatinga domain of the state of Piauí at the west of NAF is another natural barrier. Thus, several biogeographic patterns are likely to be revealed by thorough studies of the biodiversity in the NAF.

Although fern diversity has been extensively studied in Pernambuco (Barros & Santiago 2007, 2010; Barros & Xavier 2007, 2009; Barros et al. 2006; Farias et al. 2017; Pereira et al. 2007, 2011; Pietrobom & Barros 2007; Santiago et al. 2004, 2014; Xavier & Barros 2003), the information and records of Serpocaulon species are still scarce.

In the list presented by Schwartsburd (2023) in the Flora e Funga do Brasil, the author mentions only two species of Serpocaulon from Pernambuco, and the possible occurrence of Serpocaulon fraxinifolium (Jacq.) A.R. Sm. Schwartsburd (2023) recognized S. polystichum as a synonym of S. fraxinifolium. In this respect, we agree with Sanín et al. (2023) at recognize both as different species, the former recorded in Pernambuco, and the latter a species hitherto not recorded in Brazil. However, Santiago et al. (2004) had already recorded four species of Serpocaulon previously included in Polypodium for Pernambuco. This demonstrates that the information about the genus is still to be clarified in the state.

Serpocaulon is composed of 37 species (Sanín et al. 2023) mostly occurring in the northern Andes (Sanín 2018), with a secondary center of diversity in the Brazilian Atlantic Forest (Labiak & Prado 2008; Schwartsburd & Smith 2013; Sanín et al. 2023).

With the objective of consolidating the knowledge of the fern and lycophyte flora of Pernambuco (Barros & Santiago 2007, 2010; Barros & Xavier 2007, 2009), we present a synopsis of the genus Serpocaulon (Polypodiaceae) for this state.

Material and Methods

This study was based on specimens deposited in the three main herbaria of Pernambuco (UFP, IPA, and PEUFR) and other reference collections in Brazil (BHCB, RB, and SP) (acronyms according to Thiers, continuously updated), and data from the databases Flora e Funga do Brasil 2023 (continuously updated), Specieslink (2023), and Tropicos (2023).

In the last decades, much effort has been made to increase the knowledge of ferns and lycophytes in Pernambuco through collections carried out in the four phytogeographic zones of the state: “Zona do Litoral”, “Zona da Mata”, “Zona das Caatingas”, and “Zona das Savanas” (Andrade Lima 2007). In this study, we follow IBGE (2012) to describe the vegetation of the areas, namely, Mangroves and “Restinga” in the “Zona do Litoral”; variations of Seasonal and Ombrophilous Forests in the “Zona da Mata”; and variations of Steppe-Savanna in the “Zona das Caatinga” and “Zona das Savanas” (IBGE 2012; Andrade Lima 2007). We adopted the term “Brejo de Altitude” or “Brejos Nordestinos” (Andrade Lima 2007) for areas of Montane Seasonal Semideciduous Forest and of Montane Ombrophilous Dense Forest without connections with the coastal forest. This term has been widely used in the context of the northeastern Atlantic Forest and in some cases, “Brejos de Altitude” encompass areas of ecological tension (see Pôrto et al. 2004; Ferraz & Rodal 2006).

Morphological data from the analyzed herbarium specimens, field observations, and the literature (Rödl-Linder 1990; Lellinger 2002; Smith et al. 2006; Labiak & Prado 2008; Schwartsburd & Smith 2013; Sanín 2018; Sanín et al. 2023) were used to build an identification key with descriptions, illustrations, field pictures, and comments for each species. Maps were prepared in ArcMAP version 10.4 (2023) using the coordinates of the collection sites of the specimens available in the data sources and when the coordinates were not available, they were inferred using the Google Maps geocoding service.

Results and Discussion

Five species of Serpocaulon were recorded in Pernambuco, distributed from 0 to 1,100 m in 23 (less than 12%) of the 185 municipalities of the state (Fig. 1). The genus was absent in the “Restinga” and Mangroves of the “Zona do Litoral”; was represented by only one species, S. triseriale (Sw.) A.R. Sm., in the “Zona das Caatingas”; and was represented by the five species in the “Zona da Mata”, mostly in “Brejos de Altitude”, which are therefore important areas for the study and conservation of the genus. The occurrence and comparatively high abundance of Serpocaulon in this phytogeographic zone is likely associated with the high number of studies conducted in the Atlantic Forest (Stehmann et al. 2009) and also the typically high humidity of these ecosystems (Sanín et al. 2023).

Figure 1
Distribution of Serpocaulon species from Pernambuco state - a. location of Brazil in South America; b. silhouette of Pernambuco state; c. record of Serpocaulon species in relation to Federal protected areas (green shape). Yellow circle = S. catharinae; black square = S. menisciifolium; red circle = S. polystichum; full triangle = S. richardii; white dotted circle = S. triseriale.

Although Serpocaulon triseriale is the most widely distributed species of the genus (Smith et al. 2006; Sanín 2018; Sanín et al. 2023), it was present in only twelve of the 185 municipalities of the state and was not recorded in the west region of the state, where it was expected to occur (Xavier et al. 2012). The occurrence of Serpocaulon attenuatum (Humb. & Bonpl. ex Willd.) A.R. Sm. was also expected in Pernambuco state, because the species has been found in the neighboring states (Sanín et al. 2023).

In this study, we confirm the presence of S. polystichum in Pernambuco. This species was not reported from Brazil according to Smith et al. (2006). Schwartsburd (2023) considered S. polystichum as a synonym of S. fraxinifolium, but we agree with Smith et al. (2006) and Sanín et al. (2023) who consider them as two distinct species. Sanín et al. (2023) supported that specimens identified as S. fraxinifolium must represents S. polystichum in Brazil. Additionally, we report the occurrence of S. richardii in the state of Pernambuco. This species has been frequently identified in Brazil as S. adnatum (Kunze ex Klotzsch) A.R. Sm. (e.g., Zuquim et al. 2008, 2009; Góes- Neto & Pietrobom 2012; Schwartsburd & Smith 2013). S. adnatum is an Andean species that also has a long-creeping rhizome, pinnate laminae, and adnate pinnae at the apex (Sanín et al. 2023).

The species Serpocaulon catharinae and S. menisciifolium presents their northernmost distribution limit in Pernambuco (Fig. 1). This information is important for the conservation of these species because specimens at the boundaries of the species distribution range are key for population genetic studies (Barbosa et al. 2012).

None of the five species of Serpocaulon found in Pernambuco occur within sustainable use or integral protection federal conservation units (CUs). However, they occur in privately managed areas, such the Frei Caneca Private Natural Heritage Reserve (RPPN) (in the municipality of Jaqueira) and Pedra D’antas RPP (Lagoa dos Gatos) both sustainable use CUs; state-managed areas such as the Matas do Siriji Wildlife Refuge (RVS) (São Vicente Férrer) and Dois Irmãos State Park (Recife), both integral protection CUs and the Serras e Brejos do Capibaribe Environmental Protection Area (APA) (Brejo da Madre de Deus and Taquaritinga do Norte) a sustainable use CU. The Atlantic Forest in Pernambuco has been currently reduced to less than 2% of its original cover (Capobianco 2001) and most of the remnants are still vulnerable to anthropic pressure (Santiago, personal observation).

Serpocaulon A.R. Sm. Taxon 55(4): 928. 2006. Figs. 1-4

Figure 2
Pinnae/segments of Serpocaulon species from Pernambuco state - a. segment of S. catharinae with rounded/truncate apex; b. pinna of S. menisciifolium showing pubescence and one row of sori; c. segment of S. polystichum with four rows of areoles and sori; d. pinna of S. richardii showing pubescence and four rows of sori; e. pinna of S. triseriale with four rows of areoles and tree of sori. [a. D. Sanín & A. Santiago 4194 (BHCB); b. I.C.L. Barros 36 et al. (PEUFR); c. D. Sanín & A. Santiago 4197 (BHCB); d. D. Sanín & Santiago 7267 (BHCB); e. D. Sanín & A. Santiago 4195 (BHCB)]. Scale bars = 1 cm.

Figure 3
a-g. Field pictures of Serpocaulon species from Pernambuco state - a. S. catharinae; b. S. menisciifolium; c. S. polystichum; d-f. S. richardii - d. lamina; e. apex rhizome; f. adnate pinna; g. S. triseriale. [a. D. Sanín & A. Santiago 4194 (BHCB); b. D. Sanín & A. Santiago (BHCB); c. D. Sanín & A. Santiago 4197 (BHCB); d-f. D. Sanín & A. Santiago 7267 (BHCB); g. D. Sanín & A. Santiago 7267 (BHCB)].

Figure 4
a-e. Rhizome scales from Serpocaulon species from Pernambuco state - a. subulate scale of S. catharinae; b. wide ovate-lanceolate scale of S. menisciifolium; c. narrow ovate-lanceolate scale of S. polystichum; d. subulate scale of S. richardii; e. lanceolate scale of S. triseriale. [a. D. Sanín & A. Santiago 4194 (BHCB); b. I.C.L. Barros 36 et al. (PEUFR); c. D. Sanín & A. Santiago 4197 (BHCB); d. D. Sanín & Santiago 7267 (BHCB); e. D. Sanín & A. Santiago 7267 (BHCB)].

Epiphytic and terrestrial, rhizomes short (phyllopodia < 2 times the rhizome width apart) to long-creeping (phyllopodia 2 or more times the rhizome width apart), pruinose or not, rhizome scales clathrate, dense or scattered, patent, lanceolate to subulate, bicolorous or concolorous, dark brown to reddish, pale or iridescent, rounded base and acuminate to long tapering apex. Laminae pinnate or pinnatisect, when pinnate 5-15 pairs of pinnae, when pinnatisect 28-44 pairs of segments, linear to ovate-lanceolate, papyraceous to coriaceous, proximally reflexed, distally gradually tapering (S. catharinae) or with a conform pinna apex, base of the pinna or segments decurrent, sessile to adnate (S. richardii and rarely S. triseriale), glabrous to densely pubescent (in juvenile specimens of S. menisciifolium and S. richardii), venation regularly anastomosing (goniophlebioid) with individual chevron-shaped areoles and each with a single, free, included excurrent veinlet, 1-5 rows of sori between the costa and the pinnae or segments margins.

Key for species of Serpocaulon in Pernambuco

• 1. Laminae pinnatisect 1. Serpocaulon catharinae

• 1’. Laminae pinnate 2

  • 2. Rhizome scales subulate with reflexed apex 4. Serpocaulon richardii

  • 2’. Rhizome scales lanceolate with straight apex 3

    • 3. Rhizome long-creeping 3. Serpocaulon polystichum

    • 3’. Rhizome short-creeping 4

      • 4. Rhizome scales bicolorous, middle pinnae adnate 2. Serpocaulon menisciifolium

      • 4’. Rhizome scales concolorous, middle pinnae sessile 5. Serpocaulon triseriale

1. Serpocaulon catharinae (Langsd. & Fisch.) A.R. Sm. Taxon 55(4): 928. 2006. Polypodium catharinae Langsd. & Fisch. Pl. Voy. Russes Monde 9, pl. 9. 1810. Type: Brazil. Insula Catharinae, G.H. Langsdorff 12 (lectotype, designated by Hensen [1990: 292], LE [bc] 00000033 image!). Figs. 1, 2a, 3a, 4a

Epiphytic, rhizome short-creeping, pruinose; scales patent, subulate, disperse, bicolorous, brown whitish to the margin, dark brown to the center, opaque, base rounded, apex long acuminate and reflexed. Laminae pinnatisect, linear, base truncate, apex gradually tapering; segments 28-44 pairs, chartaceous, glabrous, base decurrent, apex rounded; sori 1 row between the costa and the segment margin.

Examined material: Amaraji, Engenho Animoso, 25.VI.1999, I. Fernandes 1451 (JPB). Bonito, Mata da Reserva Biológica Municipal, 08º30’30.5”S, 35º43’18.2”W, 750-800 m, 11.VII.2000, A. Santiago 222 (PEUFR). Caruaru, aprox. 450 m, 10.IX.1971, Andrade-Lima 71-6724 (IPA); Mata do Brejo dos Cavalos, 16.VII.1988, I.C.L. Barros 1988 (PEUFR); Brejo dos Cavalos, 1,100 m, 08º18’36”S, 36º00’00”W, 3.VIII.1995, M.R.C. Sales de Melo 143 (PEUFR). Jaqueira/Lagoa dos Gatos, Serra do Urubu, trilha do Mosquito, 672 m, 08º42’2”S, 35º51’10”W, 3.XII.2014, L.S.B. Calazans 467 (BHCB); Mata da Colônia, 800 m, 08º30’14”S, 35º42’56”W, 800 m, 29.XI.2000, A. Santiago & M. Pietrobom-Silva 343 (PEUFR). Maraial/Lagoa dos Gatos, Serra do Urubú, 19.IV.1994, A.M. Miranda 1538 (PEUFR). Tapera, Mata do Toró, 3.III.1974, 50 m, I. Pontual 74-1271 (PACA-AGP).

The species is registered in Lowland Semideciduous Seasonal Forest; Lowland, Submontane, and Montane Ombrophilous Dense Forest; “Brejos de Altitude”, 50-1,100 m.

Serpocaulon catharinae is widely distributed species in Brazil (Labiak & Prado 2008). Specimens from the NAF present narrow laminae with several segments.

Hensen (1990) designate a lectotype from LE without link to any specific herbarium specimen. After revising the protologue (Langsdorff & Fisch 1810, Fig. 9) and comparing the digital specimens at LE, where three plants are found in the same sheet, Sanín et al. (2023) found that the line drawing in the protologue matches the right-corner plant of the specimen L. Langsdorff 12 (LE) in size, shape, number of segments, and sori in the middle pinna. Since Langsdorff & Fisch (1810) did not associate this name with any specific collection, Sanín et al. (2023) designate the specimen L. Langsdorff 12 (LE) as the lectotype of this species.

2. Serpocaulon menisciifolium (Langsd. & Fisch.) A.R. Sm. Taxon 55(4): 928. 2006. Polypodium menisciifolium Langsd. & Fisch. Pl. Voy. Russes Monde 11, pl. 12. 1810. Type: Brazil. “Insula St. Catharina”, G.H. Langsdorff 15 (lectotype, designated by Hensen [1990: 304], LE [bc] 00000044 image!).

Serpocaulon hirsutulum (T. Moore) Schwartsb. & A.R. Sm., J. Bot. Res. Ins. Texas 7(1): 90. 2013. Polypodium hirsutulum T. Moore, Index. Fil. (T. Moore) 390. 1862, nom. nov. for Polypodium hirsutulum Raddi, Pl. Bras. Nov. Gen. 21, t 29, fig 2. 1825, nom. illeg. (non G. Forst 1786). Type: Brazil. “Super palmarum annosarum truncus inviciniis Urbis Rio-Janeiro” [On trunk of aged palms in the vicinity of the city of Rio de Janeiro], Nov 1817-May 1818, Raddi s.n. (lectotype, designated by Schwartsburd & Smith, [2013: 88], PI [bc] 010870 image!). Figs. 1, 2b, 3b, 4b

Epiphytic, rhizome short-creeping; scales patent, ovate-lanceolate, densely covering the rhizome, bicolorous, reddish, iridescent, brown to the center and dark brown to the margin, base rounded, apex acute-acuminate. Laminae pinnate, lanceolate, base truncate, apex conform; pinnae 12-13 pairs, chartaceous, rarely coriaceous, densely pubescent, base adnate, acroscopically softly auriculate, apex acute; sori 1-2 rows between the costa and pinna margin.

Examined material: Bonito, Mata da Colonia, 800 m, 08º30’14”S, 35º42’56”W, 19.VI.1998, I.C.L. Barros et al. (PEUFR); Mata da Chuva, 750 m, 08º32’20.8”S, 35º43’22.3”W, 7.V.1999, I.C.L. Barros et al. (PEUFR). Jaqueira, Usina Colônia, Mata do Ageró, 415 m, 18.X.2001, M.S. Lopes & M.R.S. Pietrobom 404 (RB). Lagoa dos Gatos, RPPN Pedra D’antas, R.P. Farias et al. 150 (UFP). Maraial/Lagoa dos Gatos, Serra do Urubu, 22.V.1998, I.C.L. Barros 36 et al. (PEUFR).

These records represent the northernmost limit of the species distribution range. The species is registered in Submontane and Montane Ombrophilous Dense Forest, and “Brejos de Altitude”, 415-800 m.

Serpocaulon hirsutulum (Raddi) Schwartsb. & A.R. Sm. was segregated from S. menisciifolium by Schwartsburd & Smith (2013) suggesting that small plants with densely pubescent laminae characterize this taxon. However, we agree with Brade (1951) and Hensen (1990) that proposed P. hirsutulum Raddi as a synonym of P. meniscifolium (= S. menisciifolium) by observed that the presence of pubescence is related with the age of the plants.

3. Serpocaulon polystichum (Link) A.R. Sm. Taxon 55(4): 928. 2006. Polypodium polystichum Link, Hort. Berol. 2: 101. 1833. Type: Brazil. (Cult. in bot. gard. at Berlin), J.H.F. Link s.n. (Lectotype, designated by Hensen [1990: 297], B [bc] 200163713 image!). Figs. 1, 2c, 3c, 4c

Epiphytic or terrestrial, rhizome long-creeping; scales patent, ovate-lanceolate, scarcely covering the rhizome, bicolorous, brown to the margin, dark brown to the center, opaque, base rounded, apex acuminate to long acuminate and patent to rarely reflexed. Laminae pinnate, ovate-lanceolate, base truncate, apex conform; pinnae 9-10 pairs, chartaceous, glabrous, proximally sessile to softly adnate apically, base decurrent, apex acuminate; sori 2-4 rows between the costa and pinna margin, non-impressed.

Examined material: Jaqueira, Usina Colônia, Serra do Quengo, 713 m, 08º42’50.4”S, 35º50’25.8”W, 3.IV.2002, M.S. Lopes & M.R.S. Pietrobom 529 (UFP). Taquaritinga do Norte, Sítio Cafundó, 10.I.1977, I. Pontal 77-1438 (PEUFR); alto da serra ou Engenho Brejinho, 21.XI.1977, Andrade-Lima 77-8321 (IPA).

The species is registered in Montane Ombrophilous Dense Forest and “Brejos de Altitude”, at 713 m.

Serpocaulon polystichum has been frequently confounded in Brazil with S. fraxinifolium (Labiak & Prado 2008; Schwartsburd & Smith 2013; Sanín et al. 2023), an Andean species described based on a cultivated specimen from Schönbrunn, Austria, but collected in Venezuela (Hensen 1990). Like S. polystichum, S. fraxinifolium has long-creeping rhizomes and pinnate laminae. However, S. polystichum can be distinguished from S. fraxinifolium by its patent rhizome scales (vs. appressed) and non-impressed sori at the adaxial surface of the pinnae (vs. impressed).

4. Serpocaulon richardii (Klotzsch) A.R. Sm. Taxon 55(4): 929. 2006. Polypodium richardii Klotzsch, Linnaea 20: 394. 1847. Type: English Guyana. R. Schomburgk 1651 (lectotype, designated by Hensen [1990: 306], B, isotype frag. NY). Figs. 1, 2d, 3d-f, 4d

Epiphytic or terrestrial, rhizome long-creeping; scales patent, subulate, scarcely covering the rhizome, bicolorous, brown to the margin, reddish to the center, opaque, base rounded, apex long linear tapering, reflexed. Laminae pinnate, ovate-lanceolate, base truncate, apex conform; pinnae 4-7(-9) pairs, papyraceous to rarely chartaceous, densely pubescent, base barely sessile to completely adnate, apex long acuminate; sori (1-)3-5 rows between the costa and pinna margin.

Examined material: Bonito, margem estrada para Camocim, 21.I.1970, Andrade-Lima 70-5657 (IPA); Reserva Biológica, 21.XI.1994, L.P. Felix 7003 (PEUFR); Mata da Azuada, relitos a mao dereita deçendo, 709 m, 08º30’8.6”S, 35º41’49.9”W, 16.III.2019, D. Sanín & A. Santiago 7267 (BHCB); Mata da Colônia, 800 m, 08º30’14”S, 35º42’56”W, 14.VII.2000, A. Santiago 218 (PEUFR); Mata da Chuva, 750 m, 08º32’20”S, 35º43’22”W, 8.V.2001, A. Santiago & M.R. Pietrobom-Silva 453 (PEUFR); Mata da Reserva Biológica municipal de Bonito, 750-800 m, 08º30’30”S, 35º43’18”W, 19.V.2000, A. Santiago et al. 149 (PEUFR). São Vicente Ferrer, Complexo do Maciço Serra do Mascarcenhas, Mata do Estado, 640 m, 07º35’00”S, 35º30’00”W, 20.IV.1998, M.R. Pietrobom-Silva 4229 (PEUFR). Vicência, 8.VII.1974, I. Pontual (PEUFR).

The species is registered mainly in “Brejos de Altitude”, but it is also recorded in Submontane Ombrophilous Dense Forest, 480-800 m.

This species has been frequently misidentified in Brazil as S. adnatum (Smith et al. 2006; Zuquim et al. 2008, 2009; Coelho & Esteves 2011; Góes-Neto & Pietrobom 2012; Schwartsburd 2023), an Andean species distributed from Colombia and Venezuela to Peru (Sanín et al. 2023). Smith et al. (2006) suggested the occurrence of S. richardii in Brazil. In a later work, Brazil was not mentioned by Smith et al. (2018) in the distribution range of the species. By his part, Schwartsburd & Smith (2013) suggested that Brazilian plants identified as S. adnatum should be better referred to as S. richardii, but they did not cite any specific collection to exemplify this assertion. Later, Schwartsburd (2023) reinforced the occurrence of S. richardii in Brazil, but did not cite specimens from Pernambuco.

Serpocaulon richardii has long-creeping rhizomes and pinnate laminae with distally adnate pinnae, as does S. adnatum. However, it can be distinguished from S. adnatum by the subulate rhizome scales with reflexed apex (vs. ovate rhizome scales and appressed apex), and densely pubescent laminae (vs. glabrous). Serpocaulon richardii can also be confounded with S. psychotrium Mostacero, D. Sanín & A.R. Sm. because they share long-creeping rhizomes with subulate scales and pinnate laminae with distally adnate pinnae, but S. richardii can be distinguished from S. psychotrium by the densely pubescent laminae (vs. glabrous laminae) and the papyraceous to chartaceous laminae with non-impressed sori (vs. chartaceous laminae with impressed sori).

Serpocaulon richardii presents its southernmost distribution in Pernambuco and Bahia states (Sanín et al. 2023).

5. Serpocaulon triseriale (Sw.) A.R. Sm., Taxon 55(4): 929. 2006. Polypodium triseriale Sw. J. Bot. (Schrader) 2: 26. 1800 [1801]. Goniophlebium triseriale (Sw.) Wherry. Amer. Fern. J. 54(3): 144. 1964. Type: West Indies. Anonym s.n. (lectotype designated by Sanín et al. [2023: 432], UPS [bc] 24589 image!). Figs. 1; 2e; 3g; 4e

Epiphytic or terrestrial, rhizome short-creeping; scales patent, lanceolate, densely covering the rhizome, concolorous, light brown, opaque, base rounded, apex narrowly acute-acuminate. Laminae pinnate, ovate-lanceolate, base truncate, apex conform; pinnae (5-)7-9(-15) pairs, coriaceous to rarely chartaceous, glabrous, base barely sessile to barely adnate, apex acute to acuminate; sori (1-)2-3(-5) rows between the costa and pinna margin.

Examined material: Amaraji, Matas do Engenho Animoso, 2.XII.1999, C.P.L. Luna et al. 21 (UFP). Bezerros, Parque Municipal de Serra Negra, 2.VI.1995, M.R.C. Sales de Melo 63 (PEUFR); Pedra Pintada, Serra Negra de Bezerros, 800 m, 08º10’S, 35º47’W, 13.VIII.1999, L. Krause & A. Liebig 33 (PEUFR). Bonito, Mata da Chuva, 750 m, 08º32’20”S, 35º43’22”W, 19.V.2000, A. Santiago et al. 166 (PEUFR); Mata da Reserva Biológica Municipal de Bonito, 750-800 m, 08º30’30”S, 35º43’18”W, 10.V.2000, A. Santiago et al. 149 (PEUFR); Mata da Colônia, 800 m, 08º30’14”S, 35º42’56”W, 19.V.2000, A. Santiago et al. 148 (PEUFR). Cabo de Santo Agostinho, Reserva de Gurjaú, VIII.1991, K. Duarte (UFP). Cortez, 10.XI.1974, Andrade-Lima (IPA). Goiana, Eng. Itapirema do Meio, 12.X.1966, Andrade-Lima 66-4728 (IPA). Gravatá, Fazenda Harmonia, 10.X.1970, Andrade-Lima 70-6043 (IPA). Jaqueira, Usina, Colônia (Ageró), 415 m, 08º44’27”S, 35º50’37”W, 20.V.2002, M.S. Lopes & M.R.S. Pietrobom 613 (UFP). Brejo da Madre de Deus, Biruti Grande, Mata do Biturizinho, 1,000 m, 15.IX.1992, K.M.R. Santos (PEUFR). Maraial, Lagoa dos Gatos, Serra do Urubú, 22.V.1998, I.C.L. Barros 34 (PEUFR). Recife, Várzea, mata de São João, 11.V.2001, M.D. Santos et al. 5 (PEUFR); Dois Irmãos, área do Açude do Prata, Jardim Zoobotânico, 1983, 20 m, A Souto 04 (UFP). Vicência, Engenho Jundiá, Mata do Engenho Jundiá, 487 m, 07º37’20”S, 35º19’40”W, 5.III.2002, A.M.R. Carvalho et al. 55 (UFP). Tacaratu, Serra do Morcego, 835 m, 04º18’80”S, 38º7’35”W, 15.III.2011, J.A. Siqueira-Filho 2468 (HVASF).

This is the most widely distributed species of the genus (Smith et al. 2006; Sanín et al. 2023). It is recorded in all forest formations and “Caatinga” (Steppe-Savanna) of Pernambuco, 0-1,000 m.

Serpocaulon triseriale is found in several herbaria of Brazil as Polypodium brasiliense Poir., a synonym of S. triseriale (Smith et al. 2006; Sanín et al. 2023).

Serpocaulon triseriale can be confused with S. rex Schwartsb. & A.R. Sm. and S. attenuatum due to the impressed venation and lanceolate and patent rhizome scales. However, it is distinguished from S. rex by its short-creeping rhizome, 1-4 rows of sori between the costa and the margin, smooth margin of the pinnae, and flat and connected verrucae ornamentation of the spore (vs. long-creeping rhizome, 4-6 rows of sori between the costa and the margin, crenate margin of the pinnae, and prominent and not connected verrucae ornamentation of the spore) (Sanín et al. 2019). On the other hand, S. triseriale can be distinguished from S. attenuatum by its pinnate laminae with 1-4 rows of sori between the costa and the margin (vs. pinnatisect laminae and 1-2 rows of sori between the costa and the margin) (Sanín et al. 2019).

Serpocaulon triseriale has been registered in Pernambuco around roots of Vriesea procera Mez (Andrade-Lima 66-4728, IPA), and as an epiphyte on palm trees (E.S. Santana 87, PEUFR).

Acknowledgements

We would like to thank the curators and the staff of the consulted herbaria, especially Rita de Cássia Araújo Pereira (IPA), Maria Elizabeth Bandeira-Pedrosa (PEUFR), and Marlene Barbosa (UFP), for kindly allowing us to access the botanical collections. We thank the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior and the Universidade Federal de Minas Gerais, for the scholarship provided to DS; the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), for the research grants (proc. 306868/2014-8, 201414/2017-1) provided to AS; and the Fundação de Amparo à Pesquisa do Estado de Minas Gerais (Fapemig), for providing financial support (APQ-03041-17). We are sincerely grateful to reviewers and the editor in chief of this volume Claudine Mynssen, for their critical reading of the manuscript and suggestions.

Data availability statement

In accordance with Open Science communication practices, the authors inform that all data are available within the manuscript.

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Edited by

Area Editor: Dra. Claudine Mynssen

Publication Dates

• Publication in this collection
  01 Dec 2023
• Date of issue
  2023

History

• Received
  23 Mar 2023
• Accepted
  18 Aug 2023