Amauropelta platensis (Thelypteridaceae), a new combination for the endemic Southern Cone fern flora

Abstract

Athyrium decurtatum var. platense (= Thelypteris decurtata subsp. platensis) is transferred to the genus Amauropelta and raised to the species level based on morphological, palynological and geographical evidences. This fern species is endemic to the Delta of Paraná district, belonging to the Esteros del Iberá biogeographic province in the Southern Cone of South America.

Key words:
Amauropelta; Argentina; endemism; ferns; Uruguay

Resumo

Athyrium decurtatum var. platense (= Thelypteris decurtata subsp. platensis) é transferida para o gênero Amauropelta e elevada ao nível de espécie com base em estudos morfológicos, palinológicos e geográficos. Esta espécie é endêmica do distrito Delta do Paraná, pertencendo à província biogeográfica dos Esteros del Iberá, no Cone Sul da América do Sul.

Palavras-chave:
Amauropelta; Argentina; endemismo; samambaias; Uruguay

Thelypteridaceae is one of the largest families of ferns, cosmopolitan in distribution with more than 1,200 species highly diverse morphologically and ecologically (Fawcett & Smith 2021), inhabiting mainly tropical and subtropical regions of the world (Smith 1992; Smith et al. 2006, 2008; PPG I 2016). As usually construed, the family is monophyletic and recent molecular phylogenetic studies have recovered two main groups: the phegopteroid and the thelypteroid lineages (Smith & Cranfill 2002; He & Zhang 2012; Almeida et al. 2016; Fawcett & Smith 2021), which are recognized at subfamily level (PPG I 2016). In the subfamily Thelypteridoideae (thelypteroid lineage), the amauropeltoid clade comprises the genera Amauropelta Kunze, related to the genera Coryphopteris Holttum, and Metathelypteris (H.Itô) Ching (Almeida et al. 2016; Fawcett & Smith 2021; Fawcett et al. 2021). Amauropelta Kunze is a genus morphologically circumscribed by the base of the lamina gradually or abruptly reduced, with one-to-many pairs of auriculiform or glanduliform pinnae, the veins from adjacent segments usually meeting at the blade margins above the sinuses on pinnatifid pinnae, and sporangia either rarely or not setose (Ponce 1995; Salino et al. 2015; Ponce & Zanotti 2018; Fawcett & Smith 2021). Amauropelta is characterized also by the spore wall with reticulate ornamentation (Ponce 1987). According to Fawcett & Smith (2021), Amauropelta is distinguished from Metathelypteris (eastern Asia) by characters as veins running to the margins, adaxially grooved costae and a base chromosome number of x = 27, 29, 31 (vs. usually x = 35 in Metathelypteris). From Coryphopteris, Amauropelta differs by the combination of usually greatly reduced proximal pinnae, lack of sessile, resinous, reddish glands on the lamina between veins (except in the A. resinifera group) and usually creeping or suberect rhizomes (vs. upright and trunklike).

Amauropelta comprises more than 200 species mainly distributed in the Neotropical region, with 21 of them growing in Argentina (Ponce 1987, 1998, 2016; Ponce & Zanotti 2018; Yañez et al. 2022). Recent molecular phylogenetic studies in Thelypteridaceae, along with morphological data, support the monophyly of Amauropelta (Smith & Cranfill 2002; Alvarez-Fuentes 2010; Almeida et al. 2016; Fawcett et al. 2021). In this genus, species delimitation is complicated, probably due to recent diversification and hybridization events (Almeida et al. 2016).

In the Neotropics, the Delta del Paraná district, belonging to the Esteros del Iberá biogeographic province, is an area of endemism that promotes speciation processes (Arana et al. 2021). This district covers the estuary and the islands of the Paraná and Uruguay Rivers, and associated fluvial formations in the Río de la Plata coastal plains. Various endemic taxa of ferns develop in this area, such as Goniopteris burkartii Abbiatti, a member of Thelypteridaceae, among other endemic taxa of plants, fungi and animals (Arana et al. 2021).

An overlooked endemic taxon from Delta of Paraná district was originally described by Weatherby as Athyrium decurtatum var. platense Weath., quite different from Athyrium decurtatum (Kunze) Fée due to its “indusiisque minute pallideque glandulosis aliter glabris” (Weatherby 1946). The type specimen of A. decurtatum var. platense was collected by A. Burkart in river-bank woods that develop along the estuary of the Río de La Plata. The reason why the taxon was first included in Athyrium Roth was that the sori are “more or less athyrioid”. Later, de la Sota (1983) demonstrated that the morpho-anatomical and chromosomal features of A. decurtatum coincided with those of the Thelypteridaceae, particularly its chromosomal number x = 29, placing the taxon within Thelypteris subgenus Amauropelta. According to Fawcett & Smith (2021), this chromosomal number is an apomorphy of Amauropelta subgenus Amauropelta. Then, Athyrium decurtatum was transferred to the genus Thelypteris, and in the same work, the variety described by Weatherby was raised to a subspecific level, as Thelypteris decurtata subsp. platensis (Weatherby) de la Sota.

The molecular phylogenetic reconstruction of the Thelypteridaceae (Almeida et al. 2016; Fawcett et al. 2021) confirmed the monophyly of numerous subgenera of Thelypteris, which were elevated to genera, as occurred with Amauropelta. Thereby, Thelypteris decurtata (Link) de la Sota was transferred to Amauropelta by Salino et al. (2015). However, the necessary combination of Thelypteris decurtata subsp. platensis within the genus Amauropelta is missing. To rectify this taxonomic issue, we provide the combination and raise the endemic taxon occurring in the Delta del Paraná district to species level, based on morphological characteristics of the sporophyte (including palynological features), along with biogeographical data. Our decisions are based on clear morphological and geographical discontinuities that define two easily distinguishable species of Amauropelta in Argentina and Uruguay. The species delimitation used here follows the morphological cluster species concept (Mallet 1995), in combination with a very distinctive geographical range of distribution.

Nomenclatural decisions for the name treated are based upon the analysis of protologues, related bibliography, and herbarium samples lodged at BA, GH, LP, RCVC, and SI, including types. Also, living specimens of Amauropelta decurtata were collected in Yungas of Nevados del Aconquija at the banks of Cochuna River (27°19’S, 65°53’W), Chicligasta Department, in Tucuman province (Argentina), whereas those of Thelypteris decurtata subsp. platensis were collected at Punta Lara Nature Reserve, Partido de Ensenada (34º47’33’’S, 58º00’28.32’’W), located on the banks of La Plata River, 12 km north of La Plata city (Province of Buenos Aires, Argentina) (Fig. 1a). Features such as the morphology of the rhizome scales, indument morphology and distribution in the laminae, sori, and indusia were examined under a Nikon SMZ1000 stereoscopic microscope and a Nikon E200 light microscope (LM). Scales from the whole sporophyte were mounted in 20% glycerin without prior treatment. The spores of T. decurtata subsp. platensis were analyzed and described in previous work by Giudice et al. (2014). To carry out a complete study of this taxon, new fresh samples were observed with a scanning electron microscope (SEM). Materials were mounted on aluminium stubs without prior treatment and covered with gold for 10 minutes. Observations were made in a microscope FEI Quanta 200 model (Servicio de Microscopía Electrónica de Barrido y Microanálisis del LIMF, Facultad de Ingeniería, Universidad Nacional de La Plata) with a 15 kV voltage. The palynological terminology used in the descriptions refers to Tryon & Lugardon (1991) and Lellinger (2002). All the microscopic preparations are preserved in the Laboratory of Comparative Anatomy, Propagation and Conservation of Embryophytes “Dr. Elías R. de la Sota”, Faculty of Natural Sciences and Museum, National University of La Plata.

Figure 1
a-g. Amauropelta platensis diagnostic characteristics - a. living specimen inhabiting Punta Lara Nature Reserve; b. light microscope (LM) micrograph of a rhizome scale; c. a glandular hair in detail; d. abaxial surface of a pinna showing indusia (arrows) and scattered glandular hairs; e. an indusium in detail (the arrow indicates an orange glandular hair); f. LM micrograph of the indusium where the papillate hairs (arrowheads) can be seen on the margin and a glandular hair (gl) (arrow); g. SEM micrograph of a spore. gl = glandular hair; in = indusium. (a-f. from Berrueta, Ponce & Luna 320; g. from Giudice, Ramos Giacosa & Luna 110).

Amauropelta platensis (Weath.) Ponce & Arana, comb. et stat. nov. Basionym: Athyrium decurtatum var. platense Weatherby, Amer. Fern J. 36: 95. 1946.

Thelypteris decurtata subsp. platensis (Weath.) de la Sota, Lilloa 36(1): 65. 1983.

Type: ARGENTINA. BUENOS AIRES: Partido Ensenada: Río Santiago, “monte a la orilla del Río de la Plata (circa de La Plata)”, XI.1931, A. Burkart 3681 (holotype GH 00075931!; isotype SI 000042!). Paratypes: ARGENTINA. BUENOS AIRES: Partido Berazategui: Arroyo Las Conchitas, monte ribereño del Rio de la Plata, 16.IV.1927, A. Burkart 1309 (GH, SI!); Delta del Paraná, Carabelas, I.1931, Pérez Moreau s.n. (GH, BA 589!), Pérez Moreau s.n. (GH, BA 3998!), A. Burkart 4327 (GH, SI!).

Terrestrial (Fig. 1a); rhizomes erect, scaly apices, scales brown, narrowly triangular or subulate, clathrate, with glandular hairs and papillae (Fig. 1b-c); fronds polystic, 0.50-1 m; petioles stramineous, 1/3 of the total length of the frond, glabrous or papillate, with scales at the base similar those to the rhizome; laminae elliptic in outline, 10-25 cm wide, abruptly tapering at the base, herbaceous, rachis stramineous, glabrous or papillate; pinnae distal and medial linear-triangular, basal pinnae elliptic or subelliptic, 6-15 cm × 1.5-2 cm, with 2-4 pairs of reduced basal pinnae, the smallest up to 3 mm, costae glabrous or with minute papillae; segments linear, oblique, 2-3 mm wide, with acute or obtuse apex, margin entire or crenulate; with 6-9 pairs of veins, adaxial side glabrous, abaxial side with few sessile, yellowish to orange glands; sori elliptic, medial; indusium (Fig. 1d-f) developed on one side of the vein (“athyrioid”), sometimes with a basal extension on both sides (“subathyrioid”) or, more rarely, developed on both sides of the vein (reniform), glabrous or with sessile, yellowish to orange glands (Fig. 1e), margin with single-celled, whitish or hyaline papillae (Fig. 1f); spores light brown, perispore plegate-reticulated with folds or ridges (Fig. 1g), of 25-28 µm in polar diameter and 35-40 µm in equatorial diameter, with straight laesura, 23-28 µm long.

Aditional specimens examined: ARGENTINA. BUENOS AIRES: Partido Ensenada, Reserva Punta Lara, 19.XII.2012, Giudice, Ramos Giacosa & Luna 110 (LP); sendero el burrito, 15.V.2017, Berrueta, Ponce & Luna 320 (LP); Río Santiago, 9.X.1906, Pastore 110 (SI); 28.X.1909; Isla Santiago, 30.IV.1932, Cabrera 2162 (LP, SI); Partido Tigre, Tigre, Isla de Forbes, 28.XII.1902, Hicken 22 (SI); Delta del Paraná, Arroyo Gambado, 19.XII.1955, Burkart 19970 (SI); isla sobre Río Sarmiento, entre arroyos Curubica y Otamendi, cerca del recreo “Las Rosas II”, 4.IV.2009, Hurrell, Ulibarri & Bazzano 6845, 6848 (SI). Partido San Fernando, Delta del Paraná, Canal Gobernador Arana, 27.XI.1932, Burkart 5068 (SI). ENTRE RÍOS: Islas del Ibicuy, Brazo Largo, Campo Ichauspe, Rosillo 376 (SI); Delta del Paraná, Río Ceibo, 24.XI.1932, Burkart 5117 (SI). URUGUAY. COLONIA. Barra de San Juan, Doello (BA).

Amauropelta platensis is distinguished from Amauropelta decurtata (Fig. 2 a-b) by the indusia glabrous with whitish or hyaline unicellular glandular hairs on the margins (Fig. 1d-f; Fig. 2d) and some unicellular glandular orange hairs on the surface (Fig. 1d-f) versus indusia with acicular and glandular orange hairs on the whole surface (Fig. 2b), and absence of unicellular whitish glandular hairs on the margins in A. decurtata. Also A. platensis lacks acicular hairs on the lamina, rachis and costae (Fig. 2c), which are characteristic on both surfaces of the fronds of A. decurtata (Fig. 2a).

Figure 2
a-d. Segments of Amauropelta platensis and A. decurtata - a. abaxial surface of last segments of A. decurtata, showing indument, venation and disposition of sori; b. detail of sorus showing the indument of the indusium; c. abaxial surface of last segments of A. cisplatina, showing indument, venation and disposition of sori; d. detail of sorus. (a-b. from Burkart 5165 (SI); c-d. from Burkart 5117).

Field observations made on living plants, together with the examination of range-wide herbarium specimens, led us to determine the existence of many novel distinctive features for Amauropelta platensis. The species is characterized by rhizome scales with pluricellular, glandular, orange hairs on margins (Fig. 1b-c) (vs. rhizome scales with acicular hairs or glabrescent in A. decurtata); laminae without acicular hairs (vs. laminae with acicular hairs on both sides in A. decurtata, Fig. 2a); rachis and costae glabrous or papillate (vs. rachises and costae pilose, or glabrescent, with ephemeral, subulate hairy scales in A. decurtata). In Amauropelta platensis the indusia surfaces are glabrous or with some scattered orange glandular hairs similar to those of the laminae, and the margins have whitish or hyaline, papillate unicellular hairs (Fig. 1d-f; Fig. 2d) (vs. indusia with unicellular acicular hairs and orange, sessile spherical glandular hairs in A. decurtata (Fig. 2b). The spores are similar in size to the both species, but the spores of A. platensis have perispore plegate-reticulated (Giudice et al. 2014) (Fig. 1g), vs. perispore reticulated with some cristae in A. decurtata.

Amauropelta platensis has a very restricted distribution area, growing exclusively in Northeastern Buenos Aires and Southern Entre Ríos provinces in Argentina, and also in the nearer coastal places of Uruguay. In contrast, Amauropelta decurtata has a much wider distribution, with a disjunct pattern in tropical Andes and Southern Brazil (Fig. 3). Particularly in Argentina, it grows in North-western and North-eastern areas, in the provinces of Corrientes, Jujuy, Misiones and Tucumán (de la Sota 1983; Ponce 2016).

Figure 3
Complete distribution map of Amauropelta platensis and A. decurtata.

Amauropelta platensis is apparently endemic to the Delta of Paraná district, which occupies part of Buenos Aires and Entre Ríos Provinces in Argentina, and also the opposite coast of the Uruguay river, in the Oriental Republic of Uruguay (Fig. 3). In this area, a variety of ecological niches develop, as flooded savannas of Cephalanthus glabratus (Spreng.) K. Schum. (Rubiaceae), Cyperus byssaceus Pereira-Silva, Schoenoplectus californicus (C.A. Mey.) Soják (Cyperaceae), and Luziola peruviana Juss. ex J.F. Gmel. (Poaceae); intermixed with wetland vegetation and gallery forests of Tessaria integrifolia Ruiz & Pav. (Asteraceae), Myrsine laetevirens (Mez) Arechav. (Myrsinaceae), Nectandra angustifolia (Schrad.) Nees & Mart. (Lauraceae), Syagrus romanzoffiana (Cham.) Glassman (Arecaceae), Inga uraguensis Hook. & Arn. (Fabaceae), Blepharocalyx salicifolius (Kunth) O. Berg (Myrtaceae) and Sambucus australis Cham. & Schltdl. (Viburnaceae) along the rivers (Arana et al. 2021). These ecosystems dissect the grassland and xerophytic forest of the Pampa plains with riverine scrubs, gallery forest, tropical-subtropical megathermic swamp grasslands and particular aquatic communities (Arana et al. 2021). Historically cited for Delta del Paraná and Río Santiago area on the coast of the Río de La Plata (de la Sota 1983; Ponce 1987), Amauropelta platensis was discovered inhabiting the gallery forests of Punta Lara Natural Reserve, Buenos Aires Province (Giudice et al. 2014).

The genus Amauropelta has traditionally been considered a natural (probably monophyletic) subgenus within Thelypteris Schmidel (Ponce 1987, 1995; Smith 1974, 1983, 1990, 1992). This also occurs with other neotropical and paleotropical taxa, i.e. the subgenus Cyclosorus. Recent molecular studies (e.g., Almeida et al. 2016 and Fawcett et al. 2021) has demonstrated that the genus Amauropelta is a monophyletic group. Thus, these authors propose to rehabilitate the generic range established in previous classifications (i.e., Holttum 1971; Smith & Cranfill 2002), criteria followed by PPG I (2016). Within Amauropelta and following Fawcett & Smith (2021), A. platensis belongs to Amauropelta subgenus Amauropelta because the proximal pinnae typically gradually reduced, x = 29 (de la Sota 1983), and rhizomes typically erect, and > 3 mm diameter.

The often-subtle morphological characters needed to distinguish genera and to identify species have contributed to the reputation of Thelypteridaceae as a taxonomically difficult lineage (Fraser-Jenkins et al. 2017). This is particularly true in Amauropelta, where e.g., 12 of the 54 Bolivian species of Amauropelta are known only from one to three collections (Smith & Kessler 2017). This likely reflects a combination of factors, including narrow endemism, rarity, paucity of herbarium collections, and failure of general collectors to recognize subtle variations in the field (Fawcett et al. 2021). In the present work, Amauropelta platensis is designated as a distinct entity from Amauropelta decurtata, because of the diagnostic features of the sporophytes already reported, combined with the restricted geographic distribution of A. platensis, which occurs only in the estuary of the Río de la Plata, and nearer coasts of Paraná and Uruguay rivers. In this study, we found only two specimens of A. decurtata from Entre Ríos Province in the Ibicuy Islands Department (Río Bravo, Burkart 1238, SI, and Concordia, Duraznal, Castellanos s.n., BA 31/859) between more than 50 examined collections. These specimens could be treated as accidental because they may have been transported south by the periodic floodings, as was previously hypothesized by de la Sota (1983). This phenomenon occurs with a several plants and animals in the Esteros del Iberá biogeographic province (Arana et al. 2021), which represents a rare case of a transboundary system that has not yet been fragmented by dams, and still retains its regular flood pulses and free connectivity between the main channels and their vast floodplains (Baigún & Minotti 2021). Also, Amauropelta decurtata has a wide disjunct distribution tropical-Andean and Austro-Brazilian and Northern Uruguayan, in the Departments of Cerro Largo and Tacuarembó (Fig. 3). According to de la Sota (1983) this type of distribution, combined by the morphological variations observed in the different populations covering the entire range of A. decurtata may be due to this taxon could be, in fact, a species complex.

Below we provide a key to identify the species studied.

Identification key to the species of Amauropelta studied here

1. Lamina pilose on both surfaces, with simple, acicular hairs and unicellular spherical, orange glandular hairs on the abaxial surface. Indusia pilose, with acicular hairs and spherical, sessile, orange, glandular hairs throughout the adaxial surface and margins. Spores 24-30 µm in polar diameter × 34-52 µm in equatorial diameter, perispore reticulated with some cristae. Disjunct distribution, tropical Andes and Southern Brazil Amauropelta decurtata

1’. Laminae lacking acicular hairs, having unicellular spherical, orange glandular hairs on abaxial surface. Indusia glabrous on the adaxial surface, rarely with some scattered orange, spherical glands, with whitish or hyaline unicellular papillate hairs on the margins. Spores 25-28 µm in polar diameter × 35-40 µm in equatorial diameter, perispore plegate-reticulated. Restricted distribution area, growing exclusively in the marginal forest of the Delta del Paraná district in Argentina and Uruguay Amauropelta platensis

Acknowledgements

We want to thank the staff of Reserva Natural Punta Lara, Buenos Aires, Argentina, for their help during field trips. We are grateful to P. Albornoz, G. Romagnoli, R. Delgado, M. Catania, M. Taboada and E. Bulacio (Fundación Miguel Lillo, Tucumán), for their hospitality and technical assistance during field trips in Tucumán. We also want to acknowledge to Carlos Lehn (Instituto Federal de Educação, Ciência e Tecnologia Farroupilha), for his assistance with the Portuguese abstract. We are also very grateful with the editors and reviewers, whose comments improved the manuscript. This study was supported by the Research Projects of Universidad Nacional de La Plata, Argentina 11N/850 (Directora G. E. Giudice) and 11N/940 (Directora M. L. Luna) and FONCYT (PICT 02227, Directora M. L. Luna).

Data availability statement

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

References

• Almeida TE, Hennequin S, Schneider H, Smith AR, Batista JANA, Ramalho AJ, Proite K & Salino A (2016) Towards a phylogenetic generic classification of Thelypteridaceae: additional sampling suggests alterations of neotropical taxa and further study of paleotropical genera. Molecular Phylogenetics and Evolution 94: 688-700. DOI: https://doi.org/10.1016/j.ympev.2015.09.009
  » https://doi.org/10.1016/j.ympev.2015.09.009
• Alvarez-Fuentes O (2010) The systematics of the genus Amauropelta (Pteridophyta: Thelypteridaceae) in the Caribbean Islands. UMI Dissertation Publishing - ProQuest LLC, Ann Arbor. 361p. DOI: https://doi.org/doi:10.25335/4tws-1r52
  » https://doi.org/10.25335/4tws-1r52
• Arana MD, Natale E, Oggero A, Ferreti N, Romano G, Martínez G, Posadas P & Morrone JJ (2021) Esquema biogeográfico de la República Argentina. Opera lilloana 56: 1-240.
• Baigún CRM & Minotti PG (2021) Conserving the Paraguay-Paraná fluvial corridor in the XXI century: conflicts, threats, and challenges. Sustainability 13: 5198. DOI: https://doi.org/10.3390/su13095198
  » https://doi.org/10.3390/su13095198
• de la Sota ER (1983) Sobre la ubicación de Athyrium decurtatum (Link) Fée (Athyriaceae-Pteridophyta). Lilloa 36: 59-68.
• Fawcett S & Smith AR (2021) A generic classification of the Thelypteridaceae. Sida, Botanical Miscellany 59. Botanical Research Institute of Texas Press, Fort Worth. 114p.
• Fawcett S, Smith AR, Sundue M, Burleigh JG, Sessa EB, Kuo L-Y, Chen C-W, Testo W, Kessler M, GoFlag Consortium & Barrington DS (2021) A global phylogenomic study of the Thelypteridaceae. Systematic Botany 46: 891-915. DOI: 10.1600/036364421X16370109698650
  » https://doi.org/10.1600/036364421X16370109698650
• Fraser-Jenkins CR, Gandhi KN, Kholia BS & Benniamin A (2017) An annotated checklist of Indian pteridophytes. Part 1: Lycopodiaceae to Thelypteridaceae. Bishen Singh Mahendra Pal Singh, Dehra Dun. 562p.
• Giudice GE, Ramos Giacosa JP & Luna ML (2014) Registro de Thelypteris decurtata (Link) de la Sota ssp. platensis (Weath.) de la Sota en la Reserva Natural Punta Lara, provincia de Buenos Aires: un aporte para la conservación de helechos nativos de distribución restringida. Historia Natural 4: 105-110.
• He L-J & Zhang X-C (2012) Exploring generic delimitation within the fern family Thelypteridaceae. Molecular Phylogenetics and Evolution 65: 757-764. DOI: https://doi.org/10.1016/j.ympev.2012.07.021
  » https://doi.org/10.1016/j.ympev.2012.07.021
• Holttum RE (1971) Studies in the family Thelypteridaceae III. A new system of genera in the Old World. Blumea 19: 17-52.
• Lellinger DB (2002) A modern multilingual glossary for taxonomic pteridology. Pteridologia 3: 1-263. DOI: https://doi.org/10.5962/bhl.title.124209
  » https://doi.org/10.5962/bhl.title.124209
• Mallet J (1995) A species definition for the modern synthesis. Trends in Ecology & Evolution 10: 294-299. DOI: https://doi.org/10.1016/0169-5347(95)90031-4
  » https://doi.org/10.1016/0169-5347(95)90031-4
• Ponce MM (1987) Revisión de las Thelypteridaceae (Pteridophyta) argentinas. Darwiniana 28: 317-390.
• Ponce MM (1995) Las especies austrobrasileñas de Thelypteris subg. Amauropelta (Thelypteridaceae, Pteridophyta). Darwiniana 33: 257-283.
• Ponce MM (1998) Citas nuevas en Thelypteris subg. Amauropelta (Thelypteridaceae, Pteridophyta) para la flora argentina. Darwiniana 35: 177-178.
• Ponce MM (2016) Thelypteridaceae. In: Ponce MM & Arana MD (coord.) Flora vascular de la República Argentina. Vol. 2. IBODA, Buenos Aires. Pp. 353-384.
• Ponce MM & Zanotti CA (2018) Amauropelta opposita (Thelypteridaceae), nueva cita para la flora Argentina. Darwiniana, nueva serie 6: 113-119. DOI: http://dx.doi.org/10.14522/darwiniana.2018.61.796
  » http://dx.doi.org/10.14522/darwiniana.2018.61.796
• PPG I (2016) A community-based classification for extant ferns and lycophytes. Journal of Systematics and Evolution 54: 563-603. DOI: https://doi.org/10.1111/jse.12229
  » https://doi.org/10.1111/jse.12229
• Salino A, Almeida TE & Smith AR (2015) New combinations in Neotropical Thelypteridaceae. Phytokeys 57: 11-50. DOI: https://doi.org/10.3897/phytokeys.57.5641
  » https://doi.org/10.3897/phytokeys.57.5641
• Smith AR (1974) A revised classification of Thelypteris subgenus Amauropelta. American Fern Journal 64: 83-95.
• Smith AR (1983) Polypodiaceae - Thelypteridoideae [Family 14 (4)]. In: Harling G & Sparre B (eds.) Flora of Ecuador. Vol. 18. Swedish Research Council, Stockholm. Pp. 1-148.
• Smith AR (1990) Thelypteridaceae. In: Kubitzki K, Kramer KU & Green PS (eds.) The families and genera of vascular plants Vol. I. Pteridophytes and Gymnosperms. Springer-Verlag, Berlin, Heidelberg, New York. Pp. 263-272.
• Smith AR (1992) Thelypteridaceae. In: Tryon RM & Stolze RG (eds.) Pteridophyta of Peru. Vol. III. Fieldiana, Botany, n. s. 29: 1-80.
• Smith AR & Cranfill R (2002) Intrafamilial relationships of the theypteroid ferns (Thelypteridaceae). American Fern Journal 92: 131-149.
• Smith AR & Kessler M (2017) Prodromus of a fern flora for Bolivia. XXX. Thelypteridaceae. Phytotaxa 331: 1-34. DOI: 10.11646/phytotaxa.331.1.1
  » https://doi.org/10.11646/phytotaxa.331.1.1
• Smith AR, Pryer KM, Schuettpelz E, Korall P, Schneider H & Wolf PG (2006) A classification for extant ferns. Taxon 55: 705-731. DOI: 10.2307/25065646
  » https://doi.org/10.2307/25065646
• Smith AR, Pryer KM, Schuettpelz E, Korall P, Schneider H & Wolf PG (2008) Fern classification. In: Ranker TA & Haufler CH (eds.) Biology and evolution of ferns and lycophytes. Cambridge University Press, Cambridge. Pp. 417-467. DOI: 10.1017/cbo9780511541827.017
  » https://doi.org/10.1017/cbo9780511541827.017
• Tryon AF & Lugardon B (1991) Spores of the Pteridophyta. Springer-Verlag, New York. 658p. DOI: https://doi.org/10.1007/978-1-4613-8991-0
  » https://doi.org/10.1007/978-1-4613-8991-0
• Weatherby CA (1946) A variety of Athyrium decurtatum from Argentina. American Fern Journal 36: 94-95. <https://doi.org/10.2307/1545143>.
  » https://doi.org/10.2307/1545143
• Yañez A, Marquez GJ, Ocampo Terraza D & Ponce M (2022) Amauropelta yabotiensis (Thelypteridaceae), A new species from Biosphere Yabotí Reserve (Misiones, Argentina) and its taxonomic relationships. Anais da Academia Brasileira de Ciencias 94: e20201870. DOI: 10.1590/0001-3765202220201870
  » https://doi.org/10.1590/0001-3765202220201870

Edited by

Area Editor: Dra. Lana Sylvestre

Publication Dates

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

History

• Received
  03 Apr 2023
• Accepted
  21 June 2023