Goniopteris C.Presl is characteristic of wet lowland forests in the Neotropics, with centers of diversity in the Greater Antilles, the eastern slope of the Andes, and southeastern Brazil. It harbors 138 species (Fawcett & Smith, 2021), or about four times as many as the average-sized fern genus, which contains about 33 species (Schuettpelz et al., 2018). Molecular phylogenetic studies have resolved Goniopteris as monophyletic (Smith & Cranfill, 2002; Schuettpelz & Pryer, 2007; Almeida et al., 2016; Fawcett et al., 2021). It bears one particularly distinctive morphological synapomorphy that supports its monophyly: hairs that are bifurcate (1-forked) or more branched apically (Christensen, 1913). No other genus of Thelypteridaceae has such hairs (Salino & Semir, 2002; Fawcett & Smith, 2021).

In his monumental and ground-breaking monograph, Christensen (1913) had three key insights about Goniopteris: 1) stellate and furcate hairs are a defining characteristic of Goniopteris; 2) G. macrotis (Hook.) Pic. Serm. and G. semihastata (Kunze) Salino & T. E. Almeida are members of Goniopteris despite the lack of such hairs; and 3) these two species, along with Goniopteris jamesonii (Hook.) Salino & T. E. Almeida form a “natural group.” We believe this natural group also includes G. yanachagae, supported by the molecular phylogenetic analyses of Fawcett et al. ( 2021). Other unsampled species, especially in South America, may also belong in this group. Although molecular data are lacking, G. deltata does have the characteristic furcate hairs of most Goniopteris species. Like G. yanachagae, it has no obvious close relatives, based on morphology.

Recently, many new combinations have been made in Goniopteris (Salino et al., 2015), and new species have been described (e.g., by Salino, 2002; Caluff & Sánchez, 2004; Krömer et al., 2007; Smith & Kessler, 2008; Matos et al., 2010; Salino et al., 2014, 2016; Moura et al., 2016; Fawcett, 2020). We report here on two more new species found during our herbarium work. Both are known from single gatherings, but they are so distinct that we do not hesitate to describe them as new.

Goniopteris deltata S.E.Fawc., R.C.Moran & A.R.Sm., sp. nov.—Type: Ecuador, Morona-Santiago: along road between Santiago and Río Morona, 33.7 km east of Santiago, steep slope with virgin forest, 02°58′55”S, 077°48′43”W [−2.9819400°Lat., −77.8119400°Long.], 523 m, 10 Jul 2004, Thomas B. Croat, Lynn P. Hannon, Gregory A. Wahlert & Tuntiak Katan Jua 90,688 (holotype: QCNE [!]; isotypes, MO barcode 177348 and accession 5850489 [!]; QCA? [n.v.]). (Fig. 1.)

Fig. 1
figure 1

Goniopteris deltata. A. Habit. B. Detail of suprabasal pinna. C. Indusiate sorus. D. Stalked stellate and furcate hairs of the abaxial surface of rachis. [From Croat 90,688 (MO).]

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Diagnosis.—Differs from its congeners by the following features: laminae deltate, pinnatifid in the distal half, leaf axes puberulent abaxially, with minute (0.1 mm long), apically 1–4-forked hairs, but lacking (also abaxially) stiff acicular (non-forked) hairs, the venation with one pair of basal veins from adjacent segments uniting to produce an excurrent vein running to the base of the sinus, and the surfaces of the indusia sparsely pubescent.

Plants terrestrial; rhizomes short-creeping, with leaves approximate; leaves monomorphic, to 52 × 25 cm; petioles tan to light brown, puberulent throughout, the hairs 0.1 mm long, 1-forked apically, erect, proximal portions of petioles scaly, the scales 2–3 × 0.5–1 mm, dull brown, lanceolate, entire, puberulent on both surfaces and margins, the hairs ca. 0.1 mm long, mostly 1-forked, but rarely with 3 or 4 arms at the tip of the hair; rachises adaxially grooved and with short simple and furcate hairs to 1 mm, abaxially glabrous or nearly so; laminae to 25 × 25 cm, deltate (basal pinnae the longest), proximally 1-pinnate with up to 4 pairs of free pinnae, distally pinnatifid, the pinnae adnate, decurrent, slightly curved toward the apices, becoming obtuse in distal segments, margins glabrous; aerophores present, visible as faint lunate depressions on the rachises at the pinna bases, sometimes slightly darker than the surrounding tissue; buds absent; rachises tan to light brown, puberulent on both surfaces, the hairs ca. 0.1 mm, 1- to 4-forked, erect; basalmost pinnae to 12 × 2.5 cm, equilateral, lobed ca. 1/4 to the costules, their base narrowed, petiolulate, the petiolules ca. 2 mm long; suprabasal pinnae with a short basal basiscopic lobe, widest at their bases; costae and costules sparsely puberulent abaxially, the hairs ca. 0.1 mm, 1–3-forked, erect, on the adaxial surfaces of the costae puberulent, the hairs 0.1 mm long, acicular (non-furcate), antrorse, the adaxial surfaces of the costules glabrous; veins in 5–7 pairs per segment, the basal pair from adjacent segments united at an acute angle with an excurrent vein to the sinus, 1 to 3 suprabasal veins running to the excurrent vein, or free, with 1 or 2 veins running to the sinus, more distal veins meeting the segment margins above the sinus, glabrous on both surfaces, on the basal pinnae with the first vein on the basiscopic side of the costule, on suprabasal pinnae, on the acroscopic side; laminar tissue between the veins not verrucose, glabrous on both surfaces; sori medial, 1 mm diam., indusiate, the indusia ca. 0.5 mm diam. (when dry), round-reniform (with a narrow invagination proximally), dark reddish brown, sparsely pubescent on the surface, with a few hairs ca. 0.1 mm long, acicular (not branched or forked); sporangial capsules and stalks glabrous.

Distribution and habitat.—Ecuador, Amazonian side of the Andes; known only from the type.

Etymology.—The specific epithet refers to the distinctive broadly deltate laminae.

Discussion.—Goniopteris deltata most resembles G. abrupta (Desv.) A.R.Sm., G. jamesonii (Hook.) Salino & T.E.Almeida, and G. macrotis (Hook.) Pic.Serm. These species have similar leaf size, laminae often widest at or near the base, similar pinna shape and lobing, and indusiate sori. Collectively, all occur in western Amazonia, from Colombia to Bolivia and in western Brazil (Smith, 1983, 1992; Smith & Kessler, 2017). The new species differs from the other three by the distal portion of the laminae with decidedly adnate pinnae or segments, especially on the basiscopic sides of the pinnae or segments. The first species, G. abrupta, differs from G. deltata by leaf axes abaxially with a mixture of acicular hairs 0.1–0.3 mm long and furcate or stellate hairs ca. 0. 1 mm long, pinna pairs 5–10(−12) and incised 1/3–1/2 their width, veins free, and a bud often present in the axil of a distal pinna.

The second species, Goniopteris jamesonii, differs from G. deltata by pinna bases with a conspicuous acroscopic lobe, leaf axes abaxially pubescent with hairs of mixed lengths 0.1–1.0 mm long, and laminar tissue between the veins adaxially pubescent with appressed hairs. Goniopteris jamesonii occurs from Ecuador to Bolivia and southern Brazil (Smith, 1983, 1992; Smith & Kessler, 2017).

The third species, Goniopteris macrotis, differs from G. deltata and nearly all other species in the genus by lacking stellate or furcate hairs; instead, it bears only stiff acicular hairs 0.3–1.5 mm long (Smith, 1992). It further differs by rhizomes erect to suberect, pinnae 0.7–1.6(−2) cm wide, buds present distally along the rachises, and laminar tissue between the veins pubescent with acicular, sometimes appressed hairs. It is known only from Peru (Smith, 1992).

Goniopteris yanachagae S.E.Fawc., R.C.Moran & A.R.Sm., sp. nov.—Type: Peru, Pasco: Oxapampa, Huancabamba, Parque Nacional Yanachaga–Chemillén, Huampal sector, across the river from the entrance/guard station, along path leading down to the big rapids, locally common, in sheltered nooks of dryish rock wall, with mosses, etc., −10.18259°Lat., −75.57367°Long., 977 m, 20 May 2016, Carl Rothfels, J.Larraín & T.Pócs 4991 (holotype: HOXA [!]; isotype: UC accession no. 2062003 [!]. (Fig. 2.)

Fig. 2
figure 2

Goniopteris yanachagae. A. Habit. B. Medial pinna. C. Detail of medial pinna. [From Rothfels 4991, (UC).]

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Diagnosis.—Differs from all other members of the genus by the following combination of character states: hairs simple (with no forked or stellate hairs), veins free, mostly simple, not anastomosing to form an excurrent veinlet to the sinus, and laminae pinnate-pinnatifid.

Plants saxicolous; rhizomes about 4 mm wide, forming a small caudex with leaf-bases tightly clustered, the scales dark-castaneous, opaque, entire, with abundant hyaline acicular hairs on both surfaces, hairs to 0.3 mm long; leaves monomorphic; petioles stramineous, to 5 cm long, with spreading hyaline acicular hairs to 0.8 mm long and, proximally with dark castaneous opaque scales, 1.5–2.5 × 0.2–0.6 mm, these with hyaline acicular hairs ca. 0.3 mm long on both surfaces; rachises stramineous, adaxially grooved and with abundant spreading hairs to 2 mm, these hyaline, some with reddish tips, abaxially with similar indument; laminae drying green, thinly chartaceous, lanceolate, 8–25 cm long, 2–4.5 cm wide, pinnate-pinnatifid, with 10–16 pinna pairs, leaf apex gradually attenuate and not conform to the lateral pinnae; aerophores and buds lacking; pinnae short-petiolulate to about 1 mm, with up to 7 lobes, incised 2/3 to 3/4 towards costae, the basal two or three pinna pairs gradually reduced, the smallest about 1/2 the size of the largest pinna, reflexed; costae and costules pale, prominent adaxially and abaxially, and with hyaline acicular hairs; veins free, simple to occasionally forking, with up to six pairs per segment, the basal pairs from adjacent segments reaching the segment margins above the sinuses; laminae between the veins adaxially glabrous, abaxially with indument of abundant, spreading hyaline acicular hairs to 0.8 mm long, these flattened and sometimes twisted, relatively uniform in length; sori round, ca. 1 mm diam., exindusiate, subcostular to inframedial; sporangia capsules and stalks glabrous; spores tan, winged.

Distribution and ecology.—Peru, Amazonian side of the Andes, known only from the type locality in the Parque Nacional Yanachaga-Chemillén, where it grows in a rock wall near rapids.

Etymology.—The specific epithet refers to the type locality and only known station, in the National Park Yanachaga-Chemillén.

Discussion.—For more than 100 years (Christensen, 1913), Goniopteris has been defined by the presence of stellate or furcate hairs, which are unique in the family. These characteristic hairs, however, have been lost in few species, one of which is Goniopteris yanachagae. The others lacking stellate or furcate hairs are the Mesoamerican G. clypeata (Maxon & C.V.Morton) Salino & T.E.Almeida, G. mollis Fée (synonym: Thelypteris ghiesbreghtii (Hook.) C.V.Morton), and the South American G. macrotis (Hook.) Pic.Serm. and G. semihastata (Kunze) Salino & T.E.Almeida. These last two species are resolved together in a primarily Andean clade with G. lugubriformis (Rosenst.) Salino & T.E.Almeida, G. lugubris (Mett.) Brade, G. pinnatifida (A.R.Sm.) Salino & T.E.Almeida, and G. tryoniorum (A.R.Sm.) Salino & T.E.Almeida (Fawcett et al., 2021). This clade is in turn sister to G. yanachagae (treated as “Goniopteris sp. Pasco” by Fawcett et al., 2021).

Goniopteris yanachagae differs from most species of the genus by the absence of stellate or furcate hairs (vs. present), and free veins (vs. at least one pair of veins anastomosing below the sinus to form and excurrent vein (Presl, 1836; Fée, 1852)). Because of its small size, free veins, exclusively simple hairs, and pinnate-pinnatifid laminae, G. yanachagae is readily distinguished from other Andean members of the genus. In the Greater Antilles, many similarly diminutive saxicolous species of Goniopteris occur, some of which also have simple, free veins, and several have simple hairs (e.g., G. abdita (Proctor) Salino & T.E.Almeida, G. minutissima (Caluff & C.Sánchez) Salino & T.E.Almeida), but stellate hairs are also present in these. Based on molecular phylogenetic evidence, these species are distantly related to this Andean clade.

The closest relative of Goniopteris yanachagae is unknown, and its phylogenetic position is on a long branch, sister to a diverse clade (Fawcett et al., 2021). Morphologically it most resembles G. semihastata, with which it shares its small stature, lack of stellate or furcate hairs, laminae abaxially strigose, and proximal pinnae reflexed and gradually reduced towards the base. The type localities for both species, as well as that of the simple-haired G. macrotis, are on the eastern side of the Peruvian Andes.

Goniopteris yanachagae resembles Amauropelta and shares with it the following traits: reduced proximal pinnae, free veins, and simple hairs. However, it has winged spores, whereas those of Amauropelta have broadly folded, fenestrate or perforate perines (Alvarez-Fuentes, 2010; Patel et al., 2019).

The type locality for Goniopteris yanachagae, in the Cordillera de Yanachaga, is part of the sub-Andean Cordilleran region, a series of discontinuous mountain chains east of the central Andean chain, and separated from it by a series of valleys (Missouri Botanical Garden, 2008). The National Park Yanachaga-Chemillén, established in 1986 (Young & León, 1999), forms part of the core area of the Oxapampa-Ashaninka-Yanesha United Nations Biosphere Reserve, designated in 2010. This region includes the largest expanse of intact remaining rainforest in Peru and hosts extraordinary biodiversity including many local endemics, owing partly to its varied topography (ranging from 300 m to 4500 m), climatic stability, and diversity of habitats (UNESCO 2016). Recent and continuing study of the biodiversity of the area has led to the discovery of many new species, including flowering plants (Judziewicz & Tyrrell, 2007; Pringle, 2017; Roque & León, 2006; Santamaría-Aguilar & Lagomarsino, 2015; Daly et al., 2020), invertebrates (Razowski & Wojtusiak, 2010), and vertebrates (Lehr et al., 2007, 2012; Venegas et al., 2011), and doubtless many others are yet to be described.