Goniopteris
Creators: name: Fawcett, Susan
name: Smith, Alan R.
Publisher: name: BRIT Press
Year: 2021
Subjects: Biodiversity
Taxonomy
Plantae
Tracheophyta
Polypodiopsida
Polypodiales
Thelypteridaceae
Goniopteris

description: GONIOPTERIS
Goniopteris C. Presl, Tent.Pterid. 181–183, pl. 7, f. 9–11.1836.— Thelypteris subg. Goniopteris (C.Presl) Duek,Adansonia,n.s., 11:720. 1971.— Thelypteris sect. Goniopteris (C. Presl) C.V. Morton, Amer. Fern J. 53(4):154. 1963.
— LECTOTYPE (designated by J. Smith, Hist. Fil. 191. 1875): Goniopteris crenata (Sw.) C.Presl [= G. poiteana (Bory) Ching]
For additional generic synonymy, see Christensen (1913, 1920), Proctor (1985, 1989).
Etymology. —Gr. gonia, angle + pteris, fern, in reference to the angle of often acutely anastomosing veins below the sinus or these producing an excurrent vein joined by other connivent veins; this contrasts to the often obliquely united veins of Meniscium.
Plants terrestrial or epilithic, small to medium-sized (5–80 cm); rhizomes typically stout, short-creeping, ascending, or erect, scales generally with stellate or furcate hairs on surfaces and/or margins; fronds erect, arching, pendent, or prostrate, monomorphic, or weakly dimorphic (e.g., Goniopteris tetragona), typically pinnate-pinnatifid, rarely simple or pinnate-pinnatisect; stipes stramineous to dull brown, grooved adaxially; stipe scales brown, deltate-ovate to linear-lanceolate, typically with furcate or stellate hairs on surfaces and/or margins; blades membranaceous to thickly coriaceous, often drying dark green, apex conform or gradually reduced, base of blade unreduced, or, if reduced, without many pairs of gradually reduced pinnae, proliferous buds sometimes present adaxially along rachises, in axils of pinnae, sometimes also along costae (e.g., G. alata); pinnae petiolulate, sessile, or narrowly adnate (especially distally), margins simple, crenate to deeply lobed, bases dilated, truncate, rounded, hastate, or with acroscopic auricles; veins anastomosing, at an angle with veins from adjacent segments, forming an excurrent veinlet to the sinus, and creating one to a series of many areoles between costae and pinna margins (e.g., G. tetragona), or veins simple, free or forked in smaller 1-pinnate spp. (e.g., G. abdita); aerophores absent, rarely tuberculate at pinna bases (G. lugubriformis); indument abaxially and adaxially of stipitate or sessile, branched hairs, these furcate (T- or Y-shaped) or stellate (Fig. 2B), hairs occasionally bifurcate or multifurcate, branched hairs sometimes co-occurring with unicellular or pluricellular hyaline acicular hairs typical of other genera, branched hairs most easily observed along the adaxial groove of rachis, but sometimes also present on all axes and laminar tissue adaxially and abaxially, glands generally absent on axes and lamina in most spp. (except G. redunca), scales usually absent, except along costae abaxially in a few South American species; pustules rarely present (e.g., G. juruensis); sori round, typically medial, indusiate or exindusiate, discrete, indusia bearing hairs, these either acicular or variously branched (Fig. 2F); sporangia glabrous or with setulae (simple or branched); spores brown, consistently ornamented with reticulate crests (Patel et al. 2019a); x = 36, with 25 species counted, diploids and tetraploids, and a single triploid (G. obliterata) known. No intergeneric hybrids have been reported, but intrageneric hybrids are quite common, especially among the species of the calciphilic Antillean radiation (Smith 1993b), where more than a dozen have been reported in Cuba alone (Sánchez 2017).
Diagnosis. —The nearly invariable presence of stellate or branched hairs somewhere on the plant, especially along the axes and on stipe base and rhizome scales, is the most diagnostic feature of the genus, readily distinguishing it from all other genera in the family. The paleotropical Ampelopteris is similar in bearing proliferous buds and stellate hairs (see illustrations in Holttum et al. 1970), but the hairs are easily abraded, and less conspicuous than in Goniopteris. Meniscium differs in having exindusiate sori that are frequently coalescent, many more pairs of anastomosing veins, usually entire or only serrate pinnae, and often longer-creeping rhizomes. Steiropteris often differs in the presence of well-developed aerophores at the pinna (and sometimes costule) bases, and in most cases, a distinct cartilaginous, non-vascularized keel at the bases of the sinuses; it also lacks stellate or furcate hairs.
Biogeography and ecology. —Numbering 138 species, Goniopteris is strictly neotropical, ranging north to Florida, Mexico, through Mesoamerica and the Antilles, throughout Amazonia, and the foothills of the Andes, south to Bolivia and Argentina, and east through Brazil, the Guianas and Venezuela. They are predominantly plants of lower elevations, ranging to 1850 m in the Andes (Smith 1992), and to 1700 m in the Antilles. They are typically plants of shady forest understories, but a diverse clade of calciphiles, numbering ~30 species, has radiated in the karst regions of the Greater Antilles (Fawcett 2020). Two serpentine specialists, G. crypta and G. brittoniae, have likely independently arisen from calciphilic ancestors on Cuba, and Puerto Rico, respectively. As with many members of the family, they respond favorably to disturbance, may become weedy, and may be especially abundant on open roadcuts and in coffee and cacao plantations.
Taxonomic and phylogenetic studies. —Historically treated as a subgenus of Dryopteris, the circumscription of species of Goniopteris has remained essentially unchanged for more than a century (Christensen 1913, 1920). Christensen’s treatment was the first to rely on close observation of microscopic features, and he united the species of Goniopteris largely on the basis of their striking stellate or branched hairs. Although the hairs have been secondarily lost on rare occasions (e.g., G. macrotis), the few taxa without them were still included by Christensen (1913) on the basis of other morphological similarities; their placement has now been supported by molecular data (Fawcett et al. in press). Christensen (1913, 1920) divided Dryopteris subg. Goniopteris into two sections— Asterochlaena and Eugoniopteris — recognized by apices gradually reduced, or by a conform terminal pinna, respectively. Phylogenetic analyses have shown these subdivisions to be non-monophyletic, and therefore artificial. Necessary combinations have been made for nearly all known members of the genus (Salino et al. 2015).
Molecular phylogenetic data support the monophyly of Goniopteris (Almeida et al. 2016; Patel et al. 2019a), and show strong biogeographical signal, with distinct radiations in the Antilles, the Andes, and Brazil (Fawcett et al. in press). The genus is in an early-diverging clade of cyclosoroids, and sister to a clade that includes the neotropical genus Meniscium, the pantropical Cyclosorus s.s., the African and Southeast Asian Ampelopteris, and the Malesian Mesophlebion (Fawcett et al. in press). In contrast, in the less densely-sampled, mostly plastid dataset of Patel et al. (2019a), Goniopteris is inferred be sister to the aforementioned genera, plus the remaining cyclosoroids.
Most necessary combinations were made by Salino et al. (2015), except:
Constituent species.—* Goniopteris abdita (Proctor) Salino & T.E. Almeida; * G. abrupta (Desv.) A.R. Sm.; G. affinis Fée; * G. alan-smithiana (L.D. Gómez) Salino & T.E.Almeida; * G.alata (L.) Ching; * G.amazonica (Salino & R.S. Fern.) Salino & T.E. Almeida; * G. ancyriothrix (Rosenst.) Salino & T.E. Almeida; G. anoptera (Kunze ex Kuhn) Salino & T.E. Almeida; G. asterothrix Fée; G. aureola (A.R. Sm.) Salino & T.E. Almeida; * G. baorucensis S.E. Fawc.; * G. beckeriana (F.B. Matos, A.R. Sm. & Labiak) Salino & T.E. Almeida; G. berlinii (A.R. Sm.) Salino & T.E. Almeida; G. bermudiana (Baker) S.E. Fawc. & A.R. Sm.; * G. bibrachiata (Jenman) Salino & T.E. Almeida; * G.biformata (Rosenst.) Salino & T.E. Almeida; * G. biolleyi (Christ) Pic.Serm.; * G. blanda (Fée) Salino & T.E. Almeida; * G. bradei Salino; * G. brittoniae (Sloss.) Ching; * G. burkartii Abbiatti; G.calypso (L.D. Gómez) Salino & T.E.Almeida; G. chocoensis (A.R.Sm.& Lellinger) Salino & T.E. Almeida; G. clypeata (Maxon & C.V. Morton) Salino & T.E. Almeida; * G. cordata (Fée) Salino & T.E. Almeida; G. costaricensis Salino & T.E. Almeida; G. crassipila (Caluff & C. Sánchez) Salino & T.E. Almeida; * G. croatii (A.R. Sm.) Salino & T.E. Almeida; G. crypta (Underw. & Maxon) Ching; * G. cumingiana (Kunze) de Jonch. & U. Sen; G. cuneata (C.Chr.) Brade; * G. curta (Christ) A.R. Sm.; G. cutaitaensis (Brade) Brade; G. dissimulans (Maxon & C. Chr.) Salino & T.E. Almeida; * G. eggersii (Hieron.) Alston; G. equitans (Christ) Salino & T.E. Almeida; G. erythrothrix (A.R. Sm.) Salino & T.E. Almeida; * G. francoana (E. Fourn.) Á. LÖve & D. LÖve; * G. fraseri (Mett. ex Kuhn) Salino & A.R. Sm.; G. fuertesii (Brause) S.E. Fawc., A.R. Sm. & Y.Y. Piña; * G. gemmulifera (Hieron.) Vareschi; G. glochidiata (Mett. ex C. Chr.) Brade; ** G. goeldii (C. Chr.) Brade; * G. gonophora (Weath.) Salino & T.E. Almeida; * G. hastata Fée; * G. hatchii (A.R. Sm.) Á. LÖve & D. LÖve; * G. hildae (Proctor) Salino & T.E. Almeida; * G. holodictya (K.U. Kramer) Salino & T.E. Almeida; G. hondurensis (L.D. Gómez) Salino & T.E. Almeida; G. iguapensis (C. Chr.) Brade; * G. imbricata (Liebm.) Á. LÖve & D. LÖve; * G. imitata (C. Chr.) Salino & T.E. Almeida; G. indusiata (Salino) Salino & T.E. Almeida; * G. jamesonii (Hook.) Salino & T.E. Almeida; G. jarucoensis (Maxon ex Caluff & C. Sánchez) Salino & T.E. Almeida; * G. juruensis (C. Chr.) Brade; G. killipii (A.R. Sm. & Lellinger) Salino & T.E. Almeida; * G. kuhlmannii (Brade) Brade; G. leonina (Maxon ex Caluff & C. Sánchez) Salino & T.E. Almeida; * G. leptocladia Fée; * G. levyi (E. Fourn.) Salino & T.E. Almeida; * G. liebmannii (Maxon & C.V. Morton) Salino & T.E. Almeida; * G. littoralis (Salino) Salino & T.E. Almeida; * G. lugubriformis (Rosenst.) Salino & T.E. Almeida; * G.lugubris (Mett.) Brade; * G. macropus (Blume) Ching (as “macropoda”); * G. macrotis (Hook.) Pic.Serm.; G. martinezii (A.R. Sm.) Salino & T.E. Almeida; G. megalodus (Schkuhr) C. Presl; * G. minor (C. Chr.) Salino & T.E. Almeida; G. minutissima (Caluff & C. Sánchez) Salino & T.E. Almeida; * G. mollis Fée; * G. monosora (C. Presl) Brade; G. montana (Salino) Salino & T.E. Almeida; * G. moranii C. Sánchez; * G. multigemmifera (Salino) Salino & T.E. Almeida; G. munchii (A.R. Sm.) Salino & T.E. Almeida; * G. nephrodioides (Klotzsch) Vareschi; * G. nicaraguensis (E. Fourn.) Salino & T.E. Almeida; * G. nigricans (Ekman & C. Chr.) Salino & T.E. Almeida; * G. obliterata (Sw.) C. Presl; * G. oroniensis (L.D. Gómez) Salino & T.E. Almeida; * G. paranaensis (Salino) Salino & T.E. Almeida; * G. paucijuga (Klotzsch) A.R. Sm.; * G. paucipinnata (Donn.Sm.) Salino & T.E. Almeida; G. pellita (Willd.) A.R. Sm.; * G. pennata (Poir.) Pic.Serm.; G. peripae (Sodiro) Salino & T.E. Almeida; G. pilonensis (A.R. Sm. & M. Kessler) Salino & T.E. Almeida; * G. pinnatifida (A.R. Sm.) Salino & T.E. Almeida; * G. platypes Fée; * G. poiteana (Bory) Ching * G. praetermissa (Maxon) Salino & T.E. Almeida; G. pyramidata Fée; G. redunca (A.R. Sm.) Salino & T.E. Almeida; Goniopteris refracta (Fisch. & C.A. Mey.) Brade; * G. reptans (J.F. Gmel.) C. Presl (Fig. 2F); * G. resiliens (Maxon) Salino & T.E. Almeida; * G. retroflexa (L.) Salino & T. E. Almeida; * G. rhachiflexuosa (Riba) Salino & T.E. Almeida; * G. riograndensis (Lindm.) Ching; * G. × rolandii (C. Chr.) A.R. Sm.; * G. sagittata (Sw.) Pic.Serm.; G.salinoi I.O. Moura & L.C. Moura; G. sapechoana (A.R. Sm. & M. Kessler) Salino & T.E. Almeida; * G. scabra (C. Presl) Brade; * G. schaffneri (Fée) Salino & T.E. Almeida; G. schippii (Weath.) Salino & T.E. Almeida; G. schomburgkii (A.R. Sm.) Salino & T.E. Almeida; G. schunkei (A.R. Sm.) Salino & T.E. Almeida; * G. schwackeana (C. Chr.) Brade; * G. sclerophylla (Poepp. ex Spreng.) Wherry; * G. scolopendrioides (L.) C. Presl; G. seidleri Salino; G. semihastata (Kunze) Salino & T.E. Almeida; * G. semirii (Salino & Melo) Salino & T.E. Almeida; G. septemjuga (C. Chr.) Salino & T.E. Almeida; * G. serrulata (Sw.) J. Sm.; G. skinneri (Hook.) Salino & T.E. Almeida; G. smithii Salino; G. stephanii (A.R. Sm.& M. Kessler) Salino & T.E. Almeida; * G. stolzeana (A.R. Sm.) Salino & T.E. Almeida; * G. straminea (Baker) Ching; G. strigosa Fée; * G. subdimorpha Salino; G. subsagittata (Maxon & C. Chr.) Salino & T.E. Almeida; G. tenebrica (Jenman) Salino & T.E. Almeida; G. tenera Fée; * G. tetragona (Sw.) C. Presl; * G. toganetra (A.R. Sm.) Á. LÖve & D. LÖve; * G. tristis (Kunze) Brade; * G. tryoniorum (A.R. Sm.) Salino & T.E. Almeida; * G. tuxtlensis (T. KrÖmer, Acebey & A.R.Sm.) Salino & T.E. Almeida; * G. urbanii (Sodiro) Salino & T.E. Almeida; G. venusta (Heward) Pic.Serm.; * G. verecunda (Proctor) Salino & T.E. Almeida; * G. vivipara (Raddi) Brack.; G.windischii Salino; * G. yaucoensis (Proctor) Salino & T.E. Almeida (Fig. 2B).
description: Published as part of Fawcett, Susan & Smith, Alan R., 2021, A Generic Classification of the Thelypteridaceae, Fort Worth, Texas, USA :BRIT Press on pages 43-45, DOI: 10.17348/jbrit.v15.i2.1206, http://zenodo.org/record/14111022