Lytoneuron (Klotzsch) J.C.Yesilyurt

Yesilyurt, Jovita C., Barbará, Thelma, Schneider, Harald, Russell, Stephen, Culham, Alastair & Gibby, Mary, 2015, Identifying the generic limits of the Cheilanthoid genus Doryopteris, Phytotaxa 221 (2), pp. 101-122 : 116-117

Lytoneuron (Klotzsch) J.C.Yesilyurt , comb. & stat. nov.

Basionym:— Doryopteris sect. Lytoneuron

Type species: — Doryopteris lomariacea Klotzsch, Linnaea 20: 343. 1847 = Lytoneuron lomariaceum (Klotzsch) J.C.Yesilyurt , comb. nov. Type:— BRITISH GUIANA: Without locality, without date, Schomburgk 1197 (holotype BM!, isotype K!).

Synonyms: — Pellaea sect. Doryopteridastrum Fée ex Prantl (1882: 418) , nom. nud. Type species:— Pellaea quinquelobata Fée, Crypt. Vasc. Bresil 1: 42, t. 10, f. 1. 1869.

Plants terrestrial or rupestral, to 30(–50) cm tall. Rhizome decumbent to horizontal; sales of the rhizome and petiole bases clathrate to non-clathrate; linear to linear-lanceolate ( Fig. 3 A 4–A 5), with or without toothed margin ( Fig. 3 A 5). Petiole terete, sub-terete, rarely sulcate, sometimes with conspicuous roughened to verrucose texture; indumentum of short and/or lanose hairs, fibrils (scale-like), these sometimes with glandular tips or adpressed scales; with two vascular bundle at the base, which unite towards the base of the lamina. Fronds monomorphic to dimorphic. Lamina dissection diverse, varying from simple, entire to 1–3-lobed, pedate, to pedate-pinnatifid to 2-pinnatifid; glabrous or with minute trichomes or fibrils (scale-like). Veins predominantly free (except Lytoneuron ornithopus (Mettenius 1867: 166) J.C.Yesilyurt ) which has anastomosed venation; vein-ends usually free and forming hydathodes. Sori marginal to predominantly submarginal. Receptacle forms a discontinuous line (vascular commissure) ( Fig. 4 A 2–A 3 , A 7), usually forming a narrow or broad line ( Fig. 4 A4 ), giving an aspect of a true indusium. Pseudo-indusium margin usually irregular or crenulated ( Fig. 3 B 2–B 3 ). Paraphyses often present ( Fig. 4 B 1 ), usually tri-quadricellular. Sporangia stalks usually long-stalked (more or less 1 ½ longer than sporangium length); basal cell of sporangium absent or inconspicuous. Spores predominantly smooth to rugulate.

Distribution:— South America, with the majority of the species endemic to Brazil

Notes: — Klotzsch (1847) established section Lytoneuron including Doryopteris geraniifolia (Raddi 1819: 293) ( Klotzsch 1847: 343) (= D. concolor ) and species from Cassebeera . The concept was adopted and followed by Tryon (1942). The concept excludes Doryopteris triphylla , which Tryon (1942) included in Lytoneuron due to its free venation pattern alongside with D. concolor .

The main characters that distinguish the genus Lytoneuron from Doryopteris and Ormopteris are based on the number of vascular bundles at the base of the petiole, petiole shape and indumentum (including scales at the base), lamina dissection, venation type, soral condition and spores (type rather than size).

As stated above, full systematic account and nomenclatural questions including lectotypifications, are undergoing revision and will be presented elsewhere. Type information is limited to citation of type material as further type specimens remain to be investigated to determine their status.

List of species:

Lytoneuron acutilobum (Prantl) J.C.Yesilyurt , comb. nov.

Basionym:— Pellaea acutiloba Prantl in Bot. Jahrb. Syst. 3: 425. 1882.

Type material:— Glaziou 2471, 7262 ( Brazil, RJ).

Lytoneuron crenulans (Fée) J.C.Yesilyurt , comb. nov.

Basionym:— Pellaea crenulans Fée, Crypt. Vasc. Brésil 2: 27, t. 87, fig. 3. 1872.

Type material:— Glaziou 5345, 5343 ( Brazil, RJ)

Lytoneuron itatiaiense (Fée) J.C.Yesilyurt , comb. nov.

Basionym:— Pellaea itatiaiensis Fée, Crypt. Vasc. Brésil 2: 26, t. 88, fig. 1. 1872.

Type material:— Glaziou 5348 ( Brazil, RJ)

Lytoneuron feei (Brade) J.C.Yesilyurt , comb. nov.

Basionym:— Doryopteris feei Brade, Arch. Inst. Biol. Veg. Rio de Janeiro 1: 226, fig. 4 & t. 5. 1935.

Type material:— Toledo & Brade 6496; Kuhlman s.n. ( Brazil, RJ)

Lytoneuron lomariaceum (Kloszch) J.C.Yesilyurt , comb. nov.

Basionym:— Doryopteris lomariacea Klotzsch in Linnaea 20: 343. 1847.

Type material:— R. Schomburgk 1197 ( British Guiana)

Lytoneuron ornithopus (Hook. & Baker) J.C.Yesilyurt , comb. nov.

Basionym:— Pellaea ornithopus Mett. ex Hook. & Baker, Syn. Fil. 166.1867. ( Pellaea lomariacea (Klotzsch) Hook. var. digitatopalmata Hook., Sp. Fil. 2: 133. 1858)

Type material:— Gardner 5298 ( Brazil, MG)

Lytoneuron paradoxum (Fée) J.C.Yesilyurt comb. nov.

Basionym:— Cassebeera paradoxa Fée, Mém. Fam. Foug. 7: 30, t. 20, fig. 2. 1857, as Cassebeeria .

Type material:— Gardner 5930 ( Brazil, RJ)

Lytoneuron quinquelobatum (Fée) J.C.Yesilyurt comb. nov.

Basionym:— Pellaea quinquelobata Fée, Crypt. Vasc. Brésil 1: 42, t. 10, fig. 1. 1869.

Type material:— Glaziou 2055 ( Brazil, RJ)

Lytoneuron rosenstockii (Brade) J.C.Yesilyurt comb. nov.

Basionym:— Doryopteris rosenstockii Brade, Bol. Mus. Nac. Rio de Janeiro 7: 143, t. 8. 1931.

Type material:— Brade 9257 ( Brazil, RJ)

Lytoneuron rufum (Brade) J.C.Yesilyurt comb. nov.

Basionym:— Doryopteris rufa Brade, Bol. Mus. Nac. Rio de Janeiro 5: 94, t. 2, fig. 2. 1929.

Type material:— Bruno de Godoy 28 ( Brazil, MG)

Lytoneuron subsimplex (Fée) J.C.Yesilyurt comb. nov.

Basionym:— Pellaea subsimplex Fée, Cryp. Vasc. Bresil 1: 44, t. 4, fig. 3. 1869.

Type material:— Glaziou 3160 ( Brazil, RJ)

Lytoneuron tijucanum (Brade & Rosenst.) J.C.Yesilyurt , comb. nov.

Basionym:— Doryopteris tijucana Brade & Ros., Bol. Mus. Nac. Rio de Janeiro 7: 144, t. 9. 1931.

Type material:— Brade 8598 ( Brazil, RJ)

Figures

FIGURE 1. Sketch of the phylogeny of cheilanthoid ferns based on rbcL sequence data (as available in December 2013). The branch lengths correspond to the reconstructed number of changes in the maximum likelihood reconstruction whereas stars mark clades with a posterior confidence of p ≥ 0.95 in the consensus tree of the Bayesian inference of the phylogeny. Species of major groups outside of Doryopteris and relatives discussed are collapsed. Doryopteris is shown as defined in Tryon (1942) with the sect. Doryopteris and sect. Lytoneuron recognized and Doryopteris concolor and relatives assigned as Doryopteris concolor group. Choristosoria and Pteridella are widely ignored generic names that can be attributed to grade/ clades recognized in this study. All other generic names are assigned according to our current understanding. Generic names in “ “ may require redefinition. In some case, putative segregates are named but in general the oldest available generic name is selected. The relationships of D. pedatoides and D. pilosa are shown. The relationships of Doryopteris-Lytoneuron-Ormopteris relatives are outlined but shown in more detail in figure 2. CD = crown group of the Doryopteris- Lytoneuron-Ormopteris clade,SD =stem group Doryopteris-Lytoneuron-Ormopterisclade, TD= Trachypteris+Adiantopsis+Doryopteris- Lytoneuron-Ormopteris clade. D = Doryopteris clade, L = Lytoneuron clade, O = Ormopteris clade.

FIGURE 2. Phylogram obtained in the maximum likelihood analyses of the combined rbcL and trnLF dataset. Stars mark clades with a posterior confidence of p ≥ 0.95 in the consensus tree of the Bayesian inference of the phylogeny. The Adiantopsis clade is collapsed for (Link-Perez et al. 2011). All abbreviation and lineage assignments as in Fig. 1. If more than one specimen was investigated, the number is given (e.g., “2 smp”.A = regularly anastomosed venation, F = free or irregularly anastomosed venation. F-A marked transitions from free to regularly anastomosing venation,A-F marked the transition from anastomosing to free venation. Note: The species Doryopteris palmata

FIGURE 3. A. Types of the margin and cells of the rhizome scales: A1–A3. Lanceolate scales; A4–A5. Linear scales. B. Types of margin and indusium cells. Scale bars= 100μm. (based on:A1: J. C. Yesilyurt, 732 & J. Prado (BM); A2 & B4: J.C. Yesilyurt 519 & R.N. Cislinski, R.E. Cislinski (BM); A3: J.C. Yesilyurt, 547 & J. Prado, P.H. Labiak (BM); A4: Herbarium specimen-GUA 01071; A5: J. C. Yesilyurt, 738 & J. Prado (BM); B1: J.C. Yesilyurt 726 & J. Prado (BM); B2: Herbarium specimen L0066049; B3: J.C. Yesilyurt 739 & J. Prado (BM). (Drawn by J.C. Yesilyurt).

FIGURE 4. A. Types of receptacle and indusium (sketches only which were not based on any particular specimen); B. Types of sporangia and paraphyses. Scale bar= 100μm. (based on B1: Herbarium specimen US 2626408; B2: J.C. Yesilyurt 529 & R.N. Cislinski, R. E. Cislinski (BM); B3: J.C. Yesilyurt 563 & J. Prado, P.H. Labiak (SP). (Drawn by J. C. Yesilyurt).