Doryopteris J.Sm., J. Bot. (Hooker)

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 : 115-116

Doryopteris J.Sm., J. Bot. (Hooker) 3: 404. 1841; emend. J.C.Yesilyurt, nom. cons.

Type species:— Doryopteris palmata (Willd.) J. Sm., J. Bot. [Hooker] 4: 163. 1841. = Pteris palmata Willd., Sp. Pl. , ed. 4 [Willdenow] 5:

357. 1810. Type:— VENEZUELA. Distrito Federal: Caracas, Bredemeyer s.n. ( B –image seen). Homotypic synonyms: — Cassebeera Kaulfuss (1824: 216) . Type species:— Casseebeera triphylla (Lam.) Kaulf. = Adiantum triphyllum Lam., Encycl. [ J. Lamarck

& al.] 1(1): 41. 1783. Bakeriopteris Kuntze (1891: 807), C.Chr., Index Filicum: 148. 1905. nom. superfl. for Cassebeera Kaulf. Heteropteris Fée, Crypt. Vasc. Bresil 1: 123. 1869. Type species:— Heteropteris doryopteris Fée. Tryonella Pic.Serm., Webbia 29: 14. 1974. nom. nov. for Heteropteris Fée.

Plants terrestrial or rupestral, to 30(–60) cm tall. Rhizomes erect to sub-erect or slightly decumbent; scales of rhizome and petiole bases clathrate, lanceolate to more or less ovate-lanceolate ( Fig. 3 A 1–A 3 ). Petiole black to brown or dark-brown; cross section terete, sub-terete, sulcate to ridged; indumentum of short hairs, fibrils (scale-like) or acicular trichomes; with one vascular bundle at base, usually U- or V-shaped. Fronds monomorphic to dimorphic. Laminar dissection diverse, varying from sagittate to pedate, pedate-pinnatifid to 3-pinnatifid (sometimes becoming almost pinnate at the base of the lamina) and palmate to palmate-digitate; glabrous or with trichomes, 1–3-celled, sometimes acicular and/or fibrils. Veins free, partially or totally anastomosing, without included veinlets; vein-ends usually free and may end in hydathodes. Sori predominantly marginal, only rarely submarginal, receptacle forming a continuous line (vascular commissure) ( Fig. 4 A 6) or sori separate on each vein end (no commissure/connection) ( Fig. 4 A 1 ); sporangia maybe very densely packed giving a false impression of a continuous vascular commissure. Pseudo-indusium comprises the reflexed lamina margin, which may or may not be continuous around the lamina; pseudo-indusium margin smooth to somewhat irregular ( Figs. 3 B 1 , B 2 , B 4 ). Paraphyses absent (except for D. kirkii and D. palmata which have very small, bicellular paraphyses). Sporangia stalks more or less same length as sporangium or shorter to medium-stalked; basal cell of sporangium sometimes conspicuous. Spores predominantly rugulose and cristate.

Distribution: —Central to South America, South Africa, Hawaiian Islands (one species widely distributed).

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

The concept includes Doryopteris triphylla , which Tryon (1942) included in Lytoneuron due to its free venation pattern, and Doryopteris lonchophora (Mettenius 1859: 4) Smith (1875: 289) , which was excluded by Tryon (1942) on the basis of the anatomy of the receptacle. It also includes the members from the sub-clade ( Fig. 2 ) that diverged from Doryopteris sensu stricto clade and comprised Doryopteris concolor , D. kirkii (Hooker 1861: 81) Alston (1956: 14) , as well as D. decora and its allies from Hawaiian Islands. With the exception of Doryopteris tryphylla , D. concolor and D. tryonii that have free venation, and D. lorentzii and all the rest of Hawaiian species (except D. angelica K.Wood & W.H.Wagner in Wagner et al. (1999: 147)) that have partially (rarely to) anastomosed veins, all the remaining taxa have anastomosed venation pattern ( Yesilyurt, 2005). An exception is D. lonchophora , which has a discontinuous soral line ( Figs. 4A 2 , A 3 , A7) and even has some parts of the sori naked (without any kind of indusium or reflexed margin).

The list of species presented here has been based on Tryon et al. (1990) concept for Doryopteris , which comprised 25 species. It excludes the taxa that have been referred to the sect. Lytoneuron (which is here being proposed as a new genus), except D. triphylla , and includes the recently described species. Full systematic account and other nomenclatural questions are being on their final revision and will be presented elsewhere, which will include the status of Doryopteris majestosa , as it has been flagged as putative nom. superfl. because one of the cited specimen on the examined material being an isotype of a published species.

List of species:

Doryopteris adornata J. C. Yesilyurt, Bot. J. Linn. Soc. 158: 73–77. 2008.

Doryopteris collina (Raddi) J. Sm., J. Bot. (Hooker) 4: 163. 1841.

Doryopteris concolor (Langsd. & Fisch.) Kuhn, Reisen Ost Afr. [Decken] 3 [3]. Bot.: 19. 1879. Doryopteris crenulans (Fée) H.Christ in Schwacke, Pl. Nov. Mineiras 2: 26. 1900.

Doryopteris decora Brack., U.S. Expl. Exped., Filic. 16: 103, t. 13, f. 1. 1854.

Doryopteris decipiens J. Sm., Hist. Fil. 289. 1875.

Doryopteris kirkii (Hook.) Alston, Bol. Soc. Brot. sér. 2, 30: 14. 1956.

Doryopteris leitei ( R.M.Tryon, Amer. Fern J. 36: 48. 1946) J. C. Yesilyurt, stat. nov. Doryopteris lonchophora (Mett.) J.Sm., Hist. Fil. 289. 1875.

Doryopteris lorentzii (Hieron.) Diels, Nat. Pfl. 1 [4]. 270. 1899.

Doryopteris nobilis (T.Moore) C.Chr., Ind. Fil. 244. 1905.

Doryopteris majestosa J.C.Yesilyurt, Amer. Fern J. 97(4): 212. 2008. nom. superfl.? Doryopteris palmata (Willd.) J.Sm., J. Bot. (Hooker) 4: 163. 1841.

Doryopteris pedata (L.) Fée, Mém. Foug., 5. Gen. Filic. 133. 1850–1852.

Doryopteris pentagona Pic. Serm., Webbia 60(1): 231. 2005.

Doryopteris rediviva Fée, Crypt. Vasc. Bresil 2: 30, t. 89, f. 1. 1872.

Doryopteris sagittifolia J.Sm., J. Bot. (Hooker) 4: 163. 1841.

Doryopteris stieri Rosenst., Hedwigia 46: 86. 1906.

Doryopteris surinamensis J.C.Yesilyurt, Brittonia 60(1): 63. 2008.

Doryopteris triphylla (Lam.) H.Christ, Bull. Herb. Boissier II , 2; 546. 1902.

Doryopteris varians (Raddi) J. Sm., J. Bot. (Hooker) 4: 163. 1841.

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).