soral and of the Venation patterns, characteristics, shape fronds of the microsorioid Polypodiaceae By W.L.A. Hetterscheid and E. Hennipman With 14 figures in the text Abstract W. L. A. & E.: soral and Hetterscheid, Hennipman, Venationpatterns, characteristics, shape of the fronds of the microsorioidPolypodiaceae. — Bot. Jahrb. Syst. 105: 11—47. 1984. — ISSN 0006-8152. The deals with detailed of the venation, soral and distribu- present paper a study shape referred tion, and frondshape of the microsorioidPolypodiaceae, including taxa generally the to genera Christiopteris, Colysis, Dendroconche, Dendroglossa, Diblemma, Lecanopte- ris, Leptochilus, Microsorium, Neocheiropteris, Neolepisorus, Paraleptochilus, Phymatodes, and Podosorus. The fascinating diversity found in the characters studied is used to classi- 19 which main fy the taxa into groups, can be arranged into two groups, using ontoge- netic dataof the venation. The results that the be united: suggest following genera can Colysis, Dendroglossa, and Paraleptochilus, whereas Dendroconche and Diblemma should be merged with Mi- crosorium. The latter is will be genus remarkably heterogeneous; a systematic study conducted in the nearfuture. 1. Introduction The deals with the venation and present study gross morphology, pattern, and distribution of a number of selected from the sorus shape genera family Polypodiaceae. The includes in the first the Den- group place genera Colysis, droconche, Dendroglossa, Leptochilus, Microsorium (inch Kaulinia), Paraleptochi- and which lus, Podosorus, in our opinion may constitute a monophyletic group. This that each means they are thought to be mutually more related to other than of them is other the In the second this one to any genus in family. place deals with the of Neochei- paper group consisting Christiopteris, Lecanopteris, 0006-8152/84/0105-0011 $09.25 © 1984 E. Schweizerbart'sche Verlagsbuchhandlung,D-7000 Stuttgart 1 12 W. L. A. Hetterscheid & E. Hennipman,Polypodiaceae and which ropteris (incl. Neolepisorus), Phymatodes, are added, as present-day authors have a between these and those suggested possible relationship genera attributed to the first Most of the mentioned above referred group. genera are Crabbe al. Microsorioideae which by et (1975) to the subfamily Nayar also should include and related This excludes Neochei- Crypsinus genera. subfamily which referred to the because of the ropteris they Pleopeltoideae Nayar pres- ence of peltate paraphyses. PiCHl Sermolli subdivided the 14 of (1977) Polypodiaceae into groups, one these the of listed above with comprising group genera the exclusion of Neo- cheiropteris, Lecanopteris, and Myrmecopteris (syn.: Myrmecophila — Lecanopte- ris; see & Walker referred Jermy 1975). Pichi Sermolli Neocheiropteris to a of related group genera to Pleopeltis. Lecanopteris, Myrmecopteris, and Solano- taken this author based their pteris were together by as a separate group on the time that myrmecophyly at same stating Myrmecopteris and Lecanopteris related are mutually more than either of them is to Solanopteris. Sen & Hen- NIPMAN (1981) found that the first two taxa are characterized by the occur- and rence of cyclo- cocyclocytic stomata. The inclusion of the in this results from the genus Christiopteris group work of the senior author on the included in the species formerly genus Lepto- chilus COPELAND The results that the by (1928). suggest genus Christiopteris, described by COPELAND because of its acrostichoidy, frond shape, and diplo- desmic of fertile venation its parts, is polyphyletic. Comments by Copeland the relative of the of (1947) on phylogenetic position species this genus, point- ed already to its heterogeneity. See for details Hennipman & Hetterscheid (1984). The Podosorus genus is listed in this group mainly because of the remarks made the its resemblance Mi- by publishing author (HOLTTUM 1966), on to crosorium tenuilore (J. Smith) Copel. and/or Diblemma. Sen & Hennipman found in P. Holt- (1981) exclusively copolocytic stomata to occur angustatus hence in favor of its with microsorioids turn, an argument grouping along true indication in (the Sen & Hennipman, op. cit., p. 183, that only polocytic in this is their stomata occur species, an error; compare comments on p. 196). The genus Neocheiropteris is usually placed in the alliance of Pleopeltis which includes the Polypodiaceae with peltate paraphyses. Recent investiga- tions the of the revealed the existence of on paraphyses Polypodiaceae two ma- jor types of peltate paraphyses using morphological and ontogenetical criteria The of found Neochei- (pers. comm. R. P. Baayen). type peltate paraphyses in is also in the Para- ropteris present genera Belvisia, Lemmaphyllum, Lepisorus, and of gramma, Neolepisorus. Investigations on the spores these genera carried Hennipman & Sen showed that the first four share out by (in prep.) genera a which absent also unique spore type is in Neocheiropteris (see HENNIPMAN & The latter finds of Rods 1983). genus its position in the group microsorioids under because characteristics of its venation well of its study q£ pattern as as stomata (Sen & Hennipman 1981). W. L. A. Hetterscheid & E. Hennipman, Polypodiaceae 13 Within the selection of microsorioids as delineatedabove, a character analy- sis is made of the venation pattern, shape of sterile and fertile fronds, including shape and distribution of the sori. Special attention is paid to the suggestion forward Hennipman to unite the Microsorium, put by (1977, p. 38) genera Colysis, and Leptochilus (incl. Dendroglossa and Paraleptochilus) because of the of the soral characters which these profound instability on genera were original- based. we will discuss more detail as this ly Furthermore, Microsorium into ge- be nus appeared to surprisingly heterogeneous. This is of research the of the study part a project on phylogenetics family Polypodiaceae s.str. For details see FIennipman (1984). Thanks due the Directors of the British are to Museum (Natural History) (BM), the Paris herbarium (P), Royal Botanic Gardens, Kew (K), the Rijksher- barium and the Utrecht herbarium of which herbarium material (L), (U) was received on loan, as well as the authorities of the Botanical Gardens at Leiden and Utrecht for The of the material providing living specimens. greater part studied is at and The of who present L U. cooperation Mr. J. Gubbels, pre- number of and of pared a significant projection photographs Miss G. P. Ver- DUYN, who assisted in preparing the plates, is much appreciated. Thanks are also due to Miss W. SLOOT for typing the manuscript. 2. Material and methods 2.1. Material The enumeration of in the material studied is given Chapter 5. The taxonomy fol- lowed is that Copeland number of described since largely given by (1947); a new genera are included. 2.2. Methods 2.2.1. Projection photography of Projection photographs cleared material are prepared according to the method de- scribed Hennipman of in L. and U. by (1977). Duplicates photographs are present 2.2.2. Terminology 2.2.2.1. Terminology of venation incl. areoles The from of terminology used starts the idea that the venation pattern segments should be of whether the compared irrespective segments are primary or secondary seg- ments. This practical procedure is suggested by the fact that the venation pattern of lobes in is and pinnae the family Polypodiaceae essentially the same compared to that of the undivided fronds. The holds if lobes with the same true comparing and/or pinnae lobe if is absent apical or pinna. Therefore, an apical lobe or pinna (e.g. Drynaria spe- or the venation in lateral lobes be cies) ill-developed, pattern or pinnae can extrapolated to the absent lobe or pinna. The following terms and definitions are used: of of SI Vp: primary vein a segment (Fig. la). The rhachis is the primary vein (= the lamina; definitionsee Tryon 1960). 14 W. L. A. Hetterscheid & E. Hennipman, Polypodiaceae = Fig. 1. Terminology of veins and areoles. — of veins. V primary vein. a, Terminology p = = = Vs vein. V vein. V connective. — of secondary t tertiary tc b—d, Terminology areoles. A1 = primary areole. A = costal areole. A2, A A = main areoles. 1c 3, 4 Vs: of from secondary vein a segment (Fig. la); a vein directly branching a primary vein. is is from definition. If the segment petiolate, this petiole excluded the Vt; vein of vein from tertiary a segment (Fig. la); a branching directly a secondary from vein or another tertiary vein. “connective” those less veins that Vtc: (Fig. la); more or straight tertiary connect these the two adjacent secondary veins. Usually veins are more prominent compared to rest of the tertiary venation. The tertiary venation is often copiously branched and anastomosed, thus forming areoles (with or without included veins). We distinguish: — primary areoles (Al, Fig. lb): areoles bounded by two adjacent secondary veins and two adjacent connectives. They are usually rectangular in outline, rarely quadrangular; their axis usually running parallel to the primary vein. When three or more primary is areoles are present between adjacent secondary veins, the venation called “areole- layered”. The primary areole bordering the primary vein is called: — costal areole (Ale, Fig. lb): areoles usually narrower that the other primary ones with a less complex included venation. It is bounded by the primary vein, two adjacent secondary veins and their first connective. — bounded the main areoles (A2, Fig. 1c, d): usually large areoles by primary vein, two adjacent secondary veins and one (not their first) connective. Usually the bordering ve- nation is prominent, the areole outline dome-shaped and polygonal. It should be noted In that a main areole includes by definitionthe costal areole (Fig. 1c, d). species possess- main second third of areoles in broad ing areoles, a or row prominent may develop seg- of have chosen ments. As they are implicitly connected to the existence main areoles, we it noted to name them accordingly. However, must be that they are only bounded by and veins Areoles in the first coded A3 and secondary tertiary (Fig. 1c, d). row are may second be either regularly (Fig. Id) or irregularly shaped (Fig.