New combinations in Lastreopsis and Parapolystichum - Springer Link

New combinations in Lastreopsis and Parapolystichum (Dryopteridaceae) PAULO H. LABIAK1, MICHAEL SUNDUE2, GERMINAL ROUHAN3,

AND

ROBBIN C. MORAN4

1

Dept. de Botânica, Universidade Federal do Paraná, C.P. 1903181531-980 Curitiba, PR, Brazil; e-mail: [email protected] 2 The Pringle Herbarium, Dept. of Plant Biology, University of Vermont, 27 Colchester Ave., Burlington, 05405 VT, USA; e-mail: [email protected] 3 Muséum National d’Histoire Naturelle, UMR CNRS 7205, Herbier National, CP39, 16 rue Buffon, F-75231 Paris cedex 05, France; e-mail: [email protected] 4 The New York Botanical Garden, Bronx, NY 10456-5126 USA; e-mail: [email protected]

Abstract. Recent molecular phylogenetic studies have shown that Lastreopsis is paraphyletic. Twenty-seven of its species belong to a separate clade for which the previously published generic name Parapolystichum is available. Parapolystichum is widely distributed, with four species in the Neotropics, six in Africa, nine in Madagascar, and eight in Australia and New Zealand. We make the following 25 new combinations in Parapolystichum: P. acuminatum, P. acutum, P. barterianum, P. boivinii, P. confine, P. coriaceosquamatum, P. currorii subsp. currorii, P. currorii subsp. eglandulosum, P. excultum, P. fideleae, P. glabellum, P. hornei, P. manongarivense, P. microsorum ssp. microsorum, P. microsorum ssp. pentangulare, P. munitum, P. nigritianum, P. perrierianum, P. pseudoperrierianum, P. rufescens, P. smithianum, P. subsimile, P. tinarooense, P. vogelii, and P. windsorense. Also, we designate a lectotype for Dryopteris exculta var. squamifera (=Parapolystichum excultum). No known morphological characters distinguish Parapolystichum from Lastreopsis s.s. Certain species of Parapolystichum, however, bear buds distally along the rachis, whereas Lastreopsis lacks such buds. Because no morphological character, or suite of characters, consistently separates the two genera, nine species of Lastreopsis s.l., which were not included in the previous phylogenetic analyses, could not be classified in either Lastreopsis or Parapolystichum. New combinations are also made for L. dissecta, L. tripinnata and L. poecilophlebia, species previously classified in Oenotrichia and Coveniella. Because Coveniella was considered monotypic, it is here reduced to a synonym of Lastreopsis. As re-circumscribed here, Lastreopsis now consists of about 16 species, all restricted to southeastern Asia and Oceania. Key Words: Ferns, taxonomy, phylogeny, systematics, Oenotrichia, Coveniella.

Lastreopsis Ching, a m ember of the Dryopteridaceae sensu Smith et al. (2006, 2008), is a group of medium-sized terrestrial ferns usually with 2–4-pinnate laminae. The genus was monographed by Tindale (1965), who recognized 33 species, most of which are tropical but with some extending into temperate regions of the Southern Hemisphere. Since Tindale’s monograph, other species of Lastreopsis have been described (e.g., Rakotondrainibe & Tronchet, 2009; Perrie & Brownsey, 2012). The total number of species, subspecies, and varieties now

recognized in the genus stands at about 55. The main character that distinguishes the genus from others in the family is the architecture of the adaxial surface of the axes. The margin of the green lamina is thickened and decurrent onto the axis of the next lowest order (Holttum, 1959; Tindale, 1965). The thickened margins or ridges on either side of the rachises or costae define a flat (or nearly so) surface, usually pubescent by 0.1– 0.5 mm long multicellular hairs (Labiak et al., 2014, Fig. 7 D–F), which Tindale referred to as “Ctenitis-hairs”. Among Dryopteridaceae, this

Brittonia 67(1): 79–86 (2015), DOI 10.1007/s12228-014-9351-3 ISSN: 0007-196X (print) ISSN: 1938-436X (electronic) © 2014, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A. Published Online: 1 October 2014

80

BRITTONIA

rachis-costa architecture is otherwise found only in Rumohra Raddi, and some Dryopteris Adans. (McKeown et al., 2012 ; Sundue et al., 2013; Labiak et al., 2014). Megalastrum Holttum differs in this respect by having non-sulcate rachises (or hardly so), and the lamina decurrent onto the rachis as a narrow wing that is not thickened (Labiak et al., 2014). Indusia can also be useful to distinguish genera. Phylogenetic studies have recovered Lastreopsis sensu Tindale (1965) as paraphyletic, with Rumohra and Megalastrum nested within (Schuettpelz & Pryer, 2007; Labiak et al., 2014).We call the clade containing these genera the “lastreopsid ferns.” Besides rachis-costa architecture, morphological characters that define this clade include large and highly dissected fronds, asymmetrical basal pinnae that are enlarged basiscopically (except Rumohra), creeping rhizomes (except Megalastrum and some species of Parapolystichum, which have decumbent to erect rhizomes), round discrete sori, and yellowish or orange glands on the laminar tissue, and paraphyses on the sporangial stalks (present in most of the clades). Labiak et al. (2014) found that Lastreopsis sensu Tindale (1965) is paraphyletic, being composed of two main clades. The first clade, which includes the type of Lastreopsis [L. recedens (J. Sm. ex T. Moore) Ching = L. tenera (R. Br.) Tindale] was recovered as sister to Rumohra and Megalastrum, with high support values (PP=1.0, ML-BS=100%, and MP-BS=100%) (Labiak et al., 2014). This clade can be further subdivided in two lineages, which were referred by Labiak et al. (2014) as the “Lastreopsis amplissima” and “Lastreopsis s.s.” clades. The first occurs in the Neotropics, Australia, New Zealand, and several islands in the southwest Pacific. The species that belong to it are L. amplissima (C. Presl) Tindale, L. davallioides (Brack.) Tindale, L. hispida (Sw.) Tindale, and L. killipii (C. Chr. & Maxon) Tindale. The relationships of this clade were not clearly established: while the bayesian and the maximum likelihood recovered it as sister to Lastreopsis s.s. (PP=0.91 and ML-BS=66%), the maximum parsimony analysis suggested it to be more closely related to Megalastrum+Rumohra (MP-BS=73%) (Labiak et al., 2014). The second clade of Lastreopsis s.l. is Lastreopsis s.s. It is restricted to Asia and Oceania, being completely absent from the Neotropics. Based on the sampling used by Labiak et al. (2014), the species that belong to it are

[VOL 67

L. decomposita (R. Br.) Tindale, L. marginans (F. Muell.) D. A. Sm. & Tindale, L. poecilophlebia (Hook.) Labiak, Sundue & R. C. Moran (comb. nov., see below), L. silvestris D. A. Sm. ex Tindale, L. subsericea (Mett.) Tindale, L. tenera (R. Br.) Tindale, L. tripinnata (F. Muell. ex Benth.) Labiak, Sundue & R. C. Moran (comb. nov., see below), L. walleri Tindale, and L. wurunuran (Domin) Tindale. Another species, L. subrescedensChing, was not included in the Labiak et al. (2014) phylogeny, but recent analysis performed by us have shown that is it also nested within this clade. The second main clade of Lastreopsis s.l. recovered by Labiak et al. (2014) was sister to the rest of the lastreopsids; that is, to Lastreopsis s.s. L. amplissima clade, and Rumohra + Megalastrum. The support values for this relationship were relatively high in most analyses (PP=0.96, MLBS=76%, and MP-BS=100%) (Labiak et al. (2014). This clade is pantropical, occurring in the Neotropics, Africa, Madagascar, Asia, Australia, and New Zealand. It contains the following species according to Labiak et al. (2014): L. acuminata C.V. Morton, L. acuta (Hook.) Tindale, L. barteriana (Hook.) Tardieu, L. boivinii (Baker) Tardieu, L. currorii (Mett.) Tindale, L. effusa (Sw.) Tindale, L. effusa ssp. confinis (Maxon ex C. Chr.) Tindale, L. exculta (Mett.) Tindale, L. glabella (A. Cunn. in Hook.) Tindale, L. microsora (Endl.) Tindale, L. munita (Mett.) Tindale, L. nigritiana (Baker) Tindale, L. perrieriana (C. Chr.) J.P. Roux, L. pseudoperrieriana (Tardieu) Tardieu, L. rufescens (Blume) Ching, L. smithiana Tindale, L. subsimilis (Hook.) Tindale, L. tinarooensis Tindale, L. vogelii (Hook.) Tindale, L. windsorensis D.L. Jone & B. Gray. One of these species (L. effusa (Sw.) Tindale), is the type of Parapolystichum (Keyserl.) Ching (Parapolystichum effusum (Sw.) Ching). Thus, a generic name is available for this second clade, and we adopt it here. M o r p h o l o g i c a l l y, L a s t r e o p s i s a n d Parapolystichum are indistinguishable. Compared to other genera of Dryopteridaceae, their most significant similarities are the rachis-costa architecture as mentioned above, a character they share with Rumohra. Also, both Lastreopsis and Parapolystichum bear shiny, brightly colored (usually yellow or orange), cylindrical-oblong glands on the sporangial stalks and often on the laminae, especially on the abaxial surface of the veins (Tindale, 1965). Lastreopsis generally has shortcreeping or ascending rhizomes, whereas Parapolystichum has more massive decumbent

2015]

LABIAK ET AL: PARAPOLYSTICHUM (DRYOPTERIDACEAE)

ones with the leaves set close together, but the distinction is not absolute (pers. obs.). Noteworthy is that several species of Parapolystichum bear buds distally along the rachis, a character that is absent in Lastreopsis s.s. (see optimization of this character in Fig. 5 of Labiak et al., 2014). Chromosome number in both Lastreopsis and Parapolystichum is n=41 (Tindale, 1965; Tindale & Roy, 2002). This is the base number for the Dryopteridaceae sensu Smith et al. (2006). Counts for Parapolystichum come from the Neotropics, Africa, Australia, and New Zealand. In the Neotropics, only one species, P. effusum (reported as Ctenitis effusa (Sw.) Copel.), has been reported from two separate counts. A Jamaican plant was found to be as n=41 (Walker, 1966), and a plant from Trinidad, n=82 (Walker, 1973, 1985). In Africa only L. currori (as Ctentis efulensis (Baker) Tardieu) from Ghana has been counted (see Appendix of Alston, 1959). In New Zealand, chromosome counts are known for P. glabellum (n=82) (Brownlie, 1958), and P. microsorum subsp. pentangulare (n=82) (Brownlie, 1961). As summarized by Tindale and Roy (2002), the following Australian species of Parapolystichum have been counted: P. decompositum, P. microsorum, P. rufescens, P. tinarooense, and P. acuminatum. All were based on x=41. Although Lastreopsis and Parapolystichum cannot be distinguished morphologically or cytologically, we prefer to recognize them as distinct genera. We find this more convenient than the alternative, which is lumping them with Megalastrum and Rumohra into a single genus. Such a combined “super genus” would still have the drawback of being indistinguishable morphologically. Also, lumping would require about 140 new combinations in Rumohra (the earliest genus name available), instead of the 25 made here for Parapolystichum, and the resulting genus would itself lack morphological synapomorphies (Labiak et al., 2014). Recognizing four separate genera, as favored here, accords with recent pteridologists who have accepted Megalastrum and Rumohra as distinct (e.g., Tryon & Stolze, 1991, Smith & Moran, 1995, 2008; Sundue et al., 2013). Below, we first make new combinations for species belonging to Parapolystichum, and then for three species of Lastreopsis. For completeness and convenience, we also list the three species previously combined in Parapolystichum (P. davallioides, P. effusum and P. killipii). For each species cited below, we give only the basionym

81

and the name in Lastreopsis; more complete synonymy can be found in Tindale (1965). Geographical distribution by country is based primarily on Tindale (1965), Andrews (1990), Moran (1995), Jones (1998), Roux (2009), and Rakotondrainibe and Tronchet (2009). We assign species to Parapolystichum based on the phylogenetic results of Labiak et al. (2014). Five African and Western Indian Ocean species, however, were assigned to Parapolystichum, not Lastreopsis, even though they were not included in phylogenetic analysis of Labiak et al. (2014). The species are the African P. currorii subsp. eglandulosa, the Western Indian Ocean P. c o r i a c e o s q u a m a t u m , P. f i d e l e a e , P. manongarivense, and P. hornei. They are marked in the below list with an asterisk (*).We make these combinations, without the benefit of molecular data for these species, for two reasons. First, the African species bear buds along the rachises—a character not found in Lastreopsis s.s. Second, of the species included in molecular studies so far, only those of Parapolystichum, not Lastreopsis, occur in Africa and Madagascar. Even with the assignment of these African and Western Indian Ocean species to Parapolystichum, there remained several species of Lastreopsis s.l. not included in the phylogenetic analysis of Labiak et al. (2014) that could still not be placed in either Lastreopsis s.s. or Parapolystichum. Those species are listed below under Incertae Sedis. The phylogenetic position of the monotypic Trichoneuron Ching is still unknown. It is based on T. microlepioides Ching which has been treated by the Flora of China Editorial Committee as Lastreopsis microlepioides (Ching) W. M. Chu & Z. R. He. Based on its morphology and geographical distribution, we also believe it will be nested within Lastreopsis or Parapolystichum, but this relationship needs to be confirmed by molecular phylogenetic studies. Parapolystichum (Keyserl.) Ching, Sunyatsenia 5: 239. 1940. Polystichum sect. Parapolystichum Keyserl., Polyp. Herb. Bunge 11: 4 5. 1 8 73 . D r y op t e r i s s u b ge n . Parapolystichum (Keyserl.) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd. 8, 6: 93. 1920 [“1919”]. Type: Parapolystichum effusum (Sw.) Ching. Parapolystichum acuminatum (Houlston) Labiak, Sundue & R. C. Moran, comb. nov.

82

BRITTONIA

Lastrea acuminata Houlston, Gard. Mag. Bot. 3: 317. 1851. Lastreopsis acuminata (Houlston) C. V. Morton, Contr. U.S. Natl. Herb. 30: 245. 1973. Distribution.—Australia. Parapolystichum acutum (Kuntze) Labiak, Sundue & R. C. Moran, comb. nov. Dryopteris acuta Kuntze, Revis. Gen. Pl. 2: 812. 1891. Nephrodium acutum Hook., Sp. Fil. 4: 147. 1862, non Presl (1825). Lastreopsis acuta (Hook.) Tindale, Contr. New South Wales Natl. 3: 245. 1963. comb. illeg. Distribution.—Southeast Brazil. Parapolystichum barterianum (Hook.) Rouhan, comb. nov. Polypodium barterianum Hook., Sp. Fil. 313. 1867. Lastreopsis barteriana (Hook.) Tardieu, Notul. Syst. (Paris) 14: 343. 1952. Distribution.—Cameroon, Nigeria, Equatorial Guinea, Madagascar. Parapolystichum boivinii (Baker) Rouhan, comb. nov. Nephrodium boivinii Baker, Syn. Fil. 501. 1874. Lastreopsis boivinii (Baker) Tardieu, Fl. Madagasc. 5: 347. 1958. Distribution.—Madagascar. Parapolystichum confine (Maxon ex C. Chr.) Labiak, Sundue & R. C. Moran, comb. et stat. nov. Dryopteris effusa var. confinis Maxon ex C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd. 8, 6: 101. 1920 [1919]. Lastreopsis effusa ssp. confinis (Maxon ex C. Chr.) Tindale. Contr. New South Wales Natl. Herb. 3: 299. 1965. Distribution.—Cuba, Haiti, Dominican Republic. Parapolystichum confine has long been considered a variety of P. effusum (e.g., by Christensen, 1920; Tindale, 1965), but it is quite distinct from that species. It was thought to be a variety of P. effusum because, like that species, indusia are absent. Yet in leaf size and dissection, P. confine more resembles the non-indusiate P. excultum (Mett.) Tindale. The rhizome scales of P. confine differ greatly from both P. effusum and P. excultum. Its scales are 2–4 mm wide, ovate-lanceolate, dark brown, firm and opaque, whereas those of the other two species are 1–2 mm wide, linear-lanceolate, light brown or pale yellowish brown, thin and translucent. The lamina texture of P. confine also differs,

[VOL 67

being thicker and shinier. Because of its distinctness from both species, we recognize P. confine at the rank of species instead of a variety of P. effusum. *Parapolystichum coriaceosquamatum (Rakotondr.) Rouhan, comb. nov. Lastreopsis coriaceosquamata Rakotondr., Adansonia, sér. 3, 31: 12. 2012. Distribution.—Madagascar. Parapolystichum currorii (Mett. ex Kuhn) Rouhan, comb. nov. Aspidium currori Mett ex Kuhn, Fil. Afr. 130. 1868. Lastreopsis currorii (Mett. ex Kuhn) Tindale, Vict. Nat. 73: 184. 1957. Distribution.— Angola, Burundi, Ivory Coast, Liberia, Guinea, Ghana, Nigeria, Cameroon, Equatorial Guinea, Central African Republic, Gabon, Congo, Uganda, Tanzania. *Parapolystichum currorii (Mett. ex Kuhn) Rouhan subsp. eglandulosum (Tindale) R. C. Moran & Rouhan, comb. nov. Lastreopsis currorii Mett. ex Kuhn subsp. eglandulosa Tindale, Contr. New South Wales Natl. Herb. 3: 284. 1965. Distribution.—Equatorial Guinea, Congo. Parapolystichum effusum (Sw.) Ching, Sunyatsenia 5: 239. 1940. Polypodium effusum Sw., Prodr. Fl. Ind. Occ. 134. 1788. Lastreopsis effusa (Sw.) Tindale, Vict. Nat. 73: 184. 1957. Polypodium divergens Willd. ex Schkuhr, Vier Zwan. Kl. Linn. Pfl. Krypto. Gew. 1: 27, tab. 26b. 1805. Lastreopsis effusa subsp. divergens (Willd. ex Schkuhr) Tindale, Contr. New South Wales Natl. Herb. 3: 299, t. 21. 1965. Parapolystichum effusum var. divergens (Willd. ex Schkuhr) Abbiatti, Rev. Mus. La Plata, secc. Bot. 9: 19. 1958. Polypodium dilatatum Liebm., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd., ser. 5, 1: 208. 1845. Lastropsis effusa subsp. dilatata (Liebm.) Tindale, Contr. New South Wales Natl. Herb. 3: 299. 1965. Distribution.—Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Cuba, Jamaica, Martinique, St. Vincent, Grenada, Puerto Rico, Dominican Republic, Haiti, Trinidad, Surinam, Guyana, Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil, Paraguay, Argentina. Tindale (1965) recognized three subspecies of Parapolystichum effusum (as Lastreopsis). One is here treated at the specific level as P. confine (see

2015]


above). Another, L. effusum subsp. divergens (Willd. ex Schkuhr) Tindale already has a combination in Parapolystichum (see above), but we consider it synonymous with P. effusum var. effusum. We also consider the third subspecies, L. effusa subsp. dilatata (Liebm.) Tindale, to be indistinguishable from the type subspecies; accordingly, it is placed here in synonymy. We cannot consistently distinguish the last two subspecies from herbarium material. Their distinction is based primarily on lamina division, but many intermediates occur as pointed out by Christensen (1920). The situation is complicated by polyploidy. Walker (1966, 1973, 1985) found diploids in Jamaica and tetraploids in Trinidad. More work is needed on the biosystematics of this species. Parapolystichum excultum (Mett.) Labiak, Sundue & R. C. Moran, comb. nov. Aspidium excultum Mett., Abh. Senckenberg. Naturf. Ges. 2: 69. 1858. Lastreopsis exculta (Mett.) Tindale. Lastreopsis exculta var. squamifera (C. Chr.) Tindale, Contr. New South Wales Natl. Herb. 3: 245. 1963. Dryopteris exculta (Mett.) C. Chr. var. squamifera C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Math. Afd. 8, 6: 96. 1920 [“1919”]. Type: Costa Rica. Cartago: Valle del Río Navarro, 1400 m, s.d., C. Wercklé 16741 (lectotype, here designated: P 00642630!). Christensen (1920) recognized var. squamifera based on more densely scaly rachises and costae. The variety was accepted by Tindale (1965) and Moran (1995), neither of whom saw any of the syntypes (i.e., Wercklé 16741, 16753, 16764). The lectotype designated here is the only syntype of var. squamifera found by us at P. Its rachises and costae appear just as scaly as those of typical P. exculta, and in all other respects the plant appears normal for that species. Accordingly, var. squamifera is here placed in synonymy. Distribution.—Mexico, Guatemala, Nicaragua, Costa Rica, Panama, Venezuela, Colombia, Ecuador. *Parapolystichum fideleae (Rakotondr.) Rouhan, comb. nov. Lastreopsis fideleae [as “fidelei”] Rakontondr., Adansonia, sér. 3, 31: 15. 2009. Distribution.—Madagascar.

83

Parapolystichum glabellum (A. Cunn.) Labiak, Sundue & R. C. Moran, comb. nov. Nephrodium glabellum A. Cunn. in Hooker, Companion Bot. Mag. 2: 367. 1837 [“1836”, publ. 1837]. Lastreopsis glabella (A. Cunn.) Tindale, Vict. Nat. 73: 183. 1957. Distribution.—New Zealand. *Parapolystichum hornei (Baker) Rouhan, comb. nov. Nephrodium hornei Baker in Hooker & Baker, Syn. Fil., ed. 2. 500. 1874. Lastreopsis hornei (Baker) Tindale, Vict. Nat. 73: 184. 1957. Distribution.—Seychelles. *Parapolystichum manongarivense (Rakotondr.) Rouhan, comb. nov. Lastreopsis manongarivensis Rakotondr., Adansonia, sér. 3, 31: 15. 2012. Distribution.—Madagascar. Parapolystichum microsorum (Endl.) Labiak, Sundue & R. C. Moran, comb. nov. Nephrodium microsorum Endl., Prodr. Fl. Norfolk. 9. 1833. Lastreopsis microsora (Endl.) Tindale, Vict. Nat. 73: 181. 1957. Distribution.—Australia, New Zealand. Parapolystichum microsorum (Endl.) Labiak, Sundue & R. C. Moran subsp. pentagulare (Colenso) Labiak, Sundue & R. C. Moran, comb. nov. Nephrodium pentangularum Colenso, Tasm. Journ. Nat. Sci. 2: 169. 1846. Ctenitis pentangularis (Colenso) Alston, Amer. Fern J. 45: 160. 1955. Lastreopsis microsora subsp. pentangularis (Colenso) Tindale, Contr. New South Wales Nat. Herb. 3: 334. 1965. Distribution.—New Zealand. Parapolystichum munitum (Mett.) Labiak, Sundue & R. C. Moran, comb. nov. Phegopteris munita Mett., Abh. Senckenberg. Naturf. Ges. 2: 298. 1858. Lastreopsis munita (Mett.) Tindale, Vict. Nat. 73: 183. 1957. Distribution.—Australia. Parapolystichum nigritianum (Baker) Rouhan, comb. nov. Polypodium nigritianum Baker, Syn. Fil. 313. 1867. Lastreopsis nigritiana (Baker) Tindale, Contr. New South Wales Nat. Herb. 3: 245. 1963. Distribution.—Liberia, Nigeria, Cameroon, Equatorial Guinea.

84

BRITTONIA

Parapolystichum perrierianum (C. Chr.) Rouhan, comb. nov. Dryopteris perrieriana C. Chr., Notes Pteridol. 16: 176, fig. 4a. 1925. Lastreopsis perrieriana (C. Chr.) J. P. Roux, Bothalia 34: 108. 2004. Distribution.—Madagascar. Parapolystichum pseudoperrierianum (Tardieu) Rouhan, comb. nov. Ctenitis pseudoperrieriana Tardieu, Mém. Inst. Sci. Madagascar, Sér. B, Biol. Vég. 6: 240. 1955. Lastreopsis pseudoperrieriana (Tardieu) Tardieu, Adansonia 5: 497. 1965. Distribution.—Madagascar. Parapolystichum rufescens (Blume) Labiak, Sundue & R. C. Moran, comb. nov. Aspidium rufescens Blume, Enum. Pl. Javae. 168. 1828. Polypodium rufescens (Blume) Blume, Flora Javae 194. 1829. Phegopteris rufescens (Blume) Mett., Ann. Mus. Bot. LugdunoBatavi 1: 223. 1864. Dryopteris rufescens (Blume) C. Chr., Index Filicum 290. 1905. Lastreopsis rufescens (Blume) Ching, Bull. Fan Mem. Inst. Biol. 8: 160. 1938. Distribution.—Sri Lanka, Indonesia, Australia, New Caledonia. Parapolystichum smithianum (Tindale) Labiak, Sundue & R. C. Moran, comb. nov. Lastreopsis smithiana Tindale, Contr. New South Wales Nat. Herb. 3: 90. 1961. Distribution.—Australia. Parapolystichum subsimile (Hook.) Rouhan, comb. nov. Gymnogramma subsimilis Hook., Sp. Fil. 4: 142. t. 293. 1864. Phegopteris subsimilis (Hook.) Mett., Filices Africanae 123. 1868. Lastreopsis subsimilis (Hook.) Tindale, Contr. New South Wales Nat. Herb. 3: 245. 1961. Distribution.––Ivory Coast, Gabon, Equatorial Guinea. Parapolystichum tinarooense (Tindale) Labiak, Sundue & R. C. Moran, comb. nov. Lastreopsis tinarooensis Tindale, Gard. Bull. Singapore 39: 170. 1987 [1986]. Distribution.––Australia. Parapolystichum vogelii (Hook.) Rouhan, comb. nov. Polypodium vogelii Hook., Sp. Fil. 4: 271. 1862. Lastreopsis vogelii (Hook.) Tindale, Contr. New South Wales Nat. Herb. 3: 45. 1963.

[VOL 67

Distribution.––Nigeria, Equatorial Guinea, Congo, Cameroon, Uganda. Parapolystichum windsorensis (D. L. Jones & B. Gray) Labiak, Sundue & R. C. Moran, comb. nov. Lastreopsis windsorensis D. L. Jones & B. Gray, Fl. Australia 48: 711. 1998. Distribution.—Australia. The following two species nest within Lastreopsis s.s. (Labiak et al., 2014). Accordingly, new combinations are here made for them in that genus: Lastreopsis poecilophlebia (Hook.) Labiak, Sundue & R. C. Moran, comb. nov. Polypodium poecilophlebia Hook., Sp. Fil. 5: 14. 1863. Dryopteris poecilophlebia (Hook.) C. Chr., Index Filicum 285. 1905. Coveniella poecilophlebia (Hook.) Tindale, Gard. Bull., Striats Settlem. 39: 169. 1986. Distribution.—Australia The genus Coveniella was established by Tindale (1986) on the basis of its unique morphological characters of 1-pinnate lamina with conform apex, and anastomosing veins. According to Labiak et al. (2014), Coveniella is nested within Lastreopsis s.s., and its recognition as a genus would render the later paraphyletic. Because Coveniella is monotypic (Jones, 1998), this combination reduces the genus to a synonym of Lastreopsis. Lastreopsis dissecta (C. T. White & Goy) Labiak, Sundue & R. C. Moran, comb. nov. Leptolepia dissecta C. T. White & Goy, Vict. Nat. 54: 149. 1938. Oenotrichia dissecta (C. T. White & Goy) S. B. Andrews, Austrobaileya 1: 12. 1977. Distribution.—Australia. This species is very similar to O. tripinnata, another Australian endemic (Brownsey, 1998). Although it was not included in the phylogenetic analysis of Labiak et al. (2014), its morphology suggests it also belongs to Lastreopsis. Lastreopsis tripinnata (F. Muell. ex Benth.) Labiak, Sundue & R. C. Moran, comb. nov. Davallia tripinnata F. Muell. ex Benth., Fl. Austr. 7: 717. 1878. Oenotrichia tripinnata (F. Muell. ex Benth.) Copel., Univ. California Publ. Bot. 16: 82. 1929. Distribution.—Australia.

2015]


Tryon & Tryon (1982) noted that this species does not belong in Oenotrichia, citing the presence of rhizome scales and monolete spores (vs. hairs and trilete spores in Oenotrichia). A similar conclusion was presented by Brownsey (1998), who further suggested that this species would belong to Lastreopsis or Ctenitis. In the analysis of Labiak et al. (2014), O. tripinnata was nested within Lastreopsis. Accordingly, we make the new combination of this species in Lastreopsis. Three other species of Oenotrichia are O. macgillivrayi (E. Fourn.) Brownlie, O. maxima (E. Fourn.) Copel., and O. tenera (Christ) Tagawa. Although their morphology suggests that they belong to the Dennstaedtiaceae (pers. observ.), their phylogenetic relationships remain to be tested. Oenotrichia tenera was treated in Microlepia, as M. tenera Christ, by Yeuhong et al. (2013). Incertae Sedis As mentioned above, no known morphological characteristics distinguish Lastreopsis and Parapolystichum. For this reason, nine species of Lastreopsis sensu Tindale (1965) that were not included in the molecular phylogenetic analysis of Labiak et al. (2014) cannot be placed in either Lastreopsis s.s. or Parapolystichum. The species are L. calantha (Endl.) Tindale, L. grayi D. L. Jones, L . k er m a de ce ns i s P err i e & B ro wn s e y, L. microlepioides W. M. Chu & Z. R. He, L. nephrodioides (Baker) Tindale, L. novoguineensis Holttum, L. pacifica Tindale, L. subsparsa (v.A.v.R.) Tindale, L. velutina (A. Rich.) Tindale, and L. vieillardii (Mett.) Tindale. For the time being, these species must be considered incertae sedis. Taxa excluded from Parapolystichum Parapolystichum davallioides (Brack.) Ching, Sunyatsenia 5: 239. 1940. Ctenitis davallioides (C. Presl.) Copel., J. Arn. Arbor. 30: 437. 1949. Lastrea davallioides Brack., U.S. Expl. Exped. 16: 202. 1854. Dryopteris davallioides (Brack.) O. Kuntze, Rev. Gen. Pl. 2: 812. 1891. Lastreopsis davallioides (Brack.) Tindale, Victoria Naturalist 73: 184. 1957. Distribution.—Tahiti, Samoa, Fiji, Vanuatu; 600–900 m. According to our molecular phylogenetic results (Labiak et al., 2014), this species should be classified in Lastreopsis, as has been done recently by most workers (e.g., Tindale, 1965).

85

Parapolystichum killipii (C. Chr. & Maxon) Ching, Sunyatsenia 5: 239. 1940. Dryopteris killipii C. Chr. & Maxon, Amer. Fern J. 18: 4. 1928. Lastreopsis killipii (C. Chr. & Maxon) Tindale. Distribution.—Costa Rica, Panama, Venezuela, Colombia, Ecuador, Peru, Bolivia. According to our molecular phylogenetic results (Labiak et al., 2014), this species should be classified in Lastreopsis, as has been done recently by most workers (e.g., Tindale, 1965; Tryon & Tryon, 1982). Acknowledgments This research was supported by a grant to Moran from the United States National Science Foundation (DEB-1020443), and to Labiak by a grant from CNPQ (no. 301997/2010-1). We thank Michael Grayum, Jefferson Prado, and an anonymous reviewer for their helpful comments on the manuscript.

Literature Cited Alston, A. H. G. 1959. The ferns and fern-allies of West Tropical Africa. Supplement to flora of West Tropical Africa. Ed. 2. Crown Agents for Oversea Governments and Administrations, London. Andrews, S. B. 1990. Ferns of Queensland. Queensland Department of Primary Industries Information Series Q189008, Brisbane, Australia. Brownlie, G. 1958. Chromosome numbers in New Zealand ferns. Transactions of the Royal Society of New Zealand 85: 213–216. ———. 1961. Additional chromosome numbers – New Zealand ferns. Transactions of the Royal Society of New Zealand (Botany) 1: 1–4. Brownsey, P. J. 1998. Incertae Sedis. Pp. 496. In: P. M. McCarthy (ed.), Flora of Australia, volume 48, Ferns, gymnosperms, and allied groups. ABRS/CSIRO, Melbourne. Christensen, C. 1919 [1920]. A monograph of the genus Dryopteris, Part II. The tropical American bipinnatedecompound species. Kongelige Danske Videnskabernes Selskabs Skrifter, Naturvidenskabelig of. Mathematiske Afdeling, ser. 8, 6: 3–132. Holttum, R. E. 1959. Vegetative characters distinguishing the various groups of ferns included in Dryopteris of Christensen’s Index Filicum, and other ferns of similar habit and sori. Gardens Bulletin, Singapore17: 361–367. Jones, D. L. 1998. Dryopteridaceae. Pp. 393–418. In: P. M. McCarthy (ed.), Flora of Australia, volume 48, Ferns, gymnosperms, and allied groups. ABRS/CSIRO, Melbourne. Labiak, P. H., M. Sundue, G. Rouhan, J. Garrison Hanks, J. T. Mickel & R. C. Moran. 2014. Phylogeny and historical biogeography of the lastreopsid ferns

86

BRITTONIA

(Dryopteridaceae). American Journal of Botany 101: 1207–1228. Moran, R. C. 1995. Lastreopsis. Pp.: 201–202. In: R. C. Moran & R. Riba (volume eds.). Psilotaceae a Salviniaceae. In: G. Davidse, M. Sousa S., & S. Knapp (general eds.), Flora Mesoamericana. Volumen 1. Univ. Nacional Autónoma de México, Ciudad Universitaria. Perrie, L. R. & P. J. Brownsey. 2012. Lastreopsis kermadecensis, a new species from Raoul Island in the Kermadec Islands, New Zealand, with notes on L. pacifica. New Zealand Journal of Botany 50: 29–36. Rakotondrainibe, F. & F. Tronchet. 2009. Révision synoptique du genre Lastreopsis Ching (Dryopteridaceae) à Madagascar. Adansonia, série 3, 31: 7–24. Roux, J. P. 2009. Synopsis of the Lycopodiophyta and Pteridophyta of Africa, Madagascar and neighbouring islands. Strelitzia 23. Pretoria: South African National Biodiversity Institute. Schuettpelz, E. H. & K. M. Pryer. 2007. Phylogeny of ferns inferred from 400 leptosporangiate species and three plastid genes. Taxon 56: 1037–1050. Smith, A. R. & R. C. Moran 1995. Megalastrum Holttum. In: R.C. Moran & R. Riba (eds.). Psilotaceae a Salviniaceae. In: G. Davidse, M. Sousa & S. Knapp (eds.). Flora Mesoamericana. Universidad Nacional Autónoma de México, Ciudad de México, v. 1, pp. 202–204. Smith, A. R., K. M. Pryer, E. Schuettpelz, P. Korall, H. Schneider & P. G. Wolf. 2006. A classification for extant ferns. Taxon 55: 705–731. ———, K. M. Pryer, E. Schuettpelz, P. Korall, H. Schneider & P. G. Wolf. 2008. Fern classification. Pp. 417–467. In: T. A. Ranker & C. H. Haufler (eds.), Biology and evolution of ferns and lycophytes. Cambridge: Cambridge University Press.

[VOL 67

Sundue, M., R. Hirai & J. Prado. 2013. Rumohra glandulosissima (Dryopteridaceae) a new species from the Atlantic Rainforest, and revision of the species occurring in Brazil. Systematic Botany 38: 915–924. Tryon, R. M. & A. F. Tryon. 1982. Ferns and allied plants, with special reference to tropical America. SpringerVerlag, New York. Tryon, R. M. & Stolze, R. G. 1991. Pteridophyta of Peru, part IV: 17. Dryopteridaceae. Fieldiana, Botany, new series 27: 1-176. Tindale, M. 1965. A monograph of the genus Lastreopsis Ching. Contributions from the New South Wales Herbarium, Flora Series 211: 59–78, tab. 1–23. ——— & S. K. Roy. 2002. A cytotaxonomic survey of the pteridophytes of Australia. Australian Systematic Botany 15: 839–937. Walker, T. G. 1966. A cytotaxonomic survey of the pteridophytes of Jamaica. Transactions of the Royal Society of Edinburgh 66: 169–237. ———. 1973. Evidence from cytology in the classification of ferns. Pp. 91–110. In: A. C. Jermy, J. A. Crabbe & B. A. Thomas (eds), The phylogeny and classification of the ferns. Botanical Journal of the Linnean Society 67 (Suppl. 1). ———. 1985. Cytotaxonomic studies of the ferns of Trinidad. 2. The cytological and taxonomic implications. Bulletin of the British Museum (Natural History) 13: 149–249. Yeuhong, Y., Q. Xinping & S. Serizawa. 2013. Pp. 158–168. Microlepia. In: W. Zhengyi, P. H. Raven & H. Deyuan (eds.), Flora of China, vols. 2–3, Lycopodiaceae through Polypodiaceae.