A REVISION OF THE MASCARENE SPECIES OF ELAPHOGLOSSUM (ELAPHOGLOSSACEAE) 1 David H. Lorence 2 and Germinal Rouhan 3,4 ABSTRACT A taxonomic revision of Elaphoglossum in the Mascarene Islands (southwest Indian Ocean) is presented here based on a treatment written for the Flore des Mascareignes. Seventeen species are recognized for the islands of Mauritius and Réunion. Of these species, seven are endemic. The non-endemic species also occur in Madagascar, Africa, or the Paleotropics in general. In addition, five natural interspecific hybrids (nothospecies) occur in the Mascarenes. Two lectotypes are designated for the name Acrostichum inversum and are referred pro parte to E. acrostichoides and E. angulatum. RÉSUMÉ Une révision taxinomique d’Elaphoglossum des ı̂les Mascareignes (sud-ouest de l’Océan Indien) est présentée ici, fondée sur un traitement écrit pour la Flore des Mascareignes. Dix-sept espèces dont sept endémiques sont reconnues pour les ı̂les de Maurice et de La Réunion. Les espèces non-endémiques se trouvent également à Madagascar, en Afrique, ou dans les Paléotropiques en général. De plus, cinq hybrides interspécifiques naturels (notho-espèce) sont présents dans les Mascareignes. Deux lectotypes sont désignés pour le nom Acrostichum inversum et sont attribués pro parte à E. acrostichoides and E. angulatum. Key words: Elaphoglossaceae, Elaphoglossum, hybrids, Mascarene Islands, Mauritius, nothospecies, pteridophytes, Réunion, southwest Indian Ocean. The Mascarenes are a trio of volcanic oceanic islands situated in the southwest Indian Ocean. Approximate sizes and ages (millions of years) of the islands are: Réunion, 2500 km2, 3.0 MA; Mauritius, 1865 km2, 7.8 MA; and Rodrigues, 110 km2, 1.5 MA (Fisher et al., 1967; McDougall & Chamalaun, 1969; McDougall et al., 1965). The mean annual precipitation ranges from 1000 to 6000 mm in Mauritius and Réunion and from 1100 to 1700 mm in Rodrigues. The rainy, austral summer season punctuated by occasional tropical cyclones lasts from December to April, and the relatively drier and cooler winter season lasts from May to Novem- ber (Lorence, 1985). Except in coastal areas with coral sand, Mascarene soils are derived from decomposed basaltic lavas. Relative to their small surface area the Mascarenes possess a diverse pteridophyte flora, with 273 species comprising 282 taxa including infraspecies (Badré & Cadet, 1978; Lorence, 1976b, 1978, 1992). Of these, 265 species are indigenous and 8 are adventive or naturalized. Of the indigenous species, 22.7% are endemic to the Mascarene archipelago or to a single island. Among the most species-rich fern genera on the island is Elaphoglossum Schott ex J. Sm., a large 1 Use of facilities and financial assistance provided by the Mauritius Sugar Industry Research Institute at Réduit, Mauritius, are gratefully acknowledged while the first author served as a U.S. Peace Corps Volunteer at the Mauritius Herbarium (MAU). A portion of the fieldwork by the second author was supported by the ‘‘PPF Populations fractionnées et insulaires de l’Ecole Pratique des Hautes Etudes.’’ We thank the curators of the following herbaria for making their specimens available for study: B, BM, K, MAU, MO, MPU, P, PR, PRC, PTBG, and REU. Frédéric Badré (P) provided encouragement and reviewed an earlier version of the manuscript. We are grateful to Brigitte Zimmer (B) for providing digital images of the collections in the Willdenow Herbarium (B-W) and for helpful suggestions and comments. We thank Victoria Hollowell, John Mickel, Robbin Moran, and Henk van der Werff whose comments and reviews greatly improved the quality of the paper. 2 National Tropical Botanical Garden, 3530 Papalina Road, Kalaheo, Hawaii 96741, U.S.A. lorence@ntbg.org. 3 Département Systématique et Evolution USM 0602, Herbier Plantes Vasculaires, Muséum National d’Histoire Naturelle, 57 rue Cuvier, 75231 Paris, France. rouhan@mnhn.fr. 4 Equipe Classification, Evolution et Biosystématique (EA 3496), UMR 5143, Université Pierre et Marie Curie, 12 rue Cuvier, 75005 Paris, France. ANN. MISSOURI BOT. GARD. 91: 536–565. 2004. Volume 91, Number 4 2004 Lorence & Rouhan Mascarene Elaphoglossum and taxonomically complex pantropical genus. Worldwide, the genus comprises about 600 species of primarily epiphytic, lithophytic, and casually terrestrial ferns concentrated in tropical wet forests and cloud forests (Mabberley, 1997; Mickel & Atehortúa, 1980; Mickel & Beitel, 1988). Although the greatest species diversity occurs in the Andes of South America, Elaphoglossum has undergone slight adaptive radiation in a number of oceanic islands and archipelagos such as Hawaii with 9 species (Palmer, 2003) and the Mascarene Islands with 17 species (Lorence, 1978, 1984). In the Mascarenes, Elaphoglossum occurs only on the two largest and highest islands, Mauritius and Réunion: it is absent from the smaller and drier Rodrigues Island (Lorence, 1976b). Of the 17 Mascarene species, 7 are endemic and 10 occur elsewhere primarily in Madagascar, the Comoro Islands, Africa, or the Paleotropics in general. In addition to these 17 species, five natural hybrids (nothospecies) are known, indicating the genus is in an active state of evolution in the Mascarenes (Lorence, 1984). The following revision of Elaphoglossum is based on a treatment of Lomariopsidaceae s.l. originally written in French for the Flore des Mascareignes, a joint collaborative project between the Royal Botanic Gardens Kew, the Mauritius Herbarium at the Mauritius Sugar Industry Research Institute, and Institut de Recherche pour le Développement (IRD, formerly ORSTOM) based at the Paris Muséum National d’Histoire Naturelle. Because publication of the pteridophyte fascicles for this project has not yet taken place, we decided to publish this revision to make the information available in a timely manner. In addition, we provide supplemental habit photos and drawings of rhizome scales to facilitate identification. collapsed, irregular spore shape and size) are suggestive of hybrid origin as already documented for other pteridophyte genera (Wagner & Chen, 1965; Wagner et al., 1986). Nevertheless, aborted spores alone are not an absolute criterion of hybridism because there are several other possible sources of abortion such as the metabolic conditions of the plant, which may be influenced by cold or drought shocks during spore development (Hennipman, 1977). MATERIALS AND METHODS This taxonomic revision is based on 717 herbarium specimens and field observations of all the Elaphoglossum species occurring in the Mascarenes. All measurements given in the descriptions have been taken from dried herbarium specimens. Unacetolized spores were mounted in glycerine jelly, and their size measurements were taken excluding the perispore. Secondary or lateral veins that are apically free versus united provide an important, species-specific character, but the fronds must be held up to a bright light or, in some cases, chemically cleared to see the venation. Collections with morphological characters intermediate between two species that occur in the same vicinity and have a high percentage of aborted spores (features such as 537 TAXONOMIC HISTORY The name Elaphoglossum was first published, without description, by Schott (1834), and was not validated until Smith (1841) attributed the name to Schott and published a description and cited a type species. Morton (1965) proposed Elaphoglossum Schott ex J. Sm. as a conserved generic name against Peltapteris Link, which was also published in 1841. Elaphoglossum is characterized by its dimorphic fronds with acrostichoid sori (in addition, nearly all the species have an elongated ventral meristele in the rhizome, simple and entire laminae, and free veins). Because of its soral condition, many species of Elaphoglossum were originally described under Acrostichum L. (e.g., Bojer, 1837; Fée, 1845; Baker, 1877; Cordemoy, 1895). This proved to be a polyphyletic grouping, as the type species Acrostichum aureum L., which also has acrostichoid sori, belongs to the Pteridaceae (e.g., Tryon & Tryon, 1982). Most taxa of Elaphoglossum originally described in Acrostichum were subsequently transferred to Elaphoglossum, or to Aconiopteris C. Presl or Hymenodium Fée (now both considered synonyms of Elaphoglossum). Most of these transfers were made by Moore (1857–1862). The difficulties in establishing phylogenetic relationships of Elaphoglossum to other ferns may explain why in modern times the genus has been taxonomically treated along diverse lines, namely: (1) as a part of the family of Lomariopsidaceae Alston (Alston, 1956; Kramer et al., 1990; Moran & Riba, 1995), which is the concept retained within African literature (e.g., Alston, 1959; Tardieu-Blot, 1959, 1960; Schelpe, 1969; Roux, 2001; Mickel, 2002); (2) as a member of the family Elaphoglossaceae described by Pichi Sermolli (1968); (3) as a member of a subfamily: Aspleniaceae Mett. ex A. B. Frank subfam. Elaphoglossoideae (Pic. Serm.) Crabbe, Jermy & Mickel (Crabbe et al., 1975); and (4) as a member of the tribe Bolbitideae Pic. Serm. included in the family Dryopteridaceae Herter (Tryon & Tryon, 1982). 538 Annals of the Missouri Botanical Garden The most recent comprehensive systematic account of the genus as a whole is the Monographie des Genus Elaphoglossum (Christ, 1899). This treatment covered 134 species and detailed 32 taxonomic groups. More recently Mickel and Atehortúa (1980: 47) subdivided the genus into sections and subsections, but they emphasized that ‘‘this treatment is provisional because a complete understanding of the genus can be had only after the species are better known.’’ Even with the first phylogenetic studies of the genus (Rouhan et al., 2004), knowledge of the systematics and phylogeny of Elaphoglossum is far from complete, and new species continue to be described. The present paper aims at providing new knowledge and a modern taxonomic treatment of the genus in the Mascarene region. and typus conservandus (ICBN, App. IIIA; Voss et al., 1983: 296) against Aconiopteris, which is based on a type different from that of the conserved name Elaphoglossum (Art. 14.4) as follows: Aconiopteris C. Presl. [Tent. Pterid.] in Abh. Königl. Böhm. Ges. Wiss., ser. 4, 5: 236, tab. 10(17). 1836 (ante 2 Dec.). TYPE: Aconiopteris subdiaphana (Hook. & Grev.) C. Presl, Tent. Pterid. 236, tab. 10(17). 1836 [Acrostichum subdiaphanum Hook. & Grev., Icon. Filic. 2, tab. 205. 1831]. Small to medium-sized epiphytic, epilithic, or occasionally terrestrial ferns; rhizome short- to long-creeping, branching or not, dictyostelic, dorsiventral or radial, fronds in 2 or 3 to 5 ranks (Mascarene species), paleaceous, scales basifixed. Fronds simple (all Mascarene taxa), very rarely flabellately lobed, with stipe jointed and swollen into a persistent, dark phyllopodium bearing small lateral aerenchymatous outgrowths when young; blade simple, entire, usually erect but sometimes pendent, chartaceous to coriaceous, glabrate to densely paleaceous, scales often deciduous; secondary (lateral) venation pinnate, simple or dichotomous, tips apically free and thickened or uniting into a submarginal vein or commissure, rarely branching and anastomosing reticulately (not in Mascarene taxa), margin often revolute. Fertile frond similar in shape to sterile, but blade usually smaller and longer stiped; sporangia densely covering abaxial surface of blade except along costa and margin. Spores monolete, bilateral, elliptic to reniform, exine smooth, perispore granular to spiny forming folds or crests. Gametophyte subcordate or ribbon-like, margins undulate or crisped and bearing unicellular hairs with waxy tips, rhizoids reddish brown. Chromosome number x 5 41 (Manton & Sledge, 1954). The generic name Elaphoglossum refers to the tongue-like shape of the laminae (from the Greek words elaphos 5 stag, and glossa 5 tongue). In the Mascarene Islands most species grow as low to high epiphytes or are casually terrestrial in moist and wet forests and ericoid (heath) formations. Where conditions are satisfactory, the plants usually root in mats of bryophytes on trees, at bases of shrubs and trees, or on mossy logs or boulders. One species, E. lanatum, is restricted to soil or rock of shady stream banks and cliff faces, and E. spatulatum grows specifically on large boulders in stream beds. The Mascarenes harbor seven endemic Elaphoglossum species. This suggests the genus is actively evolving in this archipelago (Lorence, 1984), especially given the relatively recent emergence of these islands. Furthermore, five natural TAXONOMIC TREATMENT Elaphoglossum Schott ex J. Sm., J. Bot. (Hook.) 4(27): 148. Aug. 1841, nom. et typ. cons. TYPE: Elaphoglossum conforme (Sw.) J. Sm., J. Bot. (Hook.) 4(27): 148. 1841 [Acrostichum conforme Sw., Syn. Fil. (Sw.) 10, 192. 1806]. Hymenodium Fée, Mém. Foug., 2: 20. 1844 [1845]. TYPE: Hymenodium crinitum (L.) Fée, Mém. Foug., 2: 90. 1844 [1845] [Acrostichum crinitum L., Sp. Pl. 2: 1068. 1753]. Peltapteris Link, Fil. Spec. 147. 1841. TYPE: Peltapteris peltata (Sw.) C. V. Morton, Amer. Fern J. 45: 13. 1955 [Osmunda peltata Sw., Prodr. 127. 1788]. Microstaphyla C. Presl, Abh. Königl. Böhm. Ges. Wiss., ser. 5, 6: 520–521. 1851. TYPE: Microstaphyla bifurcata (Jacq.) C. Presl, Epimel. Bot. 161. 1849 [Osmunda bifurcata Jacq., Collectanea 3: 282, tab. 20(4). 1789]. The following discussion clarifies the nomenclature and the typification of the genus Elaphoglossum. H. W. Schott was the first to propose the name Elaphoglossum (Gen. Fil.: 30, tab. 13. 1834), but this name lacks a description and consequently was not validly published. Validly publishing Elaphoglossum, J. Smith attributed the name to Schott (J. Bot. 4(27): 148. 1841) and referred six species to the genus, but he did not select one of them as the type. In a subsequent publication (Hist. Fil.: 125. 1875), J. Smith clearly chose E. conforme as the type. Since the date of publication of Elaphoglossum is 1841 and not 1834, Aconiopteris, which was validly published in 1836, should have priority over Elaphoglossum when these genera are united. In order to avoid replacing the well-known generic name Elaphoglossum by Aconiopteris, the name Elaphoglossum and its type E. conforme (Sw.) J. Sm. are conserved respectively as nomina conservanda Volume 91, Number 4 2004 Lorence & Rouhan Mascarene Elaphoglossum hybrids occur in the Mascarenes (see Lorence, 1984, for detailed discussion concerning the im- plications of the occurrence of these hybrids in the Mascarenes). KEY TO THE MASCARENE SPECIES OF 539 ELAPHOGLOSSUM 1a. Sterile blades glabrous or bearing (at least when young) rare to scattered, minute, appressed scales 0.5 mm or less long with margins bearing short multicellular hairs with bulbous, glandular tips. 2a. Sterile blades broadly elliptic to ovate, usually 3 cm wide or broader. 3a. Rhizome scales, filiform or bristle-like, dark reddish brown or black. 4a. Rhizome scales 3–5 mm long, stiff, shiny, black; veins in part apically free and in part uniting intramarginally at tips ----------------------------------------------------------------------- 14. E. 3revaughanii 4b. Rhizome scales 8–20 mm long, flexible, dull, brown to black; veins uniting apically into a submarginal commissure -------------------------------------------------------------------------------------------- 18. E. sieberi 3b. Rhizome scales broadly ovate to lanceolate, pale orange-brown or tan. 5a. Rhizome scales broadly ovate, the base auriculate with overlapping lobes; rhizome longcreeping ----------------------------------------------------------------------------------------------------------------- 3. E. angulatum 5b. Rhizome scales narrowly ovate to oblong, lacking overlapping basal lobes; rhizome shortcreeping. 6a. Rhizome scales 4–10 mm long, 1–1.5 mm wide, curling, with margins involute and bearing long, gland-tipped projections, these more numerous toward the base; veins free at tips ----------------------------------------------------------------------------------------------- 12. E. lepervanchei 6b. Rhizome scales (10–)15–25 mm long, 2–3 mm wide, flat, with margins entire or bearing rare short projections; veins uniting apically into submarginal commissure --------------------------------------------------------------------------------------------------------------------------- 13. E. macropodium 2b. Sterile blades narrowly elliptic, oblong or linear, usually less than 3 cm wide. 7a. Surfaces of sterile blade bearing scattered, persistent, light brown resinous punctations (seen under magnification, but not visible on all adult specimens); veins uniting apically into a submarginal commissure ------------------------------------------------------------------------------------------------------- 15. E. richardii 7b. Surfaces of sterile blade lacking resinous punctations; veins free apically. 8a. Rhizome scales thick, dark brown to black, opaque except paler basally. 9a. Sterile frond with blade at least 15 cm long ---------------------------------------------------- 6. E. coursii 9b. Sterile frond with blade 6–14 cm long --------------------------------------------------------- 5. E. 3cadetii 8b. Rhizome scales thin, papery, pale to medium brown, translucent or sometimes opaque apically. 10a. Rhizome scales with darker bands toward apex ----------------------------------- 1. E. acrostichoides 10b. Rhizome scales uniformly pale golden or reddish brown. 11a. Rhizome long-creeping, often branching; fronds spaced 1.5 cm or more apart; rhizome scales flat, broadly ovate, entire ----------------------------------------- 3. E. angulatum 11b. Rhizome short- to moderately long-creeping, rarely branched; fronds spaced 1.2 cm or less apart; rhizome scales crisped, involute, narrowly ovate to linearoblong, margins ciliate especially basally ----------------------------------- 12. E. lepervanchei 1b. Sterile blades bearing (at least when young) on both surfaces and/or the margins numerous conspicuous scales, either subulate, laminate, peltate, or substellate. 12a. Scales of sterile frond stellate or substellate, with 4 to 8 straight hair teeth ------------------ 11. E. lancifolium 12b. Scales of sterile blade basifixed or peltate, or subulate. 13a. Scales of sterile blade subulate, margins entire or occasionally minutely ciliate apically. 14a. Scales of sterile blade dark brown, confined to margins and costa -------------------- 9. E. hybridum 14b. Scales of sterile blade pale brown to reddish brown, scattered over both surfaces of the blade. 15a. Sterile blades linear, thinly chartaceous and translucent, veins readily visible when held up to light ------------------------------------------------------------------------------------------------- 4. E. aubertii 15b. Sterile blades elliptic, narrowly obovate, or spatulate, thickly chartaceous, veins not visible when held up to light. 16a. Sterile blades spatulate to narrowly obovate, 1.5–5.5 3 0.5–1.2 cm, apex obtuse ----------------------------------------------------------------------------------------------------------- 19. E. spatulatum 16b. Sterile blades elliptic, 7–10 3 1.5–2 cm, apex acute ----------------- 17. E. 3setaceum 13b. Scales of sterile blade peltate or basifixed with flat central portion, the margins fimbriate or strongly ciliate. 17a. Scales of sterile blade closely appressed, suborbicular, scarious, densely fimbriate; rhizome scales with margins scarious or fimbriate. 18a. Rhizome scales 2–3 mm long, claw-like, opaque, dull brown with fimbriate margins; sterile blade with veins all uniting apically into a submarginal commissure ---------------------------------------------------------------------------------------------------------------------------------- 22. E. tomentosum 18b. Rhizome scales 6–12 mm long, ovate-oblong with long, bristle-like apex, shiny castaneous to black with pale scarious margins; sterile blade with veins free (some uniting apically in E. 3heterophlebium). 540 Annals of the Missouri Botanical Garden 19a. Sterile blade with veins free apically with thickened tips; rhizome scales black with scarious transparent margins ------------------------------------------------------ 7. E. heterolepis 19b. Sterile blade with veins uniting apically into submarginal loops, those toward base and apex of blade free with thickened tips; rhizome scales castaneous with sparsely dentate or glandular-ciliate margins ------------------------- 8. E. 3heterophlebium 17b. Scales of sterile blade spreading, toothed with usually paired hair teeth; rhizome scales with margins bearing (at least basally) short hair teeth or sessile glands. 20a. Sterile blades paleaceous only on adaxial surface, rarely a few scales present on abaxial surface along costa ------------------------------------------------------------------------ 21. E. stipitatum 20b. Sterile blades paleaceous on both surfaces, at least when young. 21a. Fronds 2-ranked on rhizome, not densely caespitose; sterile blade covered when young with densely matted scales bearing long, tangled marginal cilia, the scales soon deciduous. 22a. Scales on adaxial surface of sterile blade forming a loose tomentum --------------------------------------------------------------------------------------------------------------- 10. E. lanatum 22b. Scales on adaxial surface of sterile blade closely appressed forming a thin tomentum ----------------------------------------------------------------------------- 2. E. 3adulterinum 21b. Fronds 2- to 5-ranked on rhizome, usually densely caespitose; sterile blade with persistent scales bearing stiff, marginal long hair teeth. 23a. Sterile blade with both surfaces obscured by a layer of reddish brown scales, with dark brown to black scales often present along abaxial surface of costa --------------------------------------------------------------------------------------- 20. E. splendens 23b. Sterile blade with both surfaces visible, covered with loose, pale reddish brown scales along the veins -------------------------------------------------- 16. E. rufidulum 1. Elaphoglossum acrostichoides (Hook. & Grev.) Schelpe, J. S. African Bot. 30: 196. 1964. Vittaria acrostichoides Hook. & Grev., Icon. Filic. 2: tab. 186. 1830. Drymoglossum acrostichoides (Hook. & Grev.) T. Moore, Index Fil.: 31. 1857. TYPE: South Africa. Cape of Good Hope, D. Thom s.n. [Hb. Hook] (holotype, K!). Figure 1A. Elaphoglossum falcatum (Fée) T. Moore, Index Fil.: 9. 1857. Acrostichum falcatum Fée, Mém. Foug., 2: 44, tab. 21(1). 1844 [1845]. TYPE: Bourbon [Réunion]. Lépervanche-Mézières & Richard s.n. [Hb. Bory 27,49] (holotype, P!). Elaphoglossum inversum (Cordem.) H. Christ, Neue Denkschr. Allg. Schweiz. Ges. Gesammten Naturwiss. 36(1): 146. 1899. Acrostichum inversum Cordem., Fl. Réunion, part 1: 84. 1891. TYPE: Réunion. Grande Montée de la Plaine des Cafres, 1899, E. J. de Cordemoy s.n. pro parte, 2 right-hand specimens only (lectotype, designated here, P!). Fronds moderately spaced 0.8–1.5 cm distant, borne in 2 ranks; rhizome long-creeping, 2–4 mm diam., occasionally branching, densely paleaceous; scales 3–5 3 1.2–1.6 mm, ovate to ovate-elliptic, brown with dark brown bands at base and apex (or often almost completely dark and only margins clearer), very often lustrous, base truncate, cordate, or auriculate, apex acuminate, margins entire or bearing toward base short teeth or bulbous-tipped cilia, cells small, rectangular. Phyllopodia to 6–10 mm long, slightly swollen, brown to dark brown, paleaceous; sterile fronds 10–25(–41) cm; stipe 3–12 cm, canaliculate, narrowly winged distally, brown to blackish, bearing scattered caducous scales like those of rhizome; blade narrowly elliptic to oblong, often somewhat falcate, 7–16(–26) 3 0.7–2(–3) cm, base narrowly cuneate to attenuate, apex acute to long-acuminate, coriaceous to subcoriaceous, costa prominulous, rounded abaxially, canaliculate adaxially, margins revolute, veins distinct to somewhat indistinct, free with thickened tips, adaxial surface of blade glabrate, abaxial surface of blade bearing scattered, brown, ovate scales 3–4 3 2 mm with margins dentate to ciliate mixed with smaller substellate scales with glandular marginal cilia. Fertile frond 9– 26(–40) cm long; stipe slender, about half total frond length; blade 6–16(–25) cm, narrowly elliptic, falcate, base narrowly cuneate, decurrent, apex acute. Spores 35–40 3 20–25 mm (excluding perispore), perispore cristate, crests 3–6 mm high, with entire to finely sinuate margins. Distribution and habitat. Réunion; also known from Madagascar, the Comoro Islands, tropical and southern Africa, Sao Tomé, and Fernando Po. In Réunion E. acrostichoides occurs from 1200 to 2300 m in moist and wet forests and ericoid formations as an epiphyte and saxicolous on mossy rocks. Discussion. Elaphoglossum acrostichoides greatly resembles E. coursii by its general appearance, but differs by paler, less stiff rhizome scales. Elaphoglossum acrostichoides resembles E. angulatum by the long-creeping rhizome, but differs by less widely spaced fronds. Elaphoglossum acrostichoides can be distinguished from the other species in the Mascarenes by non-appressed, dark-streaked rhizome and stipe scales. In Réunion it hybridizes with E. coursii (Lorence, 1984), forming E. 3cadetii Volume 91, Number 4 2004 Lorence & Rouhan Mascarene Elaphoglossum 541 Figure 1. Habit of some Mascarene Elaphoglossum species. All the illustrations show specimens with sterile and fertile fronds. —A. E. acrostichoides from Réunion, Rouhan & Grangaud 249. —B. The long-creeping rhizome of E. angulatum from Réunion, Rouhan 216. —C. E. richardii from Réunion, Rouhan et al. 231. —D. E. aubertii from Réunion, Rouhan et al. 241. —E. E. sieberi from Mauritius, Brise Fer Reserve. Scale bars 5 2 cm. 542 Annals of the Missouri Botanical Garden Lorence. Elaphoglossum acrostichoides resembles E. glabellum J. Sm. from the Neotropics in nearly all characters; certain specimens of E. acrostichoides are indistinguishable from the Neotropical ones (Moran & Smith, 2001). We found in the Paris herbarium only one sheet bearing the name Acrostichum inversum along with the annotations ‘‘La Réunion’’ and ‘‘Jacob de Cordemoy,’’ and the date ‘‘1899’’ on the label is subsequent to the original publication (1891). Of the four specimens included on this sheet, the two right-hand specimens only are Elaphoglossum acrostichoides (the two left-hand remaining specimens being E. angulatum (Blume) T. Moore), necessitating the need to designate a lectotype (ICBN, Art. 9.9; Greuter et al., 2000). We hereby designate these two right-hand specimens of E. J. de Cordemoy s.n. as the lectotype of Acrostichum inversum Cordem., herein referred to E. acrostichoides (Hook. & Grev.) Schelpe. 48.5 cm; stipe (8–)10–15 cm, 0.5–1 mm diam., stramineous, canaliculate, bearing scattered, appressed, brown caducous ovate scales 1–2 3 0.5 mm, base obtuse, apex acute, margins scarious, glandular-ciliate, these intermixed with smaller, matted scales; blade linear-oblong to linear, (19–) 25–32 3 (1.4–)1.8–2.5 cm, base narrowly cuneate, decurrent, apex acute to acuminate, subcoriaceous, costa stramineous, prominulous and rounded abaxially, shallowly canaliculate adaxially, both surfaces covered with a homogenous layer of thin, matted, appressed pale brown to buff-colored, ovate to subcircular scales 1–1.5 3 1 mm, basifixed, base rounded to cordate or sagittate, base and margins bearing long, spreading arachnoid cilia, pedicel and center darker brown, subtended by cluster of globose glands, cells hyaline, thin-walled, adaxial surface of lamina glabrescent, veins unbranched or bifurcate, in part free apically with tips thickened, in part uniting submarginally, both surfaces of blade with scattered hair-toothed to deltoid scales 0.2–0.5 mm long, glandular-ciliate at base. Fertile frond 20–25 cm long; stipe 10–11.5 cm, 1 mm diam.; blade 11–14 3 0.7–1 cm, linear-elliptic, chartaceous, base narrowly cuneate, apex acute, adaxial surface with dense layer of matted scales as in sterile frond, margins revolute. Spores 82% aborted, normal spores 35–38 3 25–27 mm (excluding perispore), perispore cristate, crests low, 1.5–3 mm high, sinuate to finely erose-dentate. Selected specimens examined. RÉUNION. Bébour, Lorence in MAU 15623 (MAU); near Rivière des Marsouins, Lorence in MAU 15614 (MAU); path to Piton de Bébour, Lorence in MAU 15621 (MAU); Col de Bébour, Cadet 4028 (REU); Boucan Launay, Boivin 801 (P); Cilaos, au dessus des Thermes, Cadet 4122 (REU); Grand Matarum, Cadet 805 (P); pente du Cirque, Cadet 1929A (P); sentier GR R1 entre ‘‘Le Bloc’’ et le gı̂te de la Caverne Dufour, Rouhan 215 (NY, P); Cirque de Salazie, Rouhan et al. 229 (P, PTBG); Dimitile, Rouhan & Grangaud 204 (P); Grand Bénard, Pic Maı̈do, Cadet 1749 (P); Grand Matarum, Lorence in MAU 15637 (MAU); Ilet à Guillaume, Badré 866 (P); Piton des Neiges, Lorence in MAU 15648 (MAU, MO); sentier de la rivière, Rouhan & Grangaud 249 (NY, P, PTBG); Plaine des Fougères, Oct. 1850, Boivin s.n. (P); Salazie, Lépervanche-Mézières s.n. (P); sentier de la Roche Ecrite, Badré s.n. (P); St. Paul, sentier de l’ı̂let Alcide, Rouhan et al. 242 (NY, P, PTBG); volcan (Massif de la Fournaise), Pas de Bellecombe, Lorence & Lorence 2520 (MO). 2. Elaphoglossum 3adulterinum Lorence [Elaphoglossum lanatum (Bojer ex Baker) Lorence 3 E. tomentosum (Bory ex Willd.) H. Christ], Fern Gaz. 12: 347, fig. 4. 1984. TYPE: Mauritius. Valley of Cascade 500 Pieds (Cascade Alexandra), 1974, D. H. Lorence 874 (holotype, MAU 16309!). Fronds caespitose, spaced 3–5 mm distant, borne in 2 ranks; rhizome short-creeping, 2–3 mm diam., rarely branched, densely paleaceous apically; scales 4–6 3 0.4–0.6 mm, narrowly ovate to lanceolate, castaneous, opaque, base cordate to auriculate, entire or bearing clear, globose marginal glands, apex narrowly acuminate, filiform, sinuate, margins sparsely glandular dentate-ciliate, cells opaque. Phyllopodia 6–8 mm long, thickened, dark brown, paleaceous basally; sterile fronds (27–)35– Distribution and habitat. Elaphoglossum 3adulterinum is known only from the type locality in Mauritius at the southern extremity of the central plateau, altitude 600 m. The hybrid occurs on boulders below cliffs in wet forests where the parental species E. lanatum and E. tomentosum occur nearby (Lorence, 1984). Discussion. Elaphoglossum 3adulterinum resembles E. tomentosum in its habitat preference (epiphytic or epipetric), erect fronds, and sterile blade shape, although its rhizome scales are glandular and resiniferous as in E. lanatum. The hybrid’s mixed venation, in part free as in E. lanatum and in part uniting intramarginally as in E. tomentosum, and intermediate frond scale morphology distinguish it from the parents. Specimens examined. MAURITIUS. Valley of Cascade 500 Pieds (Cascade Alexandra), Lorence sub MAU 16307 (MAU, MO), Lorence 1609 (MO), Lorence 1610 (MO), Lorence 847 (MAU). 3. Elaphoglossum angulatum (Blume) T. Moore, Index Fil.: 5. 1857. Acrostichum angulatum Blume, Enum. Pl. Javae 101. 1828; Fl. Javae Volume 91, Number 4 2004 3: 25 (1828), tab. 6 exhibiting a drawing of the type specimen. Olfersia angulata (Blume) C. Presl, Tent. Pterid. 234. 1836. TYPE: Java. C. L. von Blume s.n. (holotype, K!). Figure 1B. Elaphoglossum alstonii Tardieu, Notul. Syst. 15: 433. 1959. Acrostichum laurifolium Fée, Mém. Foug., 2: 36, tab. 7. 1844 [1845], non Acrostichum laurifolium Thouars, Esquisse Fl. Tristan d’Acugna: 31. 1803. TYPE: Bourbon [Réunion]. 1840, Lépervanche-Mézières 14 [Hb. Bory 27,15] (holotype, P!). Elaphoglossum inversum (Cordem.) H. Christ, Neue Denkschr. Allg. Schweiz. Ges. Gesammten Naturwiss. 36(1): 146. 1899. Acrostichum inversum Cordem., Fl. Réunion, part 1: 84. 1891. TYPE: Réunion. Grande montée de la Plaine des Cafres, 1899, E. J. de Cordemoy s.n. pro parte, 2 left-hand specimens only (lectotype, designated here, P!). Fronds widely spaced 1.5–4 cm distant, erect, borne in 2 ranks; rhizome always very long-creeping, 2–3 mm diam., occasionally branching, clothed with appressed scales; scales 3–5 3 2–3 mm, ovate, thin, light brown or reddish brown, base auriculate with overlapping lobes, apex acute to acuminate, margins entire or bearing toward base rare, short bulbous-tipped cilia, cells squarish. Phyllopodia to 15 mm long, slightly swollen, brown to dark brown, paleaceous; sterile fronds 20–40 cm; stipe 9–17 cm, canaliculate, brown to orange, bearing scattered spreading, light brown, ovate scales; blade narrowly elliptic to narrowly ovate, 10–16 3 2–4 cm, base narrowly cuneate, shortly decurrent, apex acute to acuminate, coriaceous to subcoriaceous, costa prominulous, rounded to angled abaxially, canaliculate adaxially, margins revolute, veins often indistinct, in part free apically with thickened tips and in part uniting into an intramarginal commissure, adaxial surface of blade glabrate, abaxial surface of blade bearing a few scattered, ovate scales 1–3 mm long. Fertile frond 20–40 cm long; stipe as in sterile, blade as in sterile but narrower. Spores 28–30 3 20–23 mm (excluding perispore), perispore cristate, crests 4–5 mm high, with entire to finely sinuate margins. Distribution and habitat. Réunion; also Madagascar, tropical Africa, southern India, Sri Lanka, Java, Sumatra, Borneo, New Guinea, Philippines, New Caledonia, and Taiwan. In Réunion, Elaphoglossum angulatum occurs as an epiphyte or casually terrestrial in ericoid formations from 1300 to 2200 m. Discussion. This species resembles Elaphoglossum acrostichoides by the long-creeping rhizome and general appearance, but differs by more widely spaced fronds. E. angulatum can also be distinguished by its ovate, appressed, uniformly colored, light brown or reddish brown rhizome and stipe Lorence & Rouhan Mascarene Elaphoglossum 543 scales, by the veins in part uniting into an intramarginal commissure, and by the erect fronds (see Sledge, 1967, for more details). The type of Acrostichum angulatum is from Java and was collected by Blume. Carl Ludwig von Blume worked at Leiden (L), and according to Stafleu and Cowan (1976) all his collections were deposited in L. Nevertheless, many specimens were distributed from Leiden after his death, and not always with the regard we now have for type specimens (P. Hovenkamp (L), pers. comm.). There is a specimen of Blume s.n. housed in K and bearing a label ‘‘Herb. Lugd. Batav.’’ with the mention ‘‘Acrost. angulatum’’ written by Blume. Furthermore, this specimen matches well the drawing of the type specimen in Flora Javae. For these reasons, this specimen of Blume s.n. housed in K is identified as the holotype of Acrostichum angulatum Blume. Concerning the typification of Acrostichum inversum, although the date ‘‘1899’’ written on the label is posterior to the original publication (1891), we found in the Paris herbarium only one sheet bearing the name Acrostichum inversum along with the annotations ‘‘La Réunion’’ and ‘‘Jacob de Cordemoy.’’ Of the four specimens included on this sheet, the two left-hand specimens only are Elaphoglossum angulatum (the two right-hand remaining specimens are E. acrostichoides), necessitating the need to designate a lectotype (ICBN, Art. 9.9; Greuter et al., 2000). We hereby designate these two left-hand specimens of E. J. de Cordemoy s.n. as the lectotype of Acrostichum inversum Cordem., here referred to E. angulatum (Blume) T. Moore. Selected specimens examined. RÉUNION. Bébour, near Rivière des Marsouins, Lorence in MAU 15640 (MAU, MO); Cilaos, sentier GR R1 entre ‘‘Le Bloc’’ et le gı̂te de la Caverne Dufour, Rouhan 216 (NY, P, PTBG); pente du Cirque, Cadet 1930 (P); Plaine des Palmistes rd., P. O. Wiehe 1721 (MAU); Piton des Neiges, Lorence in MAU 15612 (MAU), Lorence in MAU 15615 (MAU), Lorence 408 (MO); Plaine des Cafres, près de la source Reillac, Cadet 1778 (P); Plaine des Fougères, Rouhan et al. 233 (P); Plaine des Palmistes, la Grande Montée, Cadet 504 (P); volcan, Enclos Fouqué, Cadet 1556 (P, top 3 specimens); volcan, Pas de Bellecombe, Rouhan & Grangaud 220 (NY, P). 4. Elaphoglossum aubertii (Desv.) T. Moore, Index Fil.: 5. 1857. Acrostichum aubertii Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 309. 1811. TYPE: Bourbon [Réunion]. 1808, A. Du PetitThouars s.n. (holotype, P-JU 992!). Figure 1D. Fronds caespitose, borne in 3 ranks; rhizome very short-creeping, 2–3 mm diam., rarely branched, densely paleaceous; scales 6–8 3 0.5– 544 Annals of the Missouri Botanical Garden 1 mm, narrowly ovate to oblong, dark reddish brown, base cordate, apex narrowly acuminate, margins entire, cells fusiform, to very linear for the marginal cells. Phyllopodia 2–3 mm long, slightly flattened and constricted, dark brown to black; sterile fronds 10–30 cm; stipe 2–6 cm, 0.5–1 mm diam., stramineous, canaliculate, bearing scattered to abundant, reddish brown, narrow, spreading subulate scales 3–5 mm long, 0.25–0.5 mm wide, base tubular and closed, apex long-acuminate, margins with short, glandular-tipped cilia basally, entire above; blade linear-oblong, 8–24 3 0.8–1.4 cm, base cuneate to narrowly cuneate, apex acute to rounded, chartaceous, costa prominulous and rounded abaxially, shallowly canaliculate adaxially, both surfaces bearing spreading subulate scales as in stipe, margins thin, flat, translucent, bearing fringe of reddish brown, spreading scales as on costa, veins distinct, unbranched or bifurcate, free apically, tips thickened and dark, both surfaces of blade with scattered subulate to deltoid scales 0.2– 0.5 mm long, glandular-ciliate at base. Fertile frond about equaling or slightly shorter than sterile frond; stipe slender, 2 to 3 times longer than sterile stipe; blade 3–7 3 0.7–1.4 cm, narrowly ovate to narrowly elliptic, base cuneate to truncate or rarely subcordate, apex acute, costa sparsely and minutely paleaceous on both surfaces, sporangia lacking on margins and extreme base of blade. Spores 34– 38 3 22–25 mm (excluding perispore), perispore surface echinate, spinules 2.5–4 mm high. any other species in the genus (Rouhan et al., 2004). Thus, E. aubertii may possibly be conspecific with E. eximium (in which case E. aubertii would have priority). Distribution and habitat. Réunion; also known from Madagascar, Comoro Islands, tropical and southern Africa, Fernando Po, and São Tomé. On Réunion this species occurs in wet forests and ericoid shrubland from 200 to 1500 m elevation. The occurrence of E. aubertii on Mauritius is doubtful and based on a single specimen (Boivin s.n.), the type of A. boivinii Mett. ex Kuhn (Filic. Afr. 43. 1868), which has not been located. No other collections are known from Mauritius. Discussion. Elaphoglossum aubertii can be distinguished by spreading, subulate scales, conspicuous hydathodes, and short petioles (⅛ to ¼ the length of the blades). Elaphoglossum aubertii greatly resembles E. eximium (Mett.) H. Christ, a species of South and Central America (Moran & Smith, 2001). Both species show morphological characters so similar that distinction is very difficult without knowing the geographical origin. This close relationship is strongly supported by phylogenetic analyses based on molecular data, since E. aubertii and E. eximium appear more closely related to each other than to Selected specimens examined. RÉUNION. Bébour, Lorence in MAU 15629 (MAU); Plateau de Bébour, Bosser 21503 (P); Dimitile, Rouhan & Grangaud 206 (P); Forêt du Tévelave, Cadet 123 (P); Montée de la Plaine des Cafres, Bosser 9544 (P); Piton, Mare à Boue, Plaine des Cafres, Bosser 21759 (P); Plaine des Fougères, Rouhan et al. 226 (NY, P, PTBG); Saint-Philippe, Basse Vallée, Badré 1004 (P); St. Paul, sentier de l’ı̂let Alcide, Rouhan et al. 241 (NY, P). 5. Elaphoglossum 3cadetii Lorence [Elaphoglossum acrostichoides (Hook. & Grev.) Schelpe 3 E. coursii Tardieu], Fern Gaz. 12: 346, fig. 3. 1984. TYPE: Réunion. Piton de la Fournaise, Plaine des Sables, T. Cadet 2063 (holotype, REU!; isotype, P not seen). Fronds spaced 2–4 mm distant, borne in 2 ranks; rhizome short-creeping, 2–4 mm diam., rarely branching, densely paleaceous; scales 2–4 3 0.5–1 mm, ovate, shiny black, base cordate or auriculate, pale brown, apex acute to acuminate, black and opaque, margins bearing rare glandular-tipped cilia. Phyllopodia to 5–7.5 mm long, swollen, dark brown to black, paleaceous toward base; sterile fronds (6– )8–14 cm; stipe (2.2–)3.6–6.5 cm 3 0.5–1 mm, canaliculate, stramineous, bearing scattered dark brown, discolorous ovate scales 2–3 3 0.5–1 mm, margins sparsely ciliate; blade narrowly elliptic, 4–10 3 0.7– 1.5 cm, base narrowly cuneate, narrowly decurrent, apex acute with rounded tip, coriaceous, costa prominulous, rounded abaxially, slightly canaliculate adaxially, margins revolute, veins indistinct, dichotomous, apically free with thickened tips, adaxial surface of blade glabrate, abaxial surface bearing minute brown, substellate scales 0.5 mm diam., with a few larger scales along costa. Fertile frond 14–18.5 cm long; stipe 7.5–14 cm, about three times longer than sterile stipe; blade 6–8.5 3 0.9–1 cm, narrowly elliptic, base narrowly cuneate, slightly decurrent, apex obtuse, costa stramineous, veins indistinct. Spores ca. 90% aborted. Distribution and habitat. Réunion, known only from the type collection. Elaphoglossum 3cadetii represents a natural hybrid between E. acrostichoides and E. coursii, which both occur at the type locality on the Piton de la Fournaise massif, an active volcano (Lorence, 1984). The putative hybrid was collected from fissures in lava at 2300 m among heath vegetation. A morphologically similar collection from Mt. Ankaroka, Madagascar (H. Humbert & G. Cours Volume 91, Number 4 2004 Lorence & Rouhan Mascarene Elaphoglossum 17521, P!), may represent this hybrid, as both putative parents also occur in Madagascar. Elaphoglossum 3cadetii resembles E. coursii but differs by its longer, narrower rhizome scales, shorter coriaceous sterile fronds with indistinct veins, and fertile fronds up to twice as long as the sterile ones. Citrons, Lorence et al. 7561 (MAU, PTBG); Grande Jument, sentier, Rouhan & Grangaud 199 (P); Haut de Saint Benoit, cratère, Bosser 21012 (P). 6. Elaphoglossum coursii Tardieu, Notul. Syst. 15: 434. 1959. TYPE: Madagascar. 1951, H. Humbert & R. Capuron 24918 (holotype, P!; isotype, K!). Fronds moderately spaced 0.5–1 cm distant, borne in 2 ranks; rhizome moderately long-creeping, 1–3 mm diam., occasionally branching, sparsely paleaceous; scales 1–1.5 3 0.4–0.7 mm, ovate to ovate-elliptic, dark brown to black, paler at base and margins, opaque, rigid, base auriculate with overlapping lobes, apex acute, margins with bulbous-tipped cilia especially basally, cells square to fusiform. Phyllopodia 5–10 mm long, slightly swollen, brown to blackish, paleaceous; sterile fronds 20–35 cm; stipe 6–15 cm, slender, canaliculate, stramineous to dark brown, bearing scales like those of rhizome but larger and paler; blade linearelliptic to linear, 10–24 3 0.6–1.8 cm, base narrowly acute, attenuate and decurrent, apex acute to long-acuminate, subcoriaceous to coriaceous, costa prominulous and rounded to angulate abaxially, canaliculate adaxially, margins strongly revolute, veins distinct to somewhat indistinct, often bifurcate, apically free, adaxial surface of blade glabrous, abaxial surface of blade bearing scattered, small brown, substellate scales 0.2–1 mm diam. with glandular marginal cilia. Fertile frond 20–35 cm long, equaling or slightly shorter than sterile frond; stipe slender, slightly longer than sterile stipe; blade linear-elliptic to linear-obovate, base narrowly cuneate, decurrent, apex acute, mucronulate. Spores 40–45 3 30–35 mm (excluding perispore), perispore cristate, crests 4–8 mm high, rounded, margins entire to finely sinuate. Distribution and habitat. Réunion; also known from Madagascar, Comoro Islands, and Zambia. On Réunion E. coursii is known from 400 to 2000 m and is epiphtyic or saxicolous in open wet forests and ericoid formations. Discussion. Elaphoglossum coursii greatly resembles E. acrostichoides, but it can be distinguished by its darker, lustrous, and stiffer rhizome scales. Specimens examined. RÉUNION. Cilaos, au dessus des Thermes, Cadet 4126 (REU); Dimitile, SE rim of Cirque de Cilaos caldera, N of Entre-Deux & Ravine des 545 7. Elaphoglossum heterolepis (Fée) T. Moore, Index Fil.: 10. 1857. Acrostichum heterolepis Fée, Mém. Foug., 2: 56, tab. 15(1). 1844 [1845], non Acrostichum heterolepis Baker, Syn. Fil. 521. 1874. TYPE: Mauritius. Sieber Flora Mixta 281 (lectotype, designated by Rouhan & Lorence (2003: 838), PR!). Figure 2A, C. Fronds caespitose, spaced 4–10 mm, borne in 2 ranks; rhizome short-creeping, 4–5 mm diam., profusely branched, densely paleaceous; scales 6–12 3 0.5–0.7 mm, narrowly ovate, shiny dark brown to black with clear or white borders, spreading, persistent, base cordate to auriculate, apex slender, acute, margins entire, central cells fusiform, opaque (Fig. 5A). Phyllopodia 3–6 cm long, chocolate to blackish brown, bearing at base scales as in rhizome; sterile fronds 22–40(–55) cm; stipe 7– 14(–16) cm, 1.5–2.5 mm diam., reddish to orangebrown, canaliculate, winged in distal half, densely paleaceous with reddish brown to black, narrowly ovate, often falcate scales 3–6 3 0.5–1 mm, base truncate to auriculate and bearing a few hair teeth, apex long-acuminate, margins entire, scarious, whitish to yellowish; blade linear-ovate to linearelliptic or linear-obovate, 20–40(–50) 3 2.6–3.6(– 4) cm, base narrowly cuneate, attenuate, apex longacuminate to caudate, chartaceous, costa prominulous and rounded abaxially, prominulous and shallowly canaliculate adaxially, bearing basally scales as in stipe, margins rounded, slightly revolute, veins unbranched or bifurcate, apically free, tips thickened, both surfaces of young blade covered by a thin continuous layer of appressed, beige to gray scales of two types, the small ones 0.5 mm diam., ovate to orbicular, peltate, scarious, sometimes with brown centers, margins densely ciliate-fimbriate with hair teeth (Fig. 5C), the larger scales often concentrated along the costa, 1.5–3 3 0.5 mm, narrowly ovate, sometimes with brown centers, base flabelliform, margins densely fimbriateciliate, the cells thin-walled, translucent (Fig. 5B), adaxial blade surface glabrescent. Fertile frond 34– 50(–62) cm long; stipe 1.5 times longer than sterile stipe; blade 14–25(–28) 3 1.4–2.5(–2.8) cm, linear-elliptic, base narrowly cuneate to attenuate, apex acute to obtuse, adaxial surface blade and abaxial surface stipe bearing dense layer of rusty-colored scales as in sterile frond, intersporangial scales very often present. Spores 43–45 3 23–30 mm (excluding perispore), perispore cristate, 5–10 546 Annals of the Missouri Botanical Garden Figure 2. Habit of some Mascarene Elaphoglossum species. Except for E. heterolepis, all the illustrations show specimens with sterile and fertile fronds. —A, C. E. heterolepis with its bi-colored stipe scales and both surfaces of young blade covered by a thin continuous layer of appressed, beige to gray scales, from Mauritius, same population as Rouhan 177. —B. Typical longer fertile fronds of E. lepervanchei from Mauritius, same population of Rouhan et al. 175. —D. E. macropodium from Réunion, same population as Rouhan et al. 232. —E. E. tomentosum from Mauritius, same population as Rouhan et al. 180. Scale bars 5 2 cm. Volume 91, Number 4 2004 Lorence & Rouhan Mascarene Elaphoglossum mm high with finely crisped margins and spinulose surface. Route du Grand Etang, Badré 896 (P); Salazie, 1886, Keller s.n. (P); Takamaka, Cadet 1702 (P); Tremblet, Rouhan & Grangaud 211 (NY, P). Distribution and habitat. Réunion and Mauritius. Epiphytic to 10 m above ground or occasionally terrestrial, often forming dense clumps, rarely saxicolous. Common on both islands from 200 to 1400 m elevation in moist and wet forests. Discussion. This species resembles Elaphoglossum tomentosum by the blade covered by a thin continuous layer of appressed scales. Nevertheless, E. heterolepis differs clearly from all other Mascarenes species by its apically free veins, two types of frond scales, and slender, elongated bicolorous rhizome scales with dark centers and whitish borders. On Mauritius it hybridizes with E. tomentosum (Lorence, 1984). Since Elaphoglossum heterolepis is similar to E. tomentosum, Baker did not distinguish both species in his book Synopsis Filicum (1874). Therefore, in his treatment the name Acrostichum heterolepis Fée appears as a synonym of the earlier published name A. tomentosum. Consequently, A. tomentosum was misapplied and used in place of the correct name E. heterolepis in the following treatments: Acrostichum tomentosum sensu Bojer, Hortus Maurit. 413. 1837; sensu Cordem., Fl. Réunion, part 1: 85. 1891, Fl. Réunion. 96. 1895, non Acrostichum tomentosum Bory ex Willd. Elaphoglossum heterolepis is also the correct name for plants misidentified by Tardieu as E. obductum (Kaulf.) T. Moore and figured in her treatment in Notul. Syst. 15(4): 429, tab. 3(6–11). 1959. Typification of the name E. heterolepis is discussed in greater detail by Rouhan and Lorence (2003). Selected specimens examined. MAURITIUS. Alexandra Falls, Rouhan et al. 177 (MAU, NY, P); Bel Ombre, Lorence 1402 (MO); Kanaka Crater, Lorence 1078 (MO); Macabé forest, Lorence 1420 in MAU 17544 (K, MAU, MO); Macabé-Brise Fer rd., Lorence in MAU 16019 (K, MAU, P); Macabé, sentier menant à la Mare aux Joncs, Rouhan et al. 179 (MAU, NY, P, PTBG); Mare Longue Plateau, Lorence 2.11 (MO); Mt. Lagrave, E flank, Lorence 783 in MAU 16317 (K, MAU); flank facing Eau Bleu reservoir, Lorence in MAU 18785 (MAU); Pétrin Nat. Reserve, rd. to Grand Bassin, Lorence in MAU 15816 (MAU, MO, REU); Piton Grand Bassin, Lorence in MAU 14599 (MAU); Piton Grand Fond, Lorence 300 (MO); Plaine Champagne, rd. to Bel Ombre, Lorence in MAU 16018 (MAU); Bassin Blanc, path from Les Mares, Lorence in MAU 14609 (MAU). RÉUNION. Basse Vallée (St. Philippe), Cadet 3790 (P, REU); Bébour, path to Piton Bébour, Lorence in MAU 15645 (MAU); Boucan Launay, Boivin 800 (P); Brûlé de Sainte Rose, Badré 985 (P [2]); Brûlé, Bédier 53 (P); Chemin Arnoux, Cadet 880 (P); Cilaos, Cadet 1930bis (P); Dimitile, Rouhan & Grangaud 202 (P); Mare Longue–St. Philippe, Schlieben 10934 (MAU); Morne des Patates Durand, Lorence in MAU 15619 (MAU); Plaine des Fougères, 1850, Boivin s.n. (P); 547 8. Elaphoglossum 3heterophlebium Lorence [Elaphoglossum heterolepis (Fée) T. Moore 3 E. tomentosum (Bory ex Willd.) H. Christ], Fern Gaz. 12: 343, fig. 2. 1984. TYPE: Mauritius. Pétrin Nat. Reserve, 1973, D. H. Lorence 14.2 in MAU 15821 (holotype, MAU!). Fronds caespitose, borne in 2 ranks; rhizome short-creeping, 5 mm diam., occasionally branched, densely paleaceous; scales 5–7 3 0.5–0.7 mm, narrowly ovate to lanceolate, slightly falcate, castaneous, thick, base cordate, apex long-acute, filiform, margins scarious, bearing sparse short hair teeth or rare glandular cilia, cells opaque. Phyllopodia 1–2 cm long, dull brown, paleaceous as in rhizome; sterile fronds 33–45 cm; stipe 8–13 cm, 1.5–2 mm diam., stramineous, adaxially flattened, densely paleaceous when young, scales narrowly ovate, falcate, 2–5 3 0.8–1.5 mm, base peltate and obtuse, or basifixed and cordate or auriculate, apex acute, centers dark castaneous, opaque, margins scarious pale brown, fimbriate to ciliate-dentate with hair or glandular-tipped teeth; blade oblongelliptic or lanceolate, 22.5–32 3 2.6–3.4 cm, base narrowly cuneate, apex acute to slightly acuminate, subcoriaceous, costa prominulous on both surfaces, rounded abaxially, slightly canaliculate adaxially, margins rounded, slightly revolute, veins bifurcate, in part free with thickened tips, in part anastomosing into intramarginal arcs, both surfaces of blade covered by a thin, persistent homogeneous layer of closely appressed, matted, ovate to subcircular, peltate scales 0.5–1 3 0.5 mm, buff-colored, hyaline, base and margins fimbriate with arachnoid cilia 0.1–0.3 mm long with acicular or bulbous tips, cells thin-walled. Fertile frond 32.5–36 cm long; stipe 16.5–19.5 cm long; blade 16–16.5 3 1–1.5 cm, linear-elliptic, base and apex acute, adaxial blade surface bearing thin layer of scales as in sterile frond. Normal spores 35–38 3 25–27 mm (excluding perispore), perispore cristate, 2–3 mm high with minutely spinulose margins; 52%–85% of spores aborted. Distribution and habitat. Mauritius, known from four scattered localities on the central plateau at 600 to 700 m in wet forests and Philippia Klotzsch heath formations. In all cases, the nothospecies was growing terrestrially close to one or both parental species, E. heterolepis and E. tomentosum. Discussion. Specimens of Elaphoglossum 3heterophlebium are remarkably uniform morphologi- 548 Annals of the Missouri Botanical Garden cally. Although tending to resemble E. heterolepis somewhat more than E. tomentosum, E. 3heterophlebium is intermediate between both parents regarding color and shape of the scales and the veins, which are in part free with thickened tips (resembling E. heterolepis) and in part uniting into intramarginal arcs (resembling E. tomentosum). basally, black; sterile fronds 25–30(–55) cm; stipe 7–12(–20) cm, 1–2 mm diam., stramineous to pale brown, slightly canaliculate, bearing a dense layer of squarrose, persistent, subulate, dark brown scales 2–5 3 0.3–0.5 mm, base cochleariform and closed, apex long-acuminate, filiform, margins involute, entire, stipe also with simple capitate and irregularly branched hairs; blade ovate to narrowly ovate, elliptic, or sometimes obovate-oblong, 15– 20(–40) 3 3.5–4.5(–6) cm, base narrowly to broadly cuneate or sometimes rounded, apex acute to acuminate, chartaceous to coriaceous, costa prominulous and rounded abaxially and bearing scattered subulate scales as in stipe, shallowly canaliculate and glabrous adaxially, margins revolute, bearing fringe of dark brown, spreading, persistent, subulate scales as in stipe, veins prominulous, unbranched or 1 to 2 times dichotomous, apically free, tips thickened, both surfaces of blade bearing short, numerous, simple capitate and branched hairs, adaxially glabrescent. Fertile frond shorter than or nearly equaling sterile frond, 16–26(–33) cm long; stipe slender, about equaling sterile stipe; blade 7–12 3 1.5–2.5 cm, narrowly ovate to narrowly elliptic, base acute to narrowly cuneate, apex acute, adaxial surface with short capitate hairs. Spores 36–38 3 23–25 mm (excluding perispore), perispore cristate, crests 3–7 mm high, margins sinuate, somewhat erose. Specimens examined. MAURITIUS. Brise Fer rd., Lorence 16.2 in MAU 15819 (MAU); Plaine Champagne, Lorence sub MAU 15099 (MAU), Lorence 996 sub MAU 16518 (MAU), Lorence 1544 (MAU, MO); Pétrin Nat. Reserve, Lorence 14.2 in MAU 15821 (MAU). 9. Elaphoglossum hybridum (Bory) Brack., U.S. Expl. Exped., Filic. 16: 69. 1854. Acrostichum hybridum Bory, Voy. Iles Afrique. 3: 95. 1804, non Elaphoglossum hybridum (Bory) T. Moore, Index Fil.: 10. 1857. Olfersia hybrida (Bory) C. Presl, Tent. Pterid. 235. 1836. TYPE: Bourbon [Réunion]. Caverne le Gentil, Bory de St. Vincent s.n. [Hb. Bory 27,7] (holotype, P!). Figure 4A. Although the specific epithet hybridum means ‘‘of hybrid origin,’’ its application to this taxon is not clear. The author of the basionym, Bory, does not explain why he chose this specific epithet. Furthermore, no evidence supports a hybrid origin for this species, as the morphology is not intermediate between other species in the region, and the spores are not aborted. The names E. hybridum var. cuneatum Bonap., E. hybridum var. melanopus Farw., and E. hybridum var. nitidum (Liebm.) H. Christ have been applied to plants outside the Mascarene area and are not considered herein. 9a. Elaphoglossum hybridum (Bory) Brack. var. hybridum Acrostichum ciliatum Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 310. 1811, non Acrostichum ciliatum C. Presl, Reliq. Haenk. 1: 15. 1825. TYPE: Bourbon [Réunion]. P. Commerson s.n. (lectotype, designated by E. Schelpe (1969: 32), P-JU 1000A!). Elaphoglossum tricholepis (Baker) C. Chr., Index. Filic.: 317. 1905. Acrostichum tricholepis Baker, J. Bot. 1891: 5. 1891. TYPE: Madagascar. Bekilus Mtns., J. T. Last s.n. (holotype, K!). Fronds densely caespitose, borne in 4 ranks; rhizome very short-creeping, 10–12 mm diam., unbranched, densely paleaceous; scales 5–10 3 1–3 mm, narrowly to oblong-ovate, brown, scarious, base truncate to cordate, apex long-acuminate, margins entire or with a few short teeth, base with several short bulbous-tipped cilia, cells squarish to linear-oblong. Phyllopodia 8–18 mm long, swollen Distribution and habitat. Mauritius (400–900 m) and Réunion (600–2200 m); also Madagascar, Comoro Islands, tropical and southern continental Africa, Fernando Po, Tristan d’Acunha, and Gough Island. In the Mascarenes usually found growing on mossy rocks and boulders, rarely terrestrially, in shade of montane wet and cloud forests. Discussion. Elaphoglossum hybridum var. hybridum can be distinguished by blades 3.5–4.5 cm wide, absence of hydathodes, and blades with both surfaces and margins bearing dark brown, spreading, subulate scales. Elaphoglossum hybridum var. hybridum greatly resembles E. erinaceum (Fée) T. Moore from the Neotropics (Moran & Smith, 2001). Phylogenetic analyses based on molecular data are inconclusive concerning the relationship between both species (Rouhan et al., 2004). Although both species show morphological characters so similar that distinction is very difficult without knowing the geographical origin, evidence is inconclusive if E. hybridum may really be conspecific with E. erinaceum (in which case, E. hybridum would have priority). Selected specimens examined. MAURITIUS. Bassin Blanc crater, Julien in MAU 16494 (MAU); Corps de Gar- Volume 91, Number 4 2004 Lorence & Rouhan Mascarene Elaphoglossum de Mtn., Guého in MAU 11498 (MAU); Mont Deux Mamelles, Lorence in MAU 15024 (K, MAU, P, REU); Plaine Champagne–Chamarel rd., headwaters of Rivière Saint Denis, Julien in MAU 14595 (MAU); Rivier du Poste near Kanaka, Staub in MAU 11348 (MAU); Ruisseau Commérage, Rouhan et al. 195 (P); Tamarind Falls reservoir, Lorence in MAU 15823a (MAU); valley of Cascade 500 Pieds, Lorence 1608 (MO). RÉUNION. Bébour, Rivière des Marsouins, Lorence in MAU 15625A–D (MAU); sentier de la Rivière, Rouhan & Grangaud 250 (P); Cilaos, sentier GR R1 entre ‘‘Le Bloc’’ et le gı̂te de la Caverne Dufour, Rouhan 217 (NY, P, PTBG); sentier allant à la cascade de Bras Rouge, Rouhan 239 (P); Dimitile, sentier de la Grande Jument, Rouhan & Grangaud 198 (P); Grand Matarum, Lorence in MAU 15626 (MAU); Morne des Patates Durand, Lorence in MAU 15622 (MAU); Piton des Neiges path, Lorence in MAU 15635 (MAU); Plaine des Fougères, Rouhan et al. 228 (NY, P); Saint-Philippe, Basse Vallée, Badré 1005 (P); Takamaka, trail to Ilet à Bananes, Lorence & Rolland 6944 (PTBG); volcan, Enclos Fouqué, Cadet 1556 (isolated fertile frond, P); Crête du Volcan, Bosser 11857 (P). Rivière des Marsouins, Lorence in MAU 15625A– D, MAU). 9b. Elaphoglossum hybridum var. vulcani (Lepervanche ex Fée) H. Christ, Neue Denkschr. Allg. Schweiz. Ges. Gesammten Naturwiss. 36(1): 104. 1899. Acrostichum hybridum var. vulcani Lepervanche ex Fée, Mém. Foug., 2: 41, tab. 9(3). 1844 [1845]. TYPE: Bourbon [Réunion]. Plaine des Palmistes, Plaine des Cafres, Jun. 1831, P. Lépervanche-Mézières s.n. [Hb. Bory 27,7] (holotype, P!). Sterile fronds 8–12(–21) cm long; stipe 3–6(–13) cm long, 0.5 mm diam., somewhat flexuose, densely paleaceous with small scales 2–3.5 3 0.5 mm with large cells, short brown, capitate hairs also present; blade ovate to ovate-elliptic, 4–6(–11) 3 1–2.5(– 3.6) cm, surfaces scattered with abundant, short, pale brown, capitate, simple or branched hairs, margins fringed with squarrose subulate scales 1.5– 2 mm long. Fertile frond 4 3 1.3–1.5 cm, blade with sporangia often absent from margins and extreme base of blade. Distribution and habitat. Réunion; also Madagascar and the Comoro Islands. This diminutive variety is restricted to higher elevations (1300–2300 m) than variety hybridum. It is usually found growing on shady, mossy rock faces and in grottos. Discussion. Plants of variety vulcani differ from those of variety hybridum in their smaller habit and fronds, in the smaller scales, and in the fertile frond with sporangia often absent from margins and the extreme base of the blade. Elaphoglossum hybridum var. vulcani may, however, represent a depauperate ecotype, since collections intermediate between variety hybridum and variety vulcani occur on Réunion (e.g., see under var. hybridum: Bébour, 549 Selected specimens examined. RÉUNION. Cilaos, sentier GR R1, entre ‘‘Le Bloc’’ et le gı̂te de la Caverne Dufour, Rouhan 214 (NY, P); Cratère Commerson, Rouhan & Grangaud 219 (P, PTBG); Pas de Bellecombe, Rouhan 222 (NY, P); Pas des Sables, Sep. 1892, Cordemoy s.n. (P); path to Piton des Neiges, near le gı̂te, Lorence in MAU 15635 (MO); Plateau Kerveguen, Badré 960 (P); Route forestière de Tévelave, ligne Domaniale, Lorence 2496 (MAU); Sommet du Grand Bénard, 16 Feb. 1847, Boivin s.n. (P). 10. Elaphoglossum lanatum (Bojer ex Baker) Lorence, Fern Gaz. 11: 199, fig. 1a–g. 1976. Acrostichum viscosum Sw. var. lanatum Bojer ex Baker, Fl. Mauritius: 512. 1877. TYPE: Maurice [Mauritius]. Bois Chéri, W. Bojer s.n. pro parte, 4 left-hand specimens only (holotype, K! [Hb. Hook.]). Figure 3B. Fronds spaced 3–5 mm, borne in 2 ranks; rhizome short-creeping, 2–3 mm diam., rarely branched, resinous, densely paleaceous; scales 1.2–2.5 3 0.3–0.5 mm, narrowly ovate, castaneous, shiny, stiff, paler brown basally, base cordate to auriculate, apex narrowly acuminate, filiform, margins sinuate to shortly dentate bearing short, glandular teeth, cells square to rectangular, arranged in longitudinal rows (Fig. 6A). Phyllopodia 3–5 mm long, swollen, dark brown, resinous, densely paleaceous as in rhizome; sterile fronds (6–)12–30(–42) cm; stipe (2–)4–12(–15) cm, 1–1.5 mm diam., stramineous, slightly canaliculate, often narrowly winged distally, when young bearing densely matted, pale rusty brown, scarious, lanceolate to deltoid scales 1–2 3 0.2–0.4 mm, base truncate to auriculate, the lobes prolonged into long, often bulbous-tipped cilia, apex long-acuminate, filiform, margins with long, spreading, usually paired cilia, cells rectangular; blade pendulous, narrowly to linear-elliptic, (4–)8–20(–30) 3 1–2(–2.5) cm, base narrowly cuneate to long-attenuate, decurrent, apex acute to obtuse or rounded, subcoriaceous to coriaceous, costa rounded on both surfaces, stramineous, margins rounded, revolute, pale, veins indistinct, bifurcate, in part apically free, in part uniting into an intramarginal commissure, both surfaces of blade at first densely covered with arachnoid-tomentose layer of matted, pale reddish brown to buff-colored, deltate to rounded, hyaline scales 0.5–1 mm diam. (cilia included), base auriculate to sagittate, each lobe fringed with 3 to 5 long, spreading or crinkled, arachnoid hair teeth, apex filiform, margins bearing 1 to 2 pairs of long hair teeth and shorter teeth, cells toward base squarish to amoeboid, those to- 550 Annals of the Missouri Botanical Garden Figure 3. Habit of some Mascarene Elaphoglossum species. —A. E. splendens on mossy boulders from Réunion, same population as Rouhan 218. —B. Fronds of E. lanatum: some of them are densely covered with arachnoidtomentose layer of matted, pale scales, becoming glabrescent, from Mauritius, Rouhan et al. 192. —C. E. rufidulum with fertile fronds, from Réunion, Rouhan & Grangaud 224. —D. E. stipitatum on rocks with fertile fronds, from Réunion, same population as Rouhan 212. Scale bars 5 2 cm. Volume 91, Number 4 2004 Lorence & Rouhan Mascarene Elaphoglossum ward apex rectangular (Fig. 6B), adaxial blade surface glabrescent. Fertile frond (3–)8–14(–20) cm, slightly shorter than sterile frond; stipe slender, (2–)4–8(–11) cm; blade 5–7(–8.5) 3 1–1.2(–1.4) cm, chartaceous, shape and scales of adaxial surface as in sterile frond. Spores 35–38 3 23–28 mm (excluding perispore), perispore cristate, crests 4– 8 mm high, margins finely erose-dentate. Pouce Mtn., near summit, Lorence 21.1 in MAU 16280 (MAU); Mare Longue Plateau, Lorence in MAU 15243 (K, MAU, P, REU); Perrier Nat. Reserve, Lorence 17.2 in MAU 15830 (MAU, MO); Bord de la rivière Cogliano, Rouhan et al. 194 (MAU, NY, P [2], PTBG); Tamarind Falls Reservoir, Lorence in MAU 15823 (MAU). REUNION? [Doubtful]. Without precise locality, Bojer s.n. (pro parte, left-hand and center collections, K). Distribution and habitat. Endemic to Mauritius, where it is restricted to nearly vertical river banks and cliff faces, often in sheltered overhangs growing on soil or decomposing lava rock at 300 to 800 m. There is a Bojer s.n. sheet housed in K (not the sheet of the type) comprising three different specimens: the right-hand specimen is E. lancifolium and occurs in Réunion and Mauritius, but the two remaining specimens are E. lanatum, which is endemic to Mauritius. Consequently, the locality of Bourbon [Réunion] mentioned on the label of this sheet is probably erroneous or collections were mixed between both islands, because no other collections of E. lanatum are known from Réunion. Discussion. The holotype of Acrostichum viscosum var. lanatum, W. Bojer s.n pro parte, includes 4 left-hand specimens only; the 2 remaining righthand specimens are fertile fronds of E. lancifolium (Desv.) C. V. Morton (Lorence, 1976a: 199). Elaphoglossum viscosum (Sw.) J. Sm. is a species restricted to the Neotropics. Extremely variable are frond size and shape, the degree to which the veins are free or unite intramarginally, and the percentage of aborted spores. This variability suggests a possible hybrid origin (Lorence, 1976a, 1984). Although extremely variable, E. lanatum can be distinguished from all others in the Mascarenes by the blades with veins in part apically free, in part uniting into an intramarginal commissure, and by blades bearing an arachnoid-tomentose layer of matted, pale reddish brown, deltate to rounded, hyaline scales. Furthermore, it is unique in being the only Mauritian Elaphoglossum that is rupicolous and grows exclusively on vertical river banks and cliff faces composed of decaying lava. In the Mascarenes, Elaphoglossum lanatum hybridizes with E. hybridum forming E. 3setaceum, and with E. tomentosum resulting in E. 3adulterinum (Lorence, 1984). Selected specimens examined. MAURITIUS. Bel Ombre, Lorence 1406 (MAU, MO); Black River Gorge, N edge behind Plaine Lièvre, Lorence in MAU 15533 (MAU, MO, REU); Cascade 500 Pieds near Plaine Champagne, Lorence 1173 (KRA, MAU, MO); Florin, le long de la Mare aux Riz, Rouhan et al. 192 (MAU, NY, P, PTBG); Le 551 11. Elaphoglossum lancifolium (Desv.) C. V. Morton, Contr. U.S. Natl. Herb. 38: 32. 1967. Acrostichum lancifolium Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 310. 1811. TYPE: Mascareignes. 1801, A. Du Petit-Thouars s.n. (holotype, P-JU 1004!). Figure 4B. Acrostichum salicifolium Willd. ex Kaulf., Enum. Filic. 58. 1824. Elaphoglossum petiolatum (Sw.) Urb. var. salicifolium (Willd. ex Kaulf.) C. Chr., Trans. Linn. Soc. London, Bot. 7: 421. 1912. Elaphoglossum salicifolium (Willd. ex Kaulf.) Alston, in Exell, Cat. Vasc. Pl. San Tomé 92. 1944. Elaphoglossum petiolatum (Sw.) Urb. subsp. salicifolium (Willd. ex Kaulf.) Schelpe, Contr. Bolus Herb. 1: 34. 1969. TYPE: Bourbon [Réunion]. R. Desfontaines s.n. pro parte, right-hand specimen only (holotype, B-W 19509!). Fronds spaced 4–5 mm distant, borne in 2 ranks; rhizome short-creeping, 3–4 mm diam., sparsely branched, densely paleaceous, resinous; scales 3– 5 3 0.5–0.8 mm, narrowly to linear-ovate, dark reddish brown, shiny, base truncate to cordate or auriculate, apex long-acuminate, filiform, margins entire except with a few short cilia and glands basally, cells fusiform. Phyllopodia 5–7 mm long, slightly swollen, blackish brown; sterile fronds 25– 40(–44) cm; stipe 4–10(–23) cm, 1 mm diam., stramineous to reddish brown, slightly canaliculate, with scattered, substellate to elongate scales to 0.5– 3.5 3 0.5–1 mm, margins with long hair teeth, stipe also with brown resinous punctations; blade linearelliptic to linear, 20–30(–35) 3 1–2(–2.5) cm, base narrowly cuneate to cuneate, sometimes decurrent, apex acute to long-acuminate, chartaceous to subcoriaceous, costa prominulous and rounded abaxially, canaliculate adaxially, margins revolute, veins prominulous, unbranched or bifurcate, apically free, tips thickened, dark, both surfaces of blade when young with sparse, scattered stellate scales 0.5–1 mm diam., pale brown, margins with usually 6 to 8 long hair teeth, scales denser along costa, blade surfaces also with scattered light brown, resinous punctations. Fertile frond equaling or slightly longer than sterile frond, 20–35(–40) cm; stipe slender, equaling to twice as long as sterile stipe; blade 14–26 3 0.7–1 cm, linear or linear-elliptic, base cuneate, apex acute, adaxial surface with scattered stellate scales. Spores 35–38 3 25–28 mm 552 Annals of the Missouri Botanical Garden Volume 91, Number 4 2004 Lorence & Rouhan Mascarene Elaphoglossum (excluding perispore), perispore cristate, crests 3– 5 mm high, margins entire to sinuate. never published by Kaulfuss himself), which is different from Acrostichum (Elaphoglossum) obductum sensu Kaulfuss ex Spreng., because according to the Code the latter name is illegitimate and must now refer to Elaphoglossum lancifolium. The following misapplied name found in the early Mascarene literature should refer to E. lancifolium: Acrostichum viscosum sensu Bojer, Hortus Maurit. 412. 1837, sensu Baker, Fl. Mauritius 512. 1877, sensu Cordem., Fl. Réunion, part 1: 83. 1891, Fl. Réunion 94. 1895, non Acrostichum viscosum Sw., Syn. 10: 193. 1806. Elaphoglossum viscosum (Sw.) J. Sm. is a Neotropical species. Distribution and habitat. Mauritius (300–800 m) and Réunion (100–1300 m); also Madagascar, Comoro Islands, Seychelles, San Tomé, and tropical Africa. Occurring in moist to wet forests in light to dense shade, epiphytic to 10 m above ground or growing on fallen trunks and mossy boulders. Discussion. Elaphoglossum lancifolium resembles E. richardii by the blade shape but differs by its densely caespitose fronds and blade with free veins. Elaphoglossum lancifolium can be distinguished from all others by scales that are stellate with usually 6 to 8 long, straight hair teeth. The type of Acrostichum salicifolium, R. Desfontaines s.n., includes the right-hand specimen only. The remaining left-hand fertile specimen, an isolated fertile frond without rhizome, is of uncertain identity. Acrostichum lancifolium Desv. is the oldest available name for the species (Morton, 1967: 32–33). Morton (1973: 215–216) further stated that A. obductum Kaulf. ex Spreng. (Syst. Veg. (ed. 16): 34. 1827) is a superfluous name based on the same type as A. lancifolium Desv.: ‘‘It is likely that Kaulfuss intended his A. obductum to be based on Mauritius, Sieber, Syn. Fil. no. 25 [referring now to E. tomentosum (Bory ex Willd.) H. Christ], and it has been perhaps generally so considered, although never definitely lectotypified. But when Sprengel published the name he did not mention Sieber or indeed cite any specimens, merely citing [erroneously] Acrostichum lancifolium Desv. as a synonym. Since the name A. lancifolium was a legitimate and available name, there was no need to propose a different name, A. obductum . . . and so by Art. 63 [modern equivalent in the ICBN, Greuter et al., 2000: Art. 52] of the Code the name A. obductum must be considered superfluous, and by Art. 7 [7.5] it must be typified on the basis of the name that ought to have been adopted, thus on the type of A. lancifolium.’’ Consequently, the name Acrostcihum obductum or its combination Elaphoglossum obductum (Kaulf. ex Spreng.) T. Moore (Index Fil.: 124. 1857), has been widely misapplied. Elaphoglossum tomentosum is indeed the correct name for the plant identified by almost all authors as Acrostichum (Elaphoglossum) obductum sensu Kaulfuss (even if 553 Selected specimens examined. MAURITIUS. Bassin Blanc, Lorence 506 (MO); Bel Ombre, N of Bon Courage forest station, Lorence & Florens 7017 (MAU, PTBG); Bois Sec near Grand Bassin (date?), Lorence s.n. (MO); Curepipe, Kanaka, Bijoux 69 in MAU 2570 (MAU); Les Mares, Rochecouste & Orian s.n. (MAU); Macabé, Réserve de Brise Fer, Rouhan et al. 187 (MAU, P); Macabé, sentier menant à la Mare aux Joncs, Rouhan et al. 181 (MAU, P); Mare Longue Plateau, Mare Longue–Macabé rd., Lorence 7.3 in MAU 15813 (K, MAU, M, MO, P, REU); Mt. Lagrave, E flank, Lorence 792 in MAU 16315 (MAU); Perrier Nat. Reserve, Rouhan et al. 193 (MAU, P); Pétrin Nat. Reserve, Lorence 8.5 (MO); Piton du Fouge, Lorence 318 (MO); Piton du Milieu, Lorence 10.3 (K, MAU, MO, P); Piton Grand Bassin, Julien in MAU 14594 (MAU); Plaine Paul, Vaughan in MAU 12544 (MAU); Rivière Cogliano near Mare aux Vacoas, Lorence 1374 (MO); Rivière des Galets, below Cascade 500 Pieds, Lorence 1479 (KRA, M, MAU, MO). RÉUNION. Bois Blanc, Guého in MAU 15362 (MAU); Cascade de Bras Rouge, Rouhan 236 (NY, P, PTBG); Cilaos, au dessus des Thermes, Cadet 4127 (REU); Dimitile, Rouhan & Grangaud 201 (NY, P); Forêt du Grand Brûlé, Cadet 319 bis (P); Grande Jument, Rouhan & Grangaud 197 (P); Hauts des Trembles, Bosser 20624 (P); Morne des Patates Durand, Lorence in MAU 15628 (MAU); St. Philippe, forêt de Mare Longue, Guého in MAU 15303 (MAU); St. Philippe, Mare Longue, Schlieben 10928 in MAU 12416 (MAU); Tremblet, Rouhan & Grangaud 210 (NY, P, PTBG); Vallée de Langevin a Grand Galet, Cadet 3718 (REU). 12. Elaphoglossum lepervanchei (Bory ex Fée) T. Moore, Index Fil.: 11. 1857. Acrostichum lepervanchei Bory ex Fée, Mém. Foug., 2: 37, tab. 9(1). 1844 [1845]. TYPE: Réunion. S.n., [Hb. Bory 27,14] pro parte, right-hand specimen only (lectotype, designated by Lorence (1976: 200), P!). Figure 2B. Elaphoglossum didynamum (Fée) T. Moore, Index Fil.: 8. 1857. Acrostichum didynamum Fée, Mém. Foug., 2: ← Figure 4. Habit of some Mascarene Elaphoglossum species. —A. Epiphyte at the base of a trunk, E. hybridum with a fertile frond, from Réunion, same population as Rouhan et al. 228. —B. E. lancifolium, from Mauritius, Rouhan et al. 187. C, D. E. spatulatum on mossy boulders, from Réunion, Rouhan & Grangaud 246. —C. Shows the longer fertile fronds, conduplicate before maturity. —D. Sterile fronds. Scale bars 5 2 cm. 554 Annals of the Missouri Botanical Garden 37, tab. 16(2). 1844 [1845]. TYPE: Réunion. P. Lépervanche-Mézières s.n. [Hb. Bory 27, 13] (holotype, P!). Elaphoglossum curtisii (Baker) C. Chr., Index Fil.: 305. 1905. Acrostichum curtisii Baker, Ann. Bot. 5: 489. 1891. TYPE: Madagascar. 1881, C. Curtis 121 (holotype, K!; photo of holotype, P!). Elaphoglossum borbonicum (Baker) C. Chr., Index Fil.: 303. 1905. Acrostichum borbonicum Baker, Ann. Bot. 5: 491. 1891. TYPE: Bourbon [Réunion]. I. B. Balfour 27 (holotype, K!). or at bases of trees and shrubs. It is rarely found in tall, shady wet forests. Discussion. Elaphoglossum lepervanchei is extremely variable in size and shape of fronds, even on a single plant. This has resulted in a proliferation of synonyms and misapplied names in the literature. This species resembles E. macropodium by frond shape and reddish brown rhizome scales, but can be distinguished by having free veins, thinner rhizomes, and smaller rhizome scales with crisped, involute, glandular-ciliate margins. On Mauritius, E. 3revaughanii is a natural hybrid between E. lepervanchei and E. sieberi (Lorence, 1984). The specific epithet was originally published as ‘‘lepervanchii.’’ Since the species was dedicated to Mr. Lépervanche, the publication is valid according to the Code (Art. 32.6; Greuter et al., 2000), but the Latin termination contrary to Rec. 60C.1 is treated as an orthographic error to be corrected (Art. 60.11). Therefore, the correct spelling of the name is ‘‘lepervanchei.’’ In the literature concerning the Madagascan and Mascarenes area, the following misapplied names should actually refer to E. lepervanchei: Fronds spaced (2–)8–12(–25) mm, 2-ranked; rhizome moderately long-creeping, 3–5 mm diam., occasionally branched, densely paleaceous near apex; scales 4–10 3 0.5–1 mm, narrowly ovate, elliptic or linear, thin, pale yellowish to reddish brown, base truncate to auriculate, apex filiform with bulbous tip, margins involute, bearing especially toward base long, flexible gland-tipped cilia, the cells thin-walled, rectangular to fusiform, alternating with bands of shorter, squarish cells (Fig. 5D). Phyllopodia 15–25 mm long, slightly swollen, articulated, dark brown to black, paleaceous; sterile fronds 7–35(–60) cm; stipe 2–15(–30) cm 3 1–2 mm, stramineous to rusty brown, slightly canaliculate, narrowly winged in distal 0.5–0.7, sparsely paleaceous, the scales loose, pale brown 1–2 mm, margins ciliate; blade narrowly ovate-elliptic to linear-elliptic or narrowly obovate, 5–20(–26) 3 1.5– 4.5(–9) cm, base narrowly to broadly cuneate or truncate, usually attenuate and decurrent, apex obtuse to acuminate, subcoriaceous to coriaceous, costa prominulous, rounded abaxially, flattened to canaliculate adaxially, margins thin, revolute, veins distinct, prominulous, bifurcate 1 to 2 times, apically free, tips thickened, adaxial surface of blade glabrate, abaxial surface of blade with scattered, appressed dark brown scales 0.2–0.5 mm diam., irregularly stellate with bulbous-tipped cilia (Fig. 5E), scales larger and denser along costa. Fertile frond 12–35(–45) cm long; stipe 1.5 to 3 times longer than in sterile frond; blade narrowly elliptic to linear-elliptic, 6–20(–36) 3 1.2–2.6(–4) cm, flexible, apex acute to obtuse, base cuneate to attenuate, slightly decurrent, veins visible adaxially. Spores 35–38 3 23–26 mm (excluding perispore), the perispore cristate, crisped, 4–10 mm wide with erose-dentate margins. Distribution and habitat. Mauritius and Réunion; also Madagascar, Comoro Islands, and the Seychelles. This species occurs in humid localities from low to middle elevations, 200 to 900 m on Mauritius and 160 to 1300 m on Réunion. One of the most common Mascarene Elaphoglossum species, it abounds in ericoid shrubland and low, open Sideroxylon L. forests where it occurs terrestrially Acrostichum conforme sensu Bojer, Hortus Maurit. 412. 1837; sensu Baker, Fl. Mauritius 511. 1877; sensu Cordem., Fl. Réunion, part 1: 81. 1891, Fl. Réunion 93. 1895, non Acrostichum conforme Sw., Syn. Fil. (Sw.) 10, 192, tab.1(1). 1806. Acrostichum ovalifolium Bojer, Hortus Maurit. 412. 1837, nom. nud. Acrostichum latifolium sensu Baker, Fl. Mauritius 511. 1877, non Acrostichum latifolium Sw., Prodr. Pl. Ind. Occid.: 128. 1788. Elaphoglossum latifolium sensu C. Chr., Trans. Linn. Soc. London, Bot. 7: 420. 1912, non Elaphoglossum latifolium (Sw.) J. Sm., J. Bot. 1: 197. 1842. Elaphoglossum conforme sensu C. Chr., Trans. Linn. Soc. London, Bot. 7: 420. 1912; sensu Tardieu in Humbert, Fl. Madagasc. 5(2): 35. 1960, non Elaphoglossum conforme (Sw.) J. Sm., J. Bot. 4(27): 148. 1841. Selected specimens examined. MAURITIUS. Alexandra Falls, Rouhan et al. 175 (MAU, NY, P); Bassin Blanc, Lorence 828 in MAU 16319 (MAU); Bassin Blanc to Les Mares path, Lorence in MAU 14610 (MAU); Bel Ombre, Lorence 327 (MO); Black River Gorges from Plaine Champagne, Lorence in MAU 14880 (K, MAU, P); Chamarel rd. from Plaine Champagne, Lorence in MAU 14640 (MAU); Crown Land Declerc, Julien in MAU 13318 (MAU); Macabé to Brise Fer rd., Lorence in MAU 14588 (MAU); Mare Longue Plateau, rd. to Macabé, Lorence in MAU 15242 (MAU, REU), Rouhan et al. 188 (MAU, P, PTBG); Pétrin Nat. Reserve, rd. to Grand Bassin, Lorence 12.1 in MAU 15822 (K, MAU, P, REU), Lorence 18.3 in MAU 15812 (K, MAU, MO, P, REU); Pétrin to Macabé rd., Vaughan in MAU 12551 (MAU, P); Piton Brise Fer, Lorence 3.1 in MAU (K, M, MAU, MO, P); Piton du Fouge, Lorence 1626 (MO); Piton Grand Bassin, Lorence in MAU 14589 (MAU); Plaine Champagne to Chamarel rd., Lorence 14.1 in MAU Volume 91, Number 4 2004 Lorence & Rouhan Mascarene Elaphoglossum 555 Figure 5. Scales of some Mascarene Elaphoglossum species. A–C. E. heterolepis, from Réunion, Plaine des Fougères, Boivin s.n. —A. Rhizome scale. —B, C. Lamina scales. D, E. E. lepervanchei, from Mauritius, Lorence M255. — D. Rhizome scale. —E. Lamina scale. F, G. E. sieberi, from Mauritius, Lorence 15016. —F. Rhizome scale (own scale bar). —G. Lamina scale. —H. Lamina scale of E. tomentosum, from Mauritius, Lorence 1092. Scale bar 5 1 mm, except separate scale in F. 556 Annals of the Missouri Botanical Garden 15811 (K, KRA, MAU, MO, P, REU); Plaine Champagne, S of Black River Gorges, Lorence in MAU 15010 (K, MAU, P); Plaine Champagne, S of Plaine Paul, Lorence 2.3 in MAU 15831 (MAU, MO). RÉUNION. Brûlé, Bédier 52 (pro parte, right-hand specimen P); Cilaos, au dessus des Thermes, Cadet 4130 (REU); Forêt du Bébour, Lorence in MAU 15642 (MAU, MO); sentier de la rivière, Rouhan & Grangaud 248 (NY, P, PTBG); Petite Plaine des Palmistes, Cadet 4017 (REU); Plaine des Fougères, Rouhan et al. 227 (P); Tremblet, Rouhan & Grangaud 208 (NY, P, PTBG); Valleé de Langevin, Cadet 3766 (REU). up to 10 m above ground or rarely terrestrial, in shade of wet forests and ericoid shrublands from 100 to 1500 m. Elaphoglossum macropodium has not been collected in Mauritius since 1923 and is either extremely rare or extirpated there [vs. Réunion]. Discussion. Elaphoglossum macropodium resembles E. sieberi by the frond shape and dimensions, and it resembles E. lepervanchei by frond shape and reddish brown rhizome scales. Elaphoglossum macropodium differs from E. sieberi and E. lepervanchei by its extremely thick rhizomes (10– 15 mm diam.) densely clothed with larger, flat, thin, subentire reddish to yellowish brown scales. The following name was misapplied by Cordemoy and should now refer to E. macropodium: Acrostichum sieberi sensu Cordem., Fl. Réunion, part 1: 81. 1891, Fl. Réunion 93. 1895, non Elaphoglossum sieberi Hook. & Grev, Icon. Filic. tab. 237. 1831. 13. Elaphoglossum macropodium (Fée) T. Moore, Index Fil.: 11. 1857. Acrostichum macropodium Fée, Mém. Foug., 2.: 30, tab. 6(2). 1844 [1845]. TYPE: Bourbon [Réunion]. 1844, s.n., [Hb. Bory 30, 12] (holotype, P!). Figure 2D. Acrostichum conforme Sw. var. carmichaelii Baker, Fl. Mauritius: 511. 1877. TYPE: Mauritius. D. Carmichael s.n. (holotype, K! [Hb. Hook.]). Fronds densely caespitose, borne in 2 ranks; rhizome short-creeping, thick, 10–15 mm diam., unbranched, densely paleaceous; scales narrowly ovate, (10–)15–25 3 2–4 mm, reddish to yellowish brown, flat, thin, truncate to cordate at base, acute at apex, margins bearing rare bulbous-tipped cilia, cells linear to fusiform in vertical rows, thin-walled. Phyllopodia 1–1.5 cm long, swollen, brown to dark brown, densely paleaceous; sterile fronds 30–55(– 73) cm; stipe 8–20(–24) cm 3 3–4 mm, stramineous, adaxially canaliculate, narrowly winged distally, bearing scattered appressed scales; blade narrowly elliptic to ovate- or oblong-elliptic, 18– 36(–50) 3 4–6.5(–8.6) cm, base narrowly to broadly cuneate, decurrent, apex acute to short-acuminate, coriaceous, costa prominulous, rounded abaxially, slightly canaliculate adaxially, margins revolute, veins distinct, prominulous on both surfaces, bifurcate 1 to 2 times, uniting apically into an intramarginal commissure, adaxial surface of blade glabrate, abaxial surface of blade with sparse, scattered appressed brown scales 0.5 mm diam., stellate with bulbous-tipped cilia, scales larger and denser on abaxial surface of costa. Fertile frond 30–55 cm, equaling sterile, stipe equaling or slightly longer than in sterile frond; blade narrowly elliptic, 5–12(–15) 3 2–4 cm, base narrowly to broadly cuneate, apex acute to rounded. Spores 33– 38 3 28–30 mm (excluding perispore), the perispore cristate, the crests 4–10 mm high with finely spinulose-denticulate margins. Distribution and habitat. Mauritius and Réunion; also tropical eastern and southern Africa, Seychelles, and Comoro Islands. Common and widespread in Réunion as an epiphyte on mossy trunks Selected specimens examined. MAURITIUS. Curepipe, Kanaka, Bijoux 73 (P); Savanne, Nov. 1923, s.n. (pro parte, left-hand collection, MAU). RÉUNION. Bébour, Lorence in MAU 15638 (MAU [2], MO); Brûlé, Bédier 54 (P [2]); Saint-Philippe, Bosser 9559 (P, [2]); Montée de la Plaine des Affouches, Bosser 20432 (P); Morne des Patates Durand, Lorence in MAU 15620 (MAU); Plaine des Chicots, sentier Dos d’Ane, Bosser 22485 (P); Salazie, 1835, Lépervanche-Mézières s.n. [Hb. Bory 30,10] (P); Forêt du Grand Brûlé, Cadet 317 (P); Grand Bénard, Boivin 802 (P); Dimitile, Rouhan & Grangaud 207 (NY, P, PTBG); Tremblet, Rouhan & Grangaud 209 (NY, P); Plaine des Fougères, Rouhan et al. 232 (P), Aug. 1850, Boivin s.n. (P). 14. Elaphoglossum 3revaughanii Lorence [Elaphoglossum lepervanchei (Bory ex Fée) T. Moore 3 E. sieberi (Hook. & Grev.) T. Moore], Fern Gaz. 12: 342, fig. 1. 1984. TYPE: Mauritius. Mare Longue Plateau, 1973, D. H. Lorence 14.3 (holotype, MAU 15818!; isotypes, K!, MAU!, MEXU!, MO!, P!, REU!). Fronds caespitose, borne in 2 ranks; rhizome short-creeping, 5–6 mm diam., occasionally branched, densely paleaceous; scales 3–6 3 0.5– 0.8 mm, narrowly ovate to lanceolate, black, shiny, rigid, base truncate to cordate or auriculate with short, glandular cilia, apex long-acute, filiform, margins entire or rarely with a few long, glandular cilia, the cells small, rectangular, seriate, black and opaque. Phyllopodia 15–20 mm long, slightly swollen, grayish to yellowish brown, paleaceous; sterile fronds (30–)44–53 cm; stipe 14–16(–19) cm 3 2– 2.6 mm, stramineous, canaliculate, when young bearing scattered dark brown to black, appressedovate to lanceolate scales 0.5–4 3 0.2–0.5 mm, base cordate to auriculate, fringed by short glandular cilia, apex filiform, sinuate, margins sinuate Volume 91, Number 4 2004 with scattered glandular cilia, cells dark brown; blade narrowly elliptic to linear-elliptic, (19–)30– 35 3 (4.3–)5.5–7 cm, base narrowly cuneate, decurrent along distal 0.25–0.3 of stipe, apex acute to acuminate, subcoriaceous, costa prominulous and rounded abaxially, canaliculate adaxially, margins revolute, veins visible, raised, bifurcate 1 to 2 times, in part apically free and with thickened tips, in part uniting into intramarginal arcs, both surfaces of blade glabrate except for scattered brown, glandular-ciliate scales 0.5 mm diam. along abaxial surface of costa. Fertile frond (30–)35–42 cm long; stipe (15–)17–22 cm; blade narrowly elliptic to linear-elliptic, 15–23 3 1.5–2.6 cm, apex acute to acuminate, base narrowly cuneate, adaxial surface pallid with minute, scattered scales, veins not visible. Spores: 4%–5% very large (75–88 3 50–60 mm, excluding perispore), 65%–67% aborted, 29%–30% normal (40–50 3 30–35 mm, excluding perispore), the perispore with large, ruffled crests 10–15 mm high. Distribution and habitat. Mauritius, endemic and known only from Mare Longue Plateau (630 m) and Pétrin Nature Reserve (660 m). Discussion. This natural hybrid between E. lepervanchei and E. sieberi possesses a high percentage of aborted or abnormal spores and presents morphological characters intermediate between these two species. Elaphoglossum 3revaughanii has veins partly free (as in E. lepervanchei) and partly reunited into a submarginal commissure (as in E. sieberi), and scales on the lower portion of the stipe and phyllopodium are intermediate between those of both parents. The hybrid is more robust than either parent, which both occur in the same biotope, usually as depauperate individuals growing at the bases of stunted trees and shrubs because it is a marginal, suboptimal habitat for these shadeloving, wet forest epiphytes. Obviously more successful than either parent in this particular ecological setting of low shrub vegetation, the hybrid behaves like a terrestrial heliophilous or hemisciophilous species and apparently reproduces vegetatively by rhizome branching (Lorence, 1984). Specimens examined. MAURITIUS. Mare Longue Plateau, Lorence 11.1 (MAU, MEXU, MO), Lorence 8.6 in MAU 15531 (K, MAU, MO, P, REU), Lorence 1176 (MO), Rouhan et al. 190 (MAU, NY, P [2], PTBG); Pétrin Nat. Reserve, Lorence 1497 (MAU, MO). 15. Elaphoglossum richardii (Bory ex Fée) H. Christ, Neue Denkschr. Allg. Schweiz. Ges. Gesammten Naturwiss. 36(1): 35. 1899. Aconiopteris richardii Bory ex Fée, Mém. Foug., 2: 80, tab. 16(4). 1844 [1845]. Acrostichum ri- Lorence & Rouhan Mascarene Elaphoglossum 557 chardii (Bory ex Fée) Hook., Sp. Fil. 256. 1864. TYPE: Bourbon [Réunion]. ‘‘Dans les forêts sur les vieux bois,’’ J. M. C. Richard 10 [Hb. Bory 30,28] (holotype, P!). Figure 1C. Fronds spaced 1–2 cm distant, borne in 2 ranks; rhizome long-creeping, 2–3.5 mm diam., unbranched, paleaceous, resinous; scales 3–6 3 0.3– 0.5 mm, linear-ovate to lanceolate, sinuate, brown to dark brown, shiny, thin, base cordate, apex longacute, margins bearing short cilia or reflexed teeth toward base, cells small, oblong-fusiform (Fig. 6C). Phyllopodia 10–15 mm long, slightly swollen, dark brown; sterile fronds 25–40(–55) cm; stipe 6–12(– 15) cm, 1–2 mm diam., stramineous to brown, when young bearing minute, brown resinous punctations and abundant, ovate to oblong scales to 5 3 0.4 mm with glandular ciliate-dentate margins; blade linear-elliptic, 15–34 3 1.5–2(–3) cm, base narrowly cuneate, often decurrent, apex acute to obtuse, subcoriaceous, costa prominulous and rounded abaxially, canaliculate adaxially, margins revolute, veins distinct, bifurcate, apically uniting into an intramarginal commissure, both surfaces of blade minutely paleaceous when young, scales ovate-triangular to linear-oblong, 0.5–1 3 0.2–0.3 mm, pale brown, scarious, margins with hair teeth, with cluster of resiniferous glands at base (Fig. 6D), adaxial surface glabrescent leaving brown resinous punctations. Fertile frond equaling or slightly longer than sterile frond; stipe slender, equaling or slightly longer than sterile stipe; blade 11–24 3 0.8–1.4 cm, linear, base narrowly cuneate, decurrent, apex acute, adaxial surface paleaceous and with resinous punctations. Spores 30–33 3 23–25 mm (excluding perispore), perispore cristate, crests 2.5–5 mm high, margins finely sinuous-erose. Distribution and habitat. Réunion, endemic. Epiphytic on mossy trees and boulders up to 3 m above ground level in shady wet forests, 300 to 1460 m. Discussion. The specific epithet was published as ‘‘richardi.’’ Since the species was dedicated to Mr. Richard, the publication is valid according to the Code (Art. 32.6; Greuter et al., 2000), but the Latin termination contrary to Rec. 60C.1 is treated as an orthographic error to be corrected (Art. 60.11). Therefore, the correct spelling of the name is ‘‘richardii.’’ This species resembles Elaphoglossum lancifolium by the blade shape, but differs by more widely spaced fronds, blades with non-stellate scales, and veins uniting apically to form a submarginal commissure. In addition to the latter character, E. richardii can be distinguished by linear-elliptic 558 Annals of the Missouri Botanical Garden Figure 6. Scales of some Mascarene endemic Elaphoglossum species. A, B. E. lanatum, from Mauritius, Lorence 15014. —A. Rhizome scale. —B. Lamina scale. C, D. E. richardii, from Réunion, Lorence 15647. —C. Rhizome scale. —D. Lamina scale. E, F. E. splendens, from Réunion, Bory 42 [Hb. Bory 27,28]. —E. Rhizome scale. —F. Lamina scale. G, H. E. stipitatum, from Réunion, Cadet 2048. —G. Rhizome scale. —H. Lamina scale. Scale bar 5 1 mm. Volume 91, Number 4 2004 Lorence & Rouhan Mascarene Elaphoglossum blades (1.5–2 cm wide), paleaceous when young and bearing resinous punctations. The following name misapplied by Tardieu should refer to E. richardii: Elaphoglossum stipitatum sensu Tardieu, Notul. Syst. 15(4): 429, pl. 2, figs. 1–3. 1959, non Elaphoglossum stipitatum (Bory ex Fée) T. Moore, Index Fil.: 15. 1857. long hair teeth, veins prominulous, simple or bifurcate, apically free, both surfaces of blade covered with numerous reddish brown spreading or imbricated scales similar to those on margins but smaller. Fertile frond about as long as sterile frond; stipe longer than sterile stipe, ca. 18 cm; blade ca. 10 3 1.3 cm, narrowly elliptic, base narrowly cuneate, apex obtuse, rounded, scales of adaxial blade surface and both surfaces of costa as in sterile frond. Spores 37–38 3 28–30 mm (excluding perispore), perispore cristate, crests 2.5–5 mm high, margins finely sinuate-dentate. Selected specimens examined. RÉUNION. Basse Vallée, Ravine Pérote, Rouhan 234 (P); Bébour forest, Lorence in MAU 15644 (MAU, MO); Bois des Hauts de SaintPaul, s.n. 75 (P [2]); Boucan Launay, 1852, Bernier s.n. (P); Brûlé, Bédier 56 (P), 57 (P); Chemin Arnoux, Cadet 879 (P); Hauts de l’Ouest, sentier de l’ı̂let Alcide, Rouhan et al. 244 (P); Hauts de St. Rose (Forêt Mourouvin), Cadet 3884 (P, REU); Le Dimitile, Rouhan & Grangaud 205 (NY, P, PTBG); Mare Longue forest, Lorence in MAU 15630 (MAU); Morne des Patates Durand, Lorence in MAU 15647 (MAU, MO, P); Plaine des Fougères, Rouhan et al. 231 (NY, P, PTBG); Plaine des Palmistes, June 1851, Boivin s.n. (P). 16. Elaphoglossum rufidulum (Willd. ex Kuhn) C. Chr., Index Filic.: 314. 1905. Acrostichum rufidulum Willd. ex Kuhn, Filic. Afr. 47. 1868. Elaphoglossum deckenii (Kuhn) C. Chr. var. rufidulum (Willd. ex Kuhn) Tardieu, Notul. Syst. 15(4): 430, pl. 4, figs. 6–10. 1959. Elaphoglossum hirtum (Sw.) C. Chr. var. rufidulum (Willd. ex Kuhn) C. Chr., Dansk. Bot. Ark. 7: 170. 1932. TYPE: Madagascar. J-B. Lamark s.n. (holotype, B-W 19511; B-W microfiche MO!). Figure 3C. Fronds densely caespitose, borne in 2 to 3 ranks; rhizome very short-creeping, 3 mm diam., unbranched, densely paleaceous; scales 2–3 3 1 mm, narrowly ovate, dark brown, shiny, paler basally, base cordate to auriculate, apex acute, margins subentire or bearing black hair teeth 0.1–0.3 mm long, cells rectangular to fusiform with thick walls. Phyllopodia 0.6–1 cm long, slightly swollen, dark brown, densely paleaceous as in rhizome, but scales with long marginal cilia; sterile fronds 23– 34 cm; stipe 9–15 cm, 1 mm diam., stramineous, canaliculate, densely paleaceous, the scales spreading, pale brown to reddish brown, narrowly ovate, 3–4 3 0.5–1.2 mm, base cordate to auriculate, apex acute, margins bearing pale brown to black hair teeth 0.2–0.8 mm long, cells rectangular to oblong-fusiform, often bifurcate apically; blade narrowly elliptic, 13–19 3 2–2.5 cm, base narrowly cuneate, apex short- to long-acuminate, chartaceous, costa prominulous and rounded on both surfaces, margins rounded, weakly revolute, margins and costa bearing many, reddish brown, spreading, ovate scales 1–3 3 0.2–0.5 mm, base auriculate, apex acute, margins bearing 2 to 8 pairs of stiff, 559 Distribution and habitat. Réunion and Madagascar. This species is rare on Réunion, where it occurs on lava in the shade of fissures and grottos in ericoid vegetation from 2200 to 2340 m. Discussion. Elaphoglossum rufidulum resembles E. splendens in general appearance and by its rhizome and frond scales with stiff hair teeth, but E. rufidulum occurs at higher elevations, and its sterile fronds are less densely paleaceous, with the imbricated scales not completely obscuring the frond surface. Furthermore, E. rufidulum lacks black scales on the abaxial surface of the costa, which are present in E. splendens. Specimens from the Comoro Islands closely resemble and may be conspecific with E. rufidulum, but these collections have been referred to E. tanganjicense Krajina ex Pic. Serm. (see Pichi Sermolli, 1968: 238), which does not occur in the Mascarenes. Elaphoglossum kuhnii Hieron. of tropical Africa (absent in the Masacarenes) is also closely related. This complex, which occurs from tropical America to Macaronesia, tropical Africa, the Malagasy region, and Hawai‘i, is in need of critical study. Specimens examined. RÉUNION. Volcan, Massif de la Fournaise, l’Enclos Fouqué, près du Pas de Bellecombe, Cadet 1535 (P, REU); Pas de Bellecombe, bord du sentier descendant dans l’Enclos, Rouhan & Grangaud 224 (P), Des Abbayes 2950 (P). 17. Elaphoglossum 3setaceum Lorence [Elaphoglossum hybridum (Bory) Brack. 3 E. lanatum (Bojer ex Baker) Lorence], Fern Gaz. 12: 348, fig. 5. 1984. TYPE: Mauritius. Tamarind Falls Reservoir, 1973, D. H. Lorence s.n. (holotype, MAU 15823b!). Plant with fronds caespitose, borne in 3 ranks; rhizome very short-creeping, 3 mm diam., unbranched, densely paleaceous; scales 3–4 3 0.5 mm, lanceolate to narrowly ovate, light castaneous, base cordate, apex filiform, margins bearing short teeth each terminated by a globose gland, cells rectangular-fusiform. Phyllopodia 2–4 mm long, 560 Annals of the Missouri Botanical Garden dark brown, paleaceous; sterile fronds 8–14 cm; stipe 3–3.5 cm, 0.5–0.7 mm diam., yellowish brown, canaliculate, bearing numerous simple capitate hairs and abundant, brown, squarrose, subulate scales 1.5–2.5 3 0.2 mm, base cochleariform, apex filiform, margins involute, bearing several long cilia and short glandular teeth or cilia; blade narrowly elliptic to narrowly ovate, (5–)9–10.5 3 (1–) 1.5–2 cm, base narrowly cuneate, decurrent as low ridges along stipe, apex acuminate, chartaceous, costa prominulous on both surfaces, yellow brown, both surfaces bearing spreading subulate scales as in stipe, margins slightly revolute, rounded, ciliate with fringe of pale brown, spreading, bristle-like scales 2–3 3 0.2–0.3 mm, base cochleariform, apex filiform, margins involute basally and bearing several long, spreading glandular-tipped cilia and sparse short hair teeth, veins distinct, bifurcate, apically free, tips thickened, both surfaces of blade villous with bristle-like scales intermixed with smaller flat, deltate scales, each with a single pair of long, spreading basal cilia and scattered simple hairs. Fertile frond and spores unknown. cally entire or rarely with short, glandular projections, the cells narrowly rectangular-fusiform with thick, dark walls (Fig. 5F). Phyllopodia 6–15 mm long, swollen, articulated, dark brown to black, paleaceous; sterile fronds 20–45(–65) cm; stipe 5– 15(–25) cm 3 2–4 mm, stramineous, terete, bearing scattered brown to black, appressed-persistent scales 5–10 3 0.25–0.5 mm, base stellate with glandular cilia, apex filiform; blade narrowly ovate to elliptic or linear-elliptic, 15–30(–42) 3 4–6(–9) cm, base narrowly to broadly cuneate, sometimes attenuate and decurrent, apex acute to obtuse or rounded, subcoriaceous to coriaceous, costa prominulous, rounded abaxially, flattened to canaliculate adaxially, margins revolute, veins indistinct, bifurcate 1 to 2 times, uniting apically into an intramarginal commissure, adaxial surface of blade glabrate, abaxial surface of blade with scattered, appressed brown scales 0.5 mm diam., stellate with bulbous-tipped cilia (Fig. 5G). Fertile frond 16– 30(–45) cm long; stipe as in sterile frond; blade narrowly elliptic to linear-elliptic, 6–20(–36) 3 1.2–2.6(–4) cm, apex acute to obtuse, base cuneate to attenuate, slightly decurrent, veins visible adaxially. Spores 35–38 3 23–26 mm (excluding perispore), the perispore cristate, crisped, the crests 4–10 mm high with erose-dentate margins. Distribution and habitat. Mauritius, known only from the type locality at 500 m in secondary forest. A single sterile individual of this nothospecies was found growing in a population of E. lanatum on a shady stream bank with E. hybridum var. hybridum growing on a tree directly overhead (Lorence, 1984). Discussion. Elaphoglossum 3setaceum possesses morphological characters almost exactly intermediate between those of the putative parents, notably shape of the blade and scale characters. Furthermore, E. 3setaceum presents a rhizome bearing three ranks of fronds, whereas E. lanatum is a two-ranked species and E. hybridum var. hybridum has four ranks (Lorence, 1984). 18. Elaphoglossum sieberi (Hook. & Grev.) T. Moore, Index Fil.: 14. 1857. Acrostichum sieberi Hook. & Grev., Icon. Filic. tab. 237. 1831. Olfersia sieberi (Hook. & Grev.) C. Presl, Tent. Pterid. 235. 1836. TYPE: Mauritius. W. Bojer s.n. (lectotype, designated by Lorence (1976: 204), K! [Hb. Hook., pro parte, left-hand specimen only]). Figure 1E. Fronds densely caespitose, borne in 2 ranks; rhizome very short-creeping, 7–10 mm diam., unbranched, densely paleaceous; scales 8–20 3 0.5– 1 mm, narrowly linear-ovate, dull brown to black, base enlarged, cordate to auriculate, distally filiform and often sinuate, the margins fringed basally with short, glandular projections, medially and api- Distribution and habitat. Mauritius, endemic. One of the most commonly encountered members of the genus on Mauritius. Elaphoglossum sieberi grows as an epiphyte to 10–12 m above ground or casually terrestrial, in moist and wet forests and ericoid shrublands, at 200 to 900 m. Discussion. Elaphoglossum sieberi resembles E. macropodium by the shape, texture, and size of the fronds, but can be distinguished from E. macropodium and all others in the Mascarenes by a thinner rhizome (7–10 mm diam.) densely clothed with long (8–20 mm), narrowly linear-ovate, dull brown to black scales. Selected specimens examined. MAURITIUS. Alexandra Falls, Rouhan et al. 169 (MAU, NY, P, PTBG); Bassin Blanc, 1966, Edgerley s.n. in MAU 12464 (MAU, P); Curepipe, Kanaka, 1906, Bijoux s.n. (MAU); Curepipe, Vaughan in MAU 2566 (MAU); Deux Mamelles Mtn., Lorence in MAU 15016 (MAU); Ferney, mtn. ridge, Lorence 773 in MAU 16314 (MAU); Le Pouce Mtn., Aug. 1849, Boivin s.n. (P); Les Mares, Rouchecouste & Orian s.n. (MAU [2]); Macabé, Lorence in MAU 14584 (MAU); Macabé–Brise Fer rd., Lorence in MAU 14586 (MAU); Mt. Deux Mamelles, Lorence in MAU 15016 (MAU, P); Mt. Lagrave, E flank, Lorence 779 in MAU 16318 (MAU), Lorence 782 in MAU 16313 (MAU); Pétrin Nat. Reserve, Lorence in MO M129 (MO, P); Pétrin–Les Mares rd., Vaughan in MAU 12624 (MAU); Piton Combo, Lorence 890 (MO); Piton du Milieu, Lorence 10.2 in MAU 15814 (MAU, REU); Piton Grand Bassin, Lorence 1074 (MO); Piton Volume 91, Number 4 2004 Lorence & Rouhan Mascarene Elaphoglossum Grand Fond, Lorence 301 (MO); Plaine Champagne nr. Cascade 500 Pieds, Lorence s.n. (MO); Plaine Champagne–Chamarel rd., headwaters of Rivière Saint Denis, Julien in MAU 14585 (MAU); Plaine Champagne–Bel Ombre rd., Lorence 9.1 (MO); Plaine Champagne near Plaine Paul, Lorence 2.2 (MO); Plaine Paul near path to Mt. Cocotte, Vaughan in MAU 12593 (MAU). Discussion. Elaphoglossum spatulatum can be easily distinguished from all other Elaphoglossum in the Mascarenes by the small fronds 3–7(–9) cm long bearing subulate scales and the fertile fronds conduplicate before maturity. Furthermore, E. spatulatum is unique in growing exclusively on boulders in the beds of rivers. Elaphoglossum spatulatum greatly resembles E. piloselloides (C. Presl) T. Moore, a tropical American species (Moran & Smith, 2001), but E. spatulatum differs by having the scales of the fertile fronds orange-tan rather than black (Mickel, 2002). The close relationship between both species is strongly supported by phylogenetic analyses based on molecular data, since E. spatulatum and E. piloselloides appear more related to each other than to any other species in the genus (Rouhan et al., 2004). Thus, E. spatulatum may possibly be conspecific with E. piloselloides (in which case E. spatulatum would have priority). 19. Elaphoglossum spatulatum (Bory) T. Moore, Index Fil.: 14. 1857. Acrostichum spatulatum Bory, Voy. Iles Afrique 1: 363, tab 20, fig. 1. 1804. TYPE: Bourbon [Réunion]. Rivière St. Denis, ‘‘an X’’ [1801–1802], J. B. G. G. M. Bory de St. Vincent s.n. [Hb. Bory 27,34] (holotype, P!). Figure 4C, D. Fronds caespitose, borne in 3(to 4) ranks; rhizome very short-creeping, 1.5–2 mm diam., short and unbranched, densely paleaceous; scales 3–4 3 0.3–0.4 mm, narrowly ovate, membranaceous, reddish brown, base cordate to auriculate, apex narrowly acuminate, filiform, margins entire, cells fusiform. Phyllopodia absent; sterile fronds 3–7(–9) cm; stipe 2–4(–6) cm, 0.5 mm diam., stramineous, slightly canaliculate, densely paleaceous with spreading, narrow, reddish brown, subulate scales 2–3 3 0.3–0.4 mm, base cordate, the lobes involute, apex long-acuminate, margins with scattered short teeth basally, distally entire; blade narrowly obovate-spatulate to narrowly elliptic, 1.5–4(–5.5) 3 0.5–0.8(–1.2) cm, base narrowly cuneate and decurrent, apex obtuse or rarely acute, thickly chartaceous, costa prominulous and rounded on both surfaces, margins thin, cartilaginous, revolute, veins bifurcate, apically free, tips thickened and dark, both surfaces of blade and margins with scattered slender, subulate scales like those on stipe but smaller, mixed with simple capitate trichomes. Fertile frond slightly longer than sterile frond, 5– 8(–10) cm long; stipe slender, often longer than sterile stipe; blade 1–2.3 3 0.7–1 cm, ovate, elliptic, or obovate, conduplicate before maturity, base cuneate, shortly decurrent, apex obtuse, often emarginate, adaxial surface paleaceous as in fertile frond, margins thin, cartilaginous. Spores 30–35 3 20–25 mm (excluding perispore), surface cristate, crests 3–5 mm high with entire, sinuate margins. Distribution and habitat. Réunion, Mauritius; also known from Madagascar, tropical Africa, extratropical southern Africa, and Sri Lanka (Sledge, 1967). This diminutive species is common on Réunion where it grows exclusively on mossy boulders and rocks in the beds of rivers and streams from 900 to 1400 m. On Mauritius E. spatulatum is known only from two collections made over a century ago; it may be extirpated. 561 Selected specimens examined. MAURITIUS. Woods of Grand Port and Savanne, [probably Bijoux], s.d., s.n, ex Hb. Pamplemousses s.n. (MAU). RÉUNION. Basse Vallée, Ravine Pérote, Rouhan 235 (NY, P, PTBG); Bébour, Rouhan & Grangaud 246 (NY, P, PTBG); Brûlé, 1891, Bedier 51 (P); Hautes de Ste. Marie, Coode & Cadet 5007 (K, MAU); Montée de la Plaine des Affouches, Bosser 20460 (P); Montée de la Plaine des Palmistes, Bosser 9561 (P); Morne de Patates Durand, Lorence FR6 in MAU 15618 (MAU); Rivière des Marsouins, M. G. De L’Isle 82 (P); Takamaka, Guého in MAU 15363 (MAU). 20. Elaphoglossum splendens (Bory ex Willd.) Brack., U.S. Expl. Exped., Filic. 16: 68 1854. Acrostichum splendens Bory ex Willd., Sp. Pl. ed. 4, 5: 104. 1810. TYPE: Bourbon [Réunion]. An X [1801–1802], J. B. G. G. M. Bory de St. Vincent s.n. [Hb. Bory 27,25] (holotype, B-W 19515, B-W microfiche MO!; isotype, P!). Figure 3A. Fronds densely caespitose, borne in 4 ranks; rhizome short-creeping, 3–4 mm diam., unbranched, densely paleaceous; scales 2.5–3.5 3 0.6–1 mm, narrowly ovate, castaneous or dark brown, shiny, stiff, base cordate, apex acute, margins bearing short hair teeth (Fig. 6E). Phyllopodia 1–1.5 cm long, slightly swollen, dull brown, densely paleaceous as in rhizome; sterile fronds 25–40(–53) cm; stipe 4–16(–30) cm, 1.5–2 mm diam., pale orangebrown, canaliculate, densely paleaceous, the scales spreading, dark reddish brown, ovate to linear-oblong, 3–5 3 0.5–1.5 mm, base truncate to cordate, apex acute, often black, margins bearing many dark brown to black stiff hair teeth, cells shortly fusiform to irregularly oblong, with thick walls; blade linearovate to linear-elliptic, often falcate, 16–24(–33) 3 562 Annals of the Missouri Botanical Garden 1.5–2.5(–3.6) cm, base narrowly cuneate to attenuate, apex acute, acuminate or sometimes caudate, chartaceous to subcoriaceous, costa prominulous on both surfaces, adaxially flattened to slightly canaliculate, abaxially rounded, margins rounded, sometimes weakly revolute, fringed with spreading scales, veins bifurcate, apically free, both surfaces of blade covered with a dense, persistent layer of imbricated reddish brown scales aging grayish adaxially, those on abaxial surface of costa usually dark brown to blackish, scales ovate to narrowly ovate, (0.3–)1.5–2.5 3 0.2–0.5 mm, base cordate, apex acute, margins fringed with long, erectspreading hair teeth (Fig. 6F). Fertile frond as long as sterile frond; stipe 1 to 1.5 times as long as sterile stipe; blade 14–18(–34) 3 1.2–1.4(–1.9) cm, linear to linear-elliptic, base cuneate, truncate or cordate, apex acute to obtuse, scales of adaxial surface and abaxial surface of costa as in sterile frond. Spores 38–42 3 25–28 mm (excluding perispore), perispore weakly cristate, crests 2.5–4 mm high, margins entire to sinuate. Grande Montée de la Plaine des Cafres, Lorence 2463 (MO); Dimitile, Rouhan & Grangaud 203 (NY, P, PTBG); Grand Matarum, Lorence in MAU 15631 (MAU), Staub in MAU 13012 (MAU); Hauts de Saint-Paul, 2 June 1882, s.d. 74 (P); L’ı̂let Alcide, Rouhan et al. 243 (P); Montée de la Plaine des Palmistes, Bosser 9529 (P [pro parte, lefthand specimen]); Piton des Neiges, near le Gite, Lorence in MAU 15650 A-C (MAU); Plaine des Chicots, Bory de St. Vincent s.n. (P); Plaine des Fougères, Boivin 799 (P); St. Philippe, Schlieben 10934 I (MAU 12441); Tévelave near Les Avirons, Schlieben 10849 (MO); s. loc., Commerson s.n. (P-JU 1002). Distribution and habitat. Réunion and Mauritius. Elaphoglossum splendens is common on Réunion, where it occurs as an epiphyte in the shade of wet forests and cloud forests at 1000 to 2350 m, or occasionally on mossy boulders in ericoid vegetation. It is extremely rare or perhaps extirpated on Mauritius, where it is known only from two collections made in the early 1900s. Discussion. Elaphoglossum splendens resembles E. rufidulum in general appearance and by its rhizome and frond scales with stiff hair teeth, but E. splendens occurs at lower elevations, and its sterile fronds are more densely paleaceous, with the imbricated scales completely obscuring the frond surface. Furthermore, E. splendens has black scales on the abaxial surface of the costa, which are absent in E. splendens. The following misapplied name found in the early Mascarene literature should refer to E. splendens: Acrostichum squamosum sensu Baker, Fl. Mauritius 512. 1877, sensu Cordem., Fl. Réunion, part 1: 85. 1891, Fl. Réunion 96. 1895, non Acrostichum squamosum Sw., J. Bot. (Schrader) 1800(2): 11. 1801. Selected specimens examined. MAURITIUS. Grand Bassin, Pétrin, Bijoux 76 (MAU, P); s.d., s.n., ex Hb. Pamplemousses in MAU 17656 (MAU). RÉUNION. Bébour, Rouhan & Grangaud 247 (NY, P, PTBG); Bébour, path to Bassin des Hirondelles, Guého in MAU 15366 (MAU); Cilaos, sentier de la Mare à Joseph au coteau Kerveguen, Badré 937 (P [2]); Cilaos, sentier GR R1 entre ‘‘Le Bloc’’ et le gı̂te de la Caverne Dufour, Rouhan 213 (NY, P, PTBG), Rouhan 218 (NY, P); Cirque de Salazie, N of Piton Marmite, Lorence et al. 7544 (MO, PTBG); Cirque de Salazie, Rouhan et al. 225 (P, PTBG); Col de Bellevue, 21. Elaphoglossum stipitatum (Bory ex Fée) T. Moore, Index Fil.: 15. 1857. Acrostichum stipitatum Bory ex Fée, Mém. Foug., 2: 38, tab. 4(3). 1844 [1845]. TYPE: Bourbon [Réunion]. Salazie, s.d., s.n. [Hb. Bory 27,12] (holotype, P!). Figure 3D. Fronds spaced 0.5–1.5 cm, borne in 2 ranks; rhizome long-creeping, 1.5–2 mm diam., occasionally branched, densely paleaceous; scales 3–5 3 0.4–0.6 mm, narrowly ovate to linear-ovate, dark reddish brown to blackish, paler basally, base cordate to sagittate, apex narrowly acuminate, sinuate, filiform, margins bearing short, often paired hair teeth, cells fusiform, in rows (Fig. 6G). Phyllopodia 3–6 mm long, slightly swollen, dark brown; sterile fronds 8–22(–44) cm; stipe 3–16(–25) cm, 0.5–1 mm diam., flexuous, stramineous to yellowish brown, bearing numerous, spreading, rusty, narrowly ovate scales 2–3 3 0.3–0.8 mm, base cordate to auriculate, apex long-acute, slender, margins with short hair teeth, brown resinous punctations also present; blade oblong- to linear-ovate or oblong- to linear-elliptic, 4.5–8(–19) 3 0.9–1.5(–2) cm, base cuneate to narrowly cuneate, shortly decurrent, apex acute to obtuse, subcoriaceous to coriaceous, costa prominulous and rounded abaxially, flattened to shallowly canaliculate adaxially, both surfaces of blade bearing spreading scales as in stipe, margins thin, revolute, veins distinct, unbranched or 1 to 2 times bifurcate, apically free, adaxial surface thinly covered with rusty to gray or white, narrowly ovate scales 1–2 3 0.2–0.3 mm long, base cordate to sagittate, apex slender, margins and basal lobes bearing long, paired hair teeth, cells rectangular to fusiform (Fig. 6H), abaxial surface glabrous except costa bearing scattered scales like those on adaxial surface of blade, both surfaces of blade bearing short capitate trichomes and brown resinous punctations. Fertile frond about equaling sterile frond; stipe slender, 1.5 to 2.5 times longer than sterile stipe; blade 5–7(–8.5) 3 1–1.2(–1.4) cm, shape and scales of adaxial surface as in sterile frond. Spores 38–40 3 25–30 mm (excluding perispore), Volume 91, Number 4 2004 Lorence & Rouhan Mascarene Elaphoglossum perispore cristate, crests 3–6 mm high, margins erose-dentate. narrowly cuneate to attenuate, apex acute to longacuminate, subcoriaceous, costa prominulous and rounded abaxially, shallowly canaliculate adaxially, yellowish orange, margins rounded, slightly revolute, veins unbranched or bifurcate, tips apically uniting into an intramarginal commissure, both surfaces of young blade covered by a thin but dense layer of closely appressed, contiguous ovate to scutelliform, peltate scales 0.5–1 mm diam., pale brown or gray with darker centers, margins fimbriate, central cells square, marginal cells elongate (Fig. 5H), adaxial blade surface glabrescent. Fertile frond 20–28(–38) cm long; stipe slender, as long as or slightly exceeding sterile stipe; blade 13–19(– 23) 3 0.9–1.3(–2.3) cm, narrowly elliptic to linearelliptic, base cuneate to attenuate, apex acute, adaxial surface bearing a dense layer of contiguous scales as in sterile frond. Spores 35–38 3 23–25 mm (excluding perispore), perispore cristate, 5–7.5 mm high with finely erose-dentate margins. Distribution and habitat. Elaphoglossum stipitatum is endemic to Réunion, where it occurs on rocks or terrestrially in mats of bryophytes at 2000 to 2500 m, or in ericoid Philippia Klotzsch shrublands. Discussion. Elaphoglossum stipitatum can be distinguished by its sterile fronds with an adaxially scaly, abaxially glabrous blade surface bearing short capitate trichomes, and brown resinous punctations. Furthermore, it is not epiphytic but occurs at high elevations on rocks or terrestrially. Elaphoglossum stipitatum is very closely allied to or perhaps conspecific with E. subcinnamomeum (H. Christ) Hieron. from Kenya, Tanzania, and Cameroon. Both species grow at high elevations and exhibit short capitate trichomes in addition to similar scale characters. Nevertheless, we refrain from placing E. subcinnamomeum in synonymy until more specimens of this species have been collected and examined. Selected specimens examined. RÉUNION. Cilaos, GR R1 entre ‘‘Le Bloc’’ et le gı̂te de la Caverne Dufour, Rouhan 212 (NY, P, PTBG); Cratère Commerson, M. G. De L’Isle 396 bis (P [2]); Piton des Neiges, Lorence 407 (MO); Plaine des Salazes, Cadet 2048 (P); Salazie, s.n. [Hb. Bory 27,12] (P); Sommet du Grand Bénard, 16 Feb. 1847, Boivin 1847–1852 (P); Volcan, Pas de Bellecombe, Rouhan & Grangaud 221 (NY, P, PTBG). 22. Elaphoglossum tomentosum (Bory ex Willd.) H. Christ, Farnkr. Erde 37. 1897. Acrostichum tomentosum Bory ex Willd., Sp. Pl. ed. 4, 5: 101. 1810. TYPE: ‘‘B. Ile de France’’ [Mauritius]. J. B. G. G. M. Bory de St. Vincent s.n. [Hb. Bory 27,26] (lectotype, designated by Rouhan & Lorence (2003: 840), P!). Figure 2E. Fronds caespitose, borne in 2 ranks; rhizome short-creeping, 4 mm diam., occasionally branched, moderately paleaceous; scales 1.5–3 3 0.25–0.5 mm, triangular to unguiculate, dull dark brown, appressed, persistent, base truncate to cordate, apex acuminate, margins entire, scarious to erose, cells opaque. Phyllopodia 8–10 mm long, slightly swollen, brown to dark brown, bearing scales as in rhizome; sterile fronds 25–40(–50) cm; stipe 4–8(–11) cm, 1.2–2 mm diam., pale yellowish orange, canaliculate, narrowly winged distally, when young bearing numerous, dull brown, opaque elliptic to unguiculate scales 1.5–2.5 mm long, 0.4–0.6 mm wide, margins fimbriate, cells fusiform to irregularly polygonal-rounded, thick-walled; blade linear-oblong, 20–40(–44) 3 2–3(–4.6) cm, base cuneate to 563 Distribution and habitat. Mauritius, endemic. Wet forests from 300 to 900 m, growing epiphytically to 10 m above the ground, and occasionally on boulders or terrestrially. Although three old collections are said to be from Bourbon [Réunion], no recent collections are known from that island. Discussion. Elaphoglossum tomentosum resembles E. heterolepis by the blade covered by a thin continuous layer of appressed scales, but differs by veins that unite apically into an intramarginal commissure, by its single type of laminar scales that form a thin layer over the blade, and by the dull brown, monochrome rhizome and stipe scales. On Mauritius E. tomentosum occasionally hybridizes with E. heterolepis producing the nothospecies E. 3heterophlebium. It also hybridizes with the endemic E. lanatum producing E. 3adulterinum (Lorence, 1984). The following misapplied name is very common in the literature for the Mascarene area: Elaphoglossum (or Acrostichum) obductum sensu auct. (see discussion under E. lancifolium). Selected specimens examined. MAURITIUS. Alexandra Falls, Rouhan et al. 178 (MAU, NY, P, PTBG); Bel Ombre nat. reserve, Lorence in MAU 14605 (MAU); Brise Fer, Lorence 17.1 (MO); Crown Land Declerc, Lorence 297 (MO); Gorges de la Rivière Noire, Crown Land Le Bouton, Lorence 1092 (MO, P); Le Pouce Mt., Julien in MAU 15180 (MAU); bois de la partie supérieure, Aug. 1849, Boivin s.n. (P); Macabé forest, Lorence 1303 (MO); Macabé, sentier menant à la Mare aux Joncs, Rouhan et al. 180 (MAU, NY, P, PTBG); Macabé-Brise Fer rd., Lorence in MAU 14601 (MAU); Mare Longue, Vaughan in MAU 10084 (MAU); Mt. Lagrave, E flank, Lorence 793 in MAU 18780 (MAU); Pétrin–Macabé rd., Lorence in MAU 14587 (MAU); Pétrin Nat. Reserve, Lorence 13.5 in MAU 15817 564 Annals of the Missouri Botanical Garden (K, MAU, MO, P, REU); Plaine Champagne, Lorence in MAU 15011 (K, MAU, P, REU); valley of Cascade 500 Pieds, Lorence 1166 (K, MO); valley above Cascade 500 Pieds, Lorence et al. 6990 (MAU, PTBG). RÉUNION. [Probably in error], s.l., Habitat in insula Borboniae [Réunion], Bory de St. Vincent s.n. [legit. Flügge, in sched.], (B-W 19508), Sieber s.n. ex Hb. Bernardi (MO), s.n., ex Hb. Pamplemousses s.n. (MAU). logical reconnaissance of Rodriguez Island, Indian Ocean. Nature 206: 26–27. Mabberely, D. J. 1997. The Plant-Book, 2nd ed. Cambridge Univ. Press, Cambridge. Manton, I. & W. A. Sledge. 1954. Observations on the cytology and taxonomy of the pteridophyte flora of Ceylon. Philos. Trans., Ser. B, 238: 127–180. Mickel, J. T. 2002. Elaphoglossum. Pp. 7–27 in H. J. Beentje (editor), Flora of Tropical East Africa, Lomariopsidaceae. A. A. Balkema, Rotterdam. & L. Atehortúa. 1980. Subdivision of the genus Elaphoglossum. Amer. Fern J. 70: 47–68. & J. Beitel. 1988. Pteridophyte flora of Oaxaca, Mexico. Mem. New York Bot. Gard. 48: 1–568. Moore, T. 1857–1862. Index Filicum: A synopsis, with characters, of the genera, and an enumeration of the species of ferns, with synonymes, references. . .. London. Moran, R. C. & R. Riba. 1995. Elaphoglossum. Pp. 250– 283 in G. Davidse, M. Sousa S. & S. Knapp (editors), Flora Mesoamericana, Vol. 1. Universidad Nacional Autónoma de México, México, D.F.; Missouri Botanical Garden, St. Louis; The Natural History Museum, London. & A. R. Smith. 2001. Phytogeographic relationships between neotropical and African-Madagascan pteridophytes. Brittonia 53: 304–351. Morton, C. V. 1965. IV. Proposals in pteridophyta. Elaphoglossum Schott ex J. Smith. Regnum Veg. 40: 18– 19. . 1967. Studies of fern types, I. Contr. U.S. Natl. Herb. 38: 29–83. . 1973. Studies of fern types, II. Contr. U.S. Natl. Herb. 38: 215–281. Palmer, D. D. 2003. Hawaii’s Ferns and Fern Allies. Univ. Hawaii Press, Honolulu. Pichi Sermolli, R. E. G. 1968. Adumbratio florae aethiopicae. Elaphoglossaceae. Webbia 23: 209–246. Rouhan, G. & D. H. Lorence. 2003. On the typification of two Mascarene Elaphoglossum (Lomariopsidaceae): E. heterolepis (Fée) T. Moore and E. tomentosum (Bory ex Willd.) H. Christ. Taxon 52: 837–840. , J.-Y. Dubuisson, F. Rakotondrainibe, T. J. Motley, J. T. Mickel, J.-N. Labat & R. C. Moran. 2004. Molecular phylogeny of the fern genus Elaphoglossum (Elaphoglossaceae) based on chloroplast non-coding DNA sequences: Contributions of species from the Indian Ocean area. Molec. Phylogenet. Evol. 33: 745–763. Roux, J. P. 2001. Conspectus of Southern African Pteridophyta. Southern African Botanical Diversity Network Report No. 13. SABONET, Pretoria. Schelpe, E. A. C. L. E. 1969. Revision of tropical African pteridophytes. 4. The Lomariopsidaceae of continental tropical Africa. Contr. Bolus Herb. 1: 25–36. Schott, H. W. 1834. Genera Filicum. Wallishausser, Vienna. Sledge, W. A. 1967. The genus Elaphoglossum in the Indian peninsula and Ceylon. Bull. Brit. Mus. (Nat. Hist.), Bot. 4: 81–96. Smith, J. 1841. An arrangement and definition of the genera of ferns, with observations on the affinities of each genus. J. Bot. (Hooker) 4(27): 38–70, 147–198. Stafleu, F. A. & R. S. Cowan. 1976. Taxonomic literature: A selective guide to botanical publications and collections with dates, commentaries and types. Vol. 1: A–G. Bohn, Scheltema & Holkema, Utrecht. Tardieu-Blot, M.-L. 1959. Sur les Elaphoglossum de la région malgache avec description d’espèces nouvelles. Notul. Syst. 15: 425–443. . 1960. Polypodiacées (sensu lato). Blechnacées– Literature Cited Alston, A. H. G. 1956. The subdivision of the Polypodiaceae. Taxon 5: 23–25. . 1959. The Ferns and Fern-Allies of West Tropical Africa. A supplement to the second edition of the flora of West Tropical Africa. Crown Agents for Oversea Governments and Administrations, Millbank, London. Badré, F. & T. Cadet. 1978. The pteridophytes of Réunion Island. Fern Gaz. 11: 349–365. Baker, J. G. 1877. Flora of Mauritius and the Seychelles. London. Bojer, W. 1837. Hortus Mauritianus. Port Louis, Mauritius. Christ, H. 1899. Monographie des Genus Elaphoglossum. Neue Denkschr. Allg. Schweiz. Ges. Gesammten Naturwiss. 36(1): 1–159. Cordemoy, E. J. de. 1895. Flore de l’ı̂le de la Réunion. Paris. Crabbe, J. A., A. C. Jermy & J. T. Mickel. 1975. A new generic sequence for the pteridophyte herbarium. Fern Gaz. 11: 141–162. Fée, A. L. A. 1845. Mémoire sur les familles des fougères. Deuxième mémoire: Histoire des Acrostichées. BergerLevrault, Strasbourg. Fisher, R. L., G. L. Johnson & B. C. Heezen. 1967. Mascarene Plateau, western Indian Ocean. Bull. Geol. Soc. Amer. 78: 1247–1266. Greuter, W., J. McNeill, F. R. Barrie, H. M. Burdet, V. Demoulin, T. S. Filgueiras, D. H. Nicolson, P. C. Silva, J. E. Skog, P. Trehane, N. J. Turland & D. L. Hawksworth (editors). 2000. International Code of Botanical Nomenclature (Saint Louis Code). Regnum Veg. 138. Hennipman, E. 1977. A Monograph of the Fern Genus Bolbitis (Lomariopsidaceae). Leiden Botanical Series, Vol. 2. Leiden Univ. Press, Leiden. Kramer, K. U., P. S. Green & A. Goetz. 1990. The Families and Genera of Vascular Plants, I. Pteridophytes and Gymnosperms. Ed. K. Kubitzki. Springer-Verlag, Berlin, New York. Lorence, D. H. 1976a. Notes on some Mascarene species of Elaphoglossum (Lomariopsidaceae sensu Holttum). Fern Gaz. 11: 199–205. . 1976b. The pteridophytes of Rodrigues Island. Bot. J. Linn. Soc. 72: 269–283. . 1978. The pteridophytes of Mauritius (Indian Ocean): Ecology and distribution. Bot. J. Linn. Soc. 76: 207–247. . 1984. Hybridization in Elaphoglossum in the Mascarene Islands. Fern Gaz. 12: 341–350. . 1985. A monograph of the Monimiaceae (Laurales) in the Malagasy Region (Southwest Indian Ocean). Ann. Missouri Bot. Gard. 72: 1–165. . 1992. Phytogeography of Mascarene Pteridophytes. Amer. J. Bot. (supplement) 79: 115. [Abstract.] McDougall, I. & F. H. Chamalaun. 1969. Isotopic dating and geomagnetic polarity studies on the volcanic rocks from Mauritius, Indian Ocean. Bull. Geol. Soc. Amer. 80: 1419–1442. , B. G. J. Upton & W. J. Wadsworth. 1965. A geo- Volume 91, Number 4 2004 Polypodiacées in H. Humbert, Flore de Madagascar et des Comores (Plantes Vasculaires). 5e Famille, 2. Paris. Tryon, R. M. & A. F. Tryon. 1982. Ferns and Allied Plants with Special Reference to Tropical America. SpringerVerlag, New York. Voss, E. G., H. M. Burdet, W. G. Chaloner, V. Demoulin, P. Hiepko, J. McNeill, R. D. Meikle, D. H. Nicolson, R. C. Rollins, P. C. Silva & W. Greuter. 1983. International Code of Botanical Nomenclature (Sydney Code). Regnum Veg. 111. Wagner, W. H., Jr. & K. L. Chen. 1965. Abortion of spores and sporangia as a tool in the detection of Dryopteris hybrids. Amer. Fern J. 55: 9–29. , F. S. Wagner & W. C. Taylor. 1986. Detecting abortive spores in herbarium specimens of sterile hybrids. Amer. Fern J. 76: 129–140. APPENDIX 1. List of species and varieties. 1. 2. 3. 4. 5. 6. 7. 8. 9. 9a. 9b. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. Elaphoglossum acrostichoides (Hook. & Grev.) Schelpe Elaphoglossum 3adulterinum Lorence Elaphoglossum angulatum (Blume) T. Moore Elaphoglossum aubertii (Desv.) T. Moore Elaphoglossum 3cadetii Lorence Elaphoglossum coursii Tardieu Elaphoglossum heterolepis (Fée) T. Moore Elaphoglossum 3heterophlebium Lorence Elaphoglossum hybridum (Bory) Brack. Elaphoglossum hybridum (Bory) Brack. var. hybridum Elaphoglossum hybridum var. vulcani (Lepervanche ex Fée) H. Christ Elaphoglossum lanatum (Bojer ex Baker) Lorence Elaphoglossum lancifolium (Desv.) C. V. Morton Elaphoglossum lepervanchei (Bory ex Fée) T. Moore Elaphoglossum macropodium (Fée) T. Moore Elaphoglossum 3revaughanii Lorence Elaphoglossum richardii (Bory ex Fée) H. Christ Elaphoglossum rufidulum (Willd. ex Kuhn) C. Chr. Elaphoglossum 3setaceum Lorence Elaphoglossum sieberi (Hook. & Grev.) T. Moore Elaphoglossum spatulatum (Bory) T. Moore Elaphoglossum splendens (Bory ex Willd.) Brack. Elaphoglossum stipitatum (Bory ex Fée) T. Moore Elaphoglossum tomentosum (Bory ex Willd.) H. Christ APPENDIX 2. Index to exsiccatae. The numbers in parentheses refer to the corresponding species and varieties in the text and in Appendix 1. Specimens are listed in alphabetic order by first collector. Collection numbers in boldface type indicate type specimens. Badré 866 (1), 896 (7), 937 (20), 960 (9b), 985 (7), 1004 (4), 1005 (9a); Bédier 51 (19), 52 (12), 53 (7), 54 (13), 56 (15), 57 (15); Bijoux 69 in MAU 2570 (11), 73 (13); Boivin [1847–1852] (21), 799 (20), 800 (7), 801 (1), 802 (13); Bosser 9529 (20), 9544 (4), 9559 (13), 9561 (19), 11857 (9a), 20432 (13), 20460 (19), 20624 (11), 21012 (6), 21503 (4), 21759 (4), 22485 (13). Cadet 123 (4), 317 (13), 319 bis (11), 504 (3), 805 (1), 879 (15), 880 (7), 1535 (16), 1556 (3), 1556 (9a), 1702 (7), 1749 (1), 1778 (3), 1929A (1), 1930 (3), 1930 bis (7), 2048 (21), 2063 (5), 3718 (11), 3766 (12), 3790 (7), 3884 (15), 4017 (12), 4028 (1), 4122 (1), 4126 (6), 4127 (11), 4130 (12); Coode 5007 (19). De L’Isle 82 (19), 396 bis (21); Des Abbayes 2950 (16); Du Petit-Thouars s.n. [1801] (11), s.n. [1808] (4). Lorence & Rouhan Mascarene Elaphoglossum 565 Edgerley s.n. in MAU 12464 (18). Guého in MAU 11498 (9a), in MAU 15303 (11), in MAU 15362 (11), in MAU 15363 (19), in MAU 15366 (20). Humbert 24918 (6). Julien in MAU 13318 (12), in MAU 14585 (18), in MAU 14594 (11), in MAU 14595 (9a), in MAU 15180 (22), in MAU 16494 (9a). Lépervanche-Mézières 14 (3); Lorence 2.2 (18), 2.3 in MAU 15831 (12), 2.11 (7), 3.1 in MAU (12), FR6 in MAU 15618 (19), 7.3 in MAU 15813 (11), 8.5 (11), 8.6 in MAU 15531 (14), 9.1 (18), 10.2 in MAU 15814 (18), 10.3 (11), 11.1 (14), 12.1 in MAU 15822 (12), 13.5 in MAU 15817 (22), 14.1 in MAU 15811 (12), 14.2 in MAU 15821 (8), 14.3 in MAU 15818 (14), 16.2 in MAU 15819 (8), 17.1 (22), 17.2 in MAU 15830 (10), 18.3 in MAU 15812 (12), 21.1 in MAU 16280 (10), M129 (18), 300 (7), 301 (18), 318 (11), 327 (12), 407 (21), 408 (3), 506 (11), 773 (18), 779 in MAU 16318 (18), 782 in MAU 16313 (18), 783 in MAU 16317 (7), 792 in MAU 16315 (11), 793 in MAU 18780 (22), 828 in MAU 16319 (12), 847 (2), 890 (18), 996 in MAU 16518 (8), 1074 (18), 1078 (7), 1092 (22), 1166 (22), 1173 (10), 1176 (14), 1303 (22), 1374 (11), 1402 (7), 1406 (10), 1420 in MAU 17544 (7), 1479 (11), 1497 (14), 1544 (8), 1608 (9a), 1609 (2), 1610 (2), 1626 (12), 2463 (20), 2496 (9b), 2520 (1), 6944 (9a), 6990 (22), 7017 (11), 7544 (20), 7561 (6), in MAU 14584 (18), in MAU 14586 (18), in MAU 14587 (22), in MAU 14588 (12), in MAU 14589 (12), in MAU 14599 (7), in MAU 14601 (22), in MAU 14605 (22), in MAU 14609 (7), in MAU 14610 (12), in MAU 14640 (12), in MAU 14880 (12), in MAU 15010 (12), in MAU 15011 (22), in MAU 15016 (18), in MAU 15024 (9a), in MAU 15099 (8), in MAU 15242 (12), in MAU 15243 (10), in MAU 15533 (10), in MAU 15612 (3), in MAU 15614 (1), in MAU 15615 (3), in MAU 15619 (7), in MAU 15620 (13), in MAU 15621 (1), in MAU 15622 (9a), in MAU 15623 (1), in MAU 15625 (9b), in MAU 15625a (9a), in MAU 15626 (9a), in MAU 15628 (11), in MAU 15629 (4), in MAU 15630 (15), in MAU 15631 (20), in MAU 15637 (1), in MAU 15638 (13), in MAU 15640 (3), in MAU 15642 (12), in MAU 15644 (15), in MAU 15645 (7), in MAU 15647 (15), in MAU 15648 (1), in MAU 15650 A-C (20), in MAU 15816 (7), in MAU 15823 (10), in MAU 15823a (9a), in MAU 15823b (17), in MAU 16018 (7), in MAU 16019 (7), in MAU 16307 (2), in MAU 18785 (7). Richard 10 (15); Rouhan 169 (18), 175 (12), 177 (7), 178 (22), 179 (7), 180 (22), 181 (11), 187 (11), 188 (12), 190 (14), 192 (10), 193 (11), 194 (10), 195 (9a), 197 (11), 198 (9a), 199 (6), 201 (11), 202 (7), 203 (20), 204 (1), 205 (15), 206 (4), 207 (13), 208 (12), 209 (13), 210 (11), 211 (7), 212 (21), 213 (20), 214 (9b), 215 (1), 216 (3), 217 (9a), 218 (20), 219 (9b), 220 (3), 221 (21), 222 (9b), 224 (16), 225 (20), 226 (4), 227 (12), 228 (9a), 229 (1), 231 (15), 232 (13), 233 (3), 234 (15), 235 (19), 236 (11), 239 (9a), 241 (4), 242 (1), 243 (20), 244 (15), 246 (19), 247 (20), 248 (12), 249 (1), 250 (9a). Schlieben 10849 (20), 10928 in MAU 12416 (11), 10934 (7), 10934 I (20); Sieber Flora Mixta 281 (7); Staub in MAU 11348 (9a), in MAU 13012 (20). Vaughan in MAU 2566 (18), in MAU 10084 (22), in MAU 12544 (11), in MAU 12551 (12), in MAU 12593 (18), in MAU 12624 (18). Wiehe 1721 (3).