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Thaiszia - J. Bot., Košice, 17: 1-10, 2007 http://www.bz.upjs.sk/thaiszia/index.html

T H A I S Z I AT H A I S Z I AT H A I S Z I AT H A I S Z I A JOURNAL OF JOURNAL OF JOURNAL OF JOURNAL OF BOTANYBOTANYBOTANYBOTANY

Dracoglossum, a new Neotropical fern genus (Pteridophyta)

MAARTEN J.M. CHRISTENHUSZ

Department of Biology, University of Turku, FI-20014 Turku (Finland), email: maarten.christenhusz@utu.fi, website : http://www.botanyphotos.net

CHRISTENHUSZ M.J.M. (2007): Dracoglossum, a new Neotropical fern genus (Pteridophyta). – Thaiszia – J. Bot. 17: 1-10. – ISSN 1210-0420. Abstract: The new genus Dracoglossum is proposed and described here. It is distinguished from Tectaria, based on morphology of spores and venation. Two new combinations (D. plantagineum and D. sinuatum) are established. In the first part some nomenclatural issues are clarified. Keywords: Dracoglossum, Dryopteridaceae, ferns, Neotropics, new genus, pteridophytes, Tectariaceae.

Introduction

During field studies in French Guiana and the French Lesser Antilles, and herbarium studies in Paris, it has become evident that Tectaria plantaginea (JACQ.) MAXON, constitutes a morphologically isolated taxon deserving generic rank. The purpose of this paper is to give a taxonomic account of this new genus. Traditionally, T. plantaginea was subdivided into two varieties, T. p. var. plantaginea and T. p. var. macrocarpon C.V. MORTON, based respectively on the absence or presence of the indusium (MORAN 1995, SMITH 1995). These varieties were found morphologically and geographically isolated and I believe they constitute two separate species. First the nomenclature of these species is discussed, followed by the description of the new genus, and a comparison with related genera. The appropriate new combinations are made, and their types are assigned.

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Nomenclatural notes

The nomenclature and taxonomy associated with the name Bathmium macrocarpon FÉE are elaborate, and have a history of misinterpretation. According to FÉE (1850, 1852, 1866) B. macrocarpon and Polypodium plantagineum JACQ. represent the same species. FÉE believed it was a species variable in the presence or absence of indusia. He therefore segregated Bathmium macrocarpon from Polypodium L., because Polypodium was and is still used for species lacking indusia. Instead of establishing a new combination “Bathmium plantagineum”, he gave the species a new epithet. According to the ICBN (MCNEILL et al. 2006) FÉE thus rendered his new name illegitimate, and a nomenclatural synonym of Polypodium plantagineum JACQ. The type of Bathmium macrocarpon is therefore the same as that of Polypodium plantagineum, i.e. the plate in JACQUIN (lectotype designated by PROCTOR 1977).

Firstly FÉE (1850: 9) wrote “Il y a plus: nous avons constaté que dans une même espèce l'indusium pouvait se constituer ou bien avorter. […] le Podopeltis plantaginea que Jacquin n'a pas étudié indusié, a pris place parmi les Polypodium”, referring to the variability of the presence of an indusium in this species. Secondly FÉE (1852: 287) listed Polypodium plantagineum JACQ. as a synonym of B. macrocarpon. Later, when FÉE (1866: 72) established Drynaria plantaginea (JACQ.) FÉE, he placed that taxon as “sub Bathmio macrocarpo”.

Typical Polypodium plantagineum from the Lesser Antilles is not known to have an indusium, but FÉE saw indusiate specimens from French Guiana, and therefore expanded the circumscription of the species to include these Guianese specimens. I believe however that these indusiate specimens represent a different species.

Largely thanks to these observations (FÉE 1850, 1852), the concept of Polypodium plantagineum changed from exindusiate to occasionally indusiate. Because this meant that it could not remain in Polypodium, FÉE first decided to place it in Podopeltis, but the combination ‘Podopeltis plantaginea’ was created before the genus Podopeltis FÉE was described (FÉE 1850: 9), and the combination is therefore invalid. Later FÉE (1852: 286-288) changed his mind concerning Podopeltis and restricted that genus to Aspidium singaporianum WALL. ex HOOK. & GREV. (= Tectaria singaporiana (WALL. ex HOOK. & GREV.) COPEL.) from Southeast Asia. Like Polypodium plantagineum this species has simple blades, but it is placed in Tectaria CAV. because of its echinate, winged spores, acute-tipped blades lacking apical proliferations, by having a thin lamina with readily visible venation, with the main lateral veins straight, an erect rhizome, and a persistently peltate indusium (HOOKER & GREVILLE 1831, PIGGOTT 1988).

MOORE (1858) was the first to synonymise Bathmium macrocarpon and B. sinuatum and stated that Polypodium plantagineum was the same species but lacking the indusium. His concept has been followed by METTENIUS (1859) and HOOKER (1862). However SMITH (1875) treated B. sinuatum as separate from P. plantagineum.

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MORTON (1966) considered the three names Polypodium plantagineum, Bathmium macrocarpon and B. sinuatum as distinct, but like SMITH (1875), he recognised two species, and treated P. plantagineum as distinct from B. macrocarpon, doubtfully placing B. sinuatum in its synonymy. Taking FÉE's B. macrocarpa as a legitimate basionym, he attempted to establish a new combination: “Tectaria plantaginea (Jacq.) Maxon var. macrocarpa (FÉE) C.V.MORTON”. Nevertheless, according to art. 58 of the ICBN (MCNEILL et al. 2006), instead of a new combination, he unintentionally established a new name. Because he used an available epithet of an illegitimate name, the name should be cited as: Tectaria plantaginea (JACQ.) MAXON var. macrocarpa C.V.MORTON, without the reference to FÉE. Its type is the specimen “Poiteau s.n. (P)”, which was originally cited by FÉE as a specimen under B. macrocarpon. In the Paris herbarium there are two sheets labelled “de la Guyane par Poiteau 1825”. They are originally from the herbarium of Caen (CN), and were integrated into P in 1974. Since MORTON did not study any original Poiteau material, the holotype of MORTON’s T. p. var. macrocarpa is uncertain. It is also not sure whether Fée studied these specimens. The Fée herbarium was at Strassbourg (STR), so neither specimen can be considered to be the holotype.

Systematics

Dracoglossum CHRISTENH., gen. nov. Rhizomata angustis repentes breviter, paleis lanceolatis, clathratis, atrofuscis.

Laminae simplices, coriacae, emarginatae, ad apices saepe gemmiferae. Sori indusiati vel exindusiati, seriales duobus inter costulas. A Tectaria et Hypoderris spores teretibus (non cristato-echinatis) et alis vadosibus imprimis differt.

Typus . — Dracoglossum plantagineum (JACQ.) CHRISTENH.

Description

Evergreen, hemicryptophyte ferns, with short-creeping rhizomes, that are 3-7 mm thick, clothed in entire, lanceolate, clathrate, blackish-brown scales to 4 mm long. Scales borne on stipes and rachises abaxially, also with short hairs along stipes. Blades simple, leathery, lanceate to ovate lanceolate, margin entire to broadly crenate, to 60 cm long, the bases acute, the apices gradually acute to acuminate, mostly emarginate at tip and there bearing a proliferous bud, which may develop into a new plant when the blade touches the substrate. Tissue leathery, dark, dull green, opaque. Venation obscure, reticulate, with four or five rows of areoles between the costules, and mostly each areole with an included veinlet. Sori round to ovate, usually about 2 mm in diameter, placed (irregularly) in two rows between the main lateral veins, sometimes with a (basal) pair of sori merging into an irregularly elongately shaped sorus to 5 mm long, sori exindusiate or indusiate; indusia peltate when present. Spores shallowly ridged, surface smooth, (Fig. 1A), not crested or winged, neither spinulose nor reticulate-echinate (as in Tectaria, Fig. 1C-F).

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Fig. 1. Spores ×1000. A = Dracoglossum plantagineum; B = Hypoderris brownii; C = Tectaria prolifera; D = Tectaria incisa; E = Tectaria heracleifolia; F = Tectaria panamensis. Reproduced from Tryon & Tryon (1982: 479-483, fig s. 69 & 70 pro parte), with kind permission of Springer Science and Business Media.

Distribution and ecology

The genus Dracoglossum is restricted to the Neotropics (Fig. 2). They usually grow on rocky or clayey substrate in or along streams, in shaded gullies, or occasionally terrestrial in the shade of evergreen lowland or lower montane forests. It is mostly found at lower to middle elevations, 10-800(-1000) m.

Comparison

Dracoglossum differs from Tectaria, the genus in which the species were previously placed, primarily by the spore morphology, which is smooth and only shallowly ridged (Fig.1A). Tectaria has spores with deeply wing-like folds or many smaller crests and spinulose, perforate or cristate-echinate surfaces (Fig. 1C-F). Dracoglossum is further distinguished by the simple, leathery blades with obscure tertiary venation. Most Tectaria have pinnate blades, and the few species with simple blades are either lobed and/or show the tertiary venation readily. Examples of simple bladed species are Tectaria panamenis (HOOK.)

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R.M.TRYON & A.F.TRYON, and T. singaporiana, but both species lack the apical proliferations. Of the simple-bladed species only Tectaria prolifera (HOOK.) R.M.TRYON & A.F.TRYON bears proliferations, but the blades are very thin and dimorphic: the fertile leaves erect, bearing sori that are borne in two rows, that are covered by lunate indusia. All of these species have echinate spores with wing-like folds (Fig. 1C-F).

Fig. 2. Global distribution of Dracoglossum.

First molecular estimates segregate Dracoglossum from Tectaria (E.

Schuettpelz, pers. comm.), and therefore maintaining Tectaria in the traditional sense would render it polyphyletic. Further molecular sampling of the Tectariaceae, and molecular studies to assess the placement of Dracoglossum in the overall fern classification (SMITH et al. 2006), will be needed, but is not within the scope of this paper.

Morphologically, Dracoglossum appears to be closest to Hypoderris R.BR. ex HOOK. (Tectariaceae), which also has leathery blades, and an obscure tertiary venation, but Hypoderris differs in the prominently echinate spores (Fig. 1B), lobed blades, and the absence of the apical proliferations.

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The two species of Dracoglossum were previously placed in various genera: Aspidium SW., and Bathmium C.PRESL ex LINK are superfluous names for Tectaria. Podopeltis FÉE, and Dryomenis FÉE ex J.SM. are later synonyms of Tectaria, none of which have our species as type. Drynaria (BORY) J.SM. is an old world genus of Polypodiaceae.

Fig. 3. Dracoglossum plantagineum; photograph taken near Étang Madère, Guadeloupe, French West Indies, 13 March 2005; vouc her: Christenhusz 4065 (TUR, UC).

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Key to the species of Dracoglossum

1. Indusia absent or very reduced, if present leaving most of the sporangia uncovered; blades usually broadly (ovate-) lanceolate, the margins variable, entire to irregularly lobed (Mexico to Panama, Puerto Rico, Lesser Antilles, Trinidad, Tobago, Colombia, Venezuela, Ecuador, Peru, Bolivia, Fig. 2)

D. plantagineum 2. Indusia present, sometimes caducous on dried material, covering the

sporangia, blades usually lanceate to lanceolate, the margins usually sinuate or entire (Venezuela, Guyana, Suriname, French Guiana, northern Brazil, coastal Ecuador, Fig. 2)

D. sinuatum

1. Dracoglossum plantagineum (JACQ.) CHRISTENH., comb. nov. (Fig. 3)

Polypodium plantagineum JACQ., Collectanea 2: 104, tab. 3, fig. 1 (1788, publ. 1789). — Drynaria plantaginea (JACQ.) J.SM., J. Bot. (Hooker) 4: 61 (1841). — Podopeltis plantaginea (JACQ.) FÉE, Mém. Soc. Mus. Hist. Nat. Strasbourg 4 (1): 9 (1850), nom. nud. sub inval. nom. gen. — Bathmium macrocarpon FÉE, Mém. Foug. 5. Gen. Filic.: 288 (1852), nom. illeg. sensu stricto. — Dryomenis plantaginea (JACQ.) J.SM. in: SEEMANN B. C., Bot. Voy. Herald 229 (1854). — Aspidium plantagineum (JACQ.) GRISEB., Abh. Königl. Ges. Wiss. Göttingen 7: 286 (1857). — Bathmium plantagineum (JACQ.) E.FOURN., Bull. Soc. Bot. France 19: 254 (1872, publ. 1873). — Tectaria plantaginea (JACQ.) MAXON, Contr. U.S. Natl. Herb.10: 494 (1908). — Lectotype: Plate in JACQUIN, Collectanea 2: 104, tab. 3, fig. 1 (1788, publ. 1789), designated by Proctor 1977.

Note: According to TRYON & STOLZE (1991), the holotype of Polypodium plantagineum is probably Jacquin s.n., from Martinique, and presumably located in W or BM, but they did not see a specimen. I have also not been able to locate it. Two historical collections that must have been accessible to Jacquin are present at Paris, namely Tournefort 5365 (P-TRF-00307022!) and Vaillant s.n. (P-00398315!).

Polypodium latifolium VAHL, Eclog. Amer. 3: 50 (1807), nom. illeg. non Polypodium latifolium G.FORST. (1786). — Type: Montserrat, Ryan s.n. (holo-, C!).

Tectaria plantaginea (JACQ.) MAXON var. confluens C.V.MORTON, Amer. Fern J. 56: 123 (1966), syn. nov. — Type: Trinidad, 13.III.1921, L.H. Bailey & E.Z. Bailey T-15 (holo-, US no. 1058555), not A.C. Smith 10270 (US) as stated in PROCTOR (1977).

Note: Tectaria plantaginea var. confluens, originally described because of its confluent sori near the midrib, is within the variability of D. plantagineum (TRYON & STOLZE 1991). In my opinion this variety does not warrant recognition.

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SPECIMENS EXAMINED. — Costa Rica. Alajuela, Upala, Dos Rios, 10.X.1987, Herrera 1033 (AAU!, CR). — Heredía, La Selva, 17.IV.2002, Jones 385 (CR, TUR!). — La Selva, 5.VII.2002, Jones 512 (CR, TUR!). — Limón, Parque Nacional Tortuguero, Lomas de Sierpe, 15.VIII.1988, Robles 2061 (AAU!, CR). — Zent, VIII.1901, Tonduz 14556 (P-00518453!). — Panama. Isla Taboga, VII.1867, Le Jolis s.n. (P-00518454!). — San Blas, Cordillera de San Blas, 12.III.1993, Herrera 1330 (AAU!, MO). — Puerto Rico. Utuado, 12.III.1887, Sintenis 6431 (P-00518452!). — Guadeloupe. Grand-Étang, 9.II.1936, Allorge 40 (P-00518433!, P-00518434!). — Grand-Étang, 6.IV.2002, Christenhusz 2793 (TUR!, U!). — Étang Madère, 13.III.2005, Christenhusz 4065 (TUR!, UC!). — Bois des Bains Jaunes, Duss 767 (P-00518429!). — Forêt de Fumée, 22.II.1936, Feldmann s.n. (P-00518430!). — Forêt des Palmistes, near Gourbeyre, 3.III.1961, Le Gallo 3074 (P-00518428!). — Morne Coco, XII.1867, Hahn 25 (TUR!) —Dominica. Emerald Pool, 10.IV.2003, Christenhusz 2837 (TUR!). — in sylvis humidis prope Laudet, XI.1881, Eggers 509 (P-00518446!, P-00518447!, P-00518448!, P-00518449!, P-00518450!, P-00518451!). — Martinique. Morne de la Calebasse, II.1854, Bélanger 820 (P-00518436!, P-00518437!). — Morne de la Calebasse, 19.III.2003, Christenhusz 2684 (TUR!, U!). — Bois du camp de l'Alma, 1884, Duss 1570 (P-00518438!). — Rivière Blanche, N 3, Fort-de-France vers Morne Rouge, 550 m, 30.XII.1989, Cremers 11149 (CAY!). — Grenada. Azimar mountain woods, 18.XI.1905, Broadway s.n. (P-00518440!, P-00518441!). — Trinidad and Tobago. Trinidad, in sylvis ad Arima, 16.III.1885, Eggers s.n. (P-00518444!, P-00518445!). — Tobago, the Forest Reserve, 16.III.1911, Broadway 4230 (P-00518442!, P-00518443!). — Colombia. Campanero, XI.1843, Funck 781 (P-00518455!). — Ecuador. Napo, Parque Nacional Yasuní, 14.IV.1996, Moran 6173 (AAU!, QCA, UC, TUR!). — 18.IV.1997, Tuomisto 10649 (QCA, QCNE, TUR!) — Napo, Parque Nacional Yasuní, 20.IV.1996, Moran 6250 (AAU!, QCA, UC, TUR!). — Sucumbios, trail at Río Ushaué, 29.I.1992, Øllgaard 99712 (AAU!, QCA, TUR!). — Peru. Dept. Cuzco. Prov. Cuzco, Dist. Camisea. Campamento San Martin-C, Camisea Production Unit, S and SW of Camp 11, 24.I.1997, Acevedo-Rodriguez 9046 (SP!, US). — Dept. San Martín, near Tarapoto, Cerro Guayrapurina, I.1856, Spruce 4648 (P-00518439!).

2. Dracoglossum sinuatum (FÉE) CHRISTENH., comb. nov.

Bathmium sinuatum FÉE, Mém. Foug. 5. Gen. Filic.: 288 (1852). — Aspidium sinuatum (FÉE) T.MOORE, Index Fil. 104 (1858), nom. illeg. non Aspidium sinuatum LABILL. (1824) nec Aspidium sinuatum GAUDICH. (1828). — Podopeltis sinuata (FÉE) J.SM., Hist. Fil. 199 (1875) — Type: “Habitat in Guyana. (Leprieur.) V. S. in Herb. Moug.”, not found. Neotype (here designated): French Guiana, Montagnes Bellevue de l'Inini, Bassin de l’Inini, 15.VII.1985, G. Cremers & L. Allorge 8909 (neo-, P-00518417!; isoneo, B, BR, CAY!, G, MG, MO, U, UC, Z).

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Note: The herbarium of Mougeot has been partly integrated in MPU and P, but the original material was not located. I therefore decide to select a neotype.

Bathmium macrocarpon FÉE, Mém. Foug. 5. Gen. Filic.: 288 (1852), nom. illeg., pro parte quoad Poiteau s.n., Guyane française, Cayenne (as a cited specimen).

Tectaria plantaginea (JACQ.) MAXON var. macrocarpa C.V.MORTON, Amer. Fern J. 56: 123 (1966). — Type: French Guiana, Cayenne, Poiteau s.n.,1825 (holo-, P?, not found; iso-, P-00398318!, P-00398319!).

SPECIMENS EXAMINED. — Guyana. Potaro-Siparuni, Chenapou, amerindian village (Patumona), 50 km upstream Kaieteur Falls, 15.X.1987, Kvist 269 (AAU!, CAY!, BRG, P-00518426!, US). — Suriname. Brownsberg, 5.III.2003, Christenhusz 2554 (TUR!, U!, UC!). — Brownsberg summit, 25.VI.1924, Boschwezen (Forestry Bureau) BW 6540 (P-00518425!, U). — Inselberg Talouakem, Monts Tumuc-Humac, 12.VIII.1993, De Granville 12259 (BBS, CAY!, NY, P-00518424!, US, Z). — French Guiana. Sentier Botanique de la Réserve Volontaire Trésor, Montagne de Kaw, 20.II.2003, Christenhusz 2429 (CAY!, NY!, TUR!). — Cacao, sentier Molokoï, 23.II.2003, Christenhusz 2503 (CAY!, TUR!, U!). — Route de l’Est, km 73, Layon ONF, Bassin de l’Orapu, 10.V.1979, Cremers 5573 (CAY-060143!, NY, P-00398246!, U, Z). — Montagnes Bellevue de l'Inini, Bassin de l’Inini, 31.VIII.1985, Cremers 9204 (CAY!, P-00518419!, U). — Piste de Bélizon, Montagne Tortue, Bassin de la Comté, 2.VI.1988, Cremers 10052 (CAY-060124!, P-00398247!, UC, US, Z). — Station des Nouragues, Bassin de l’Approuague, 17.XI.1989, Cremers 10832 (B, CAY-060176!, COL, HAMAB, MG, MPU, NY, P-00518418!, SJPR, U, UC, US, VEN, Z). — D.Z. du Haut-Kourcibo, Bassin du Sinnamary, 17.IV.1991, De Granville 11293 (B, BR, BBS, CAY-009111!, G, INPA, K, MG, NY, P-00398248!, U, UC, US). — Montagnes de la Trinité, zone sud, Bassin de la Mana, 11.I.1998, De Granville 13497 (B, CAY!, NY, P-00518421!, U). — Pic Matécho, zone sommitale, Région de Saül, 10.IX.2000, Mori 25092 (CAY!, NY, P-00518422!). — Saut Pararé, Rivière Arataï, Bassin de l’Approuague, 1.IX.1977, Sastre 5856 (CAY!, P-00518423!, U). — Brazil. Prope San Gabriel da Cachoeira ad Rio Negro, 1852, Spruce 2189 (P-00518413!, P-00518414!). — Ecuador. Esmeraldas, Road Lita - San Lorenzo, km 46.7, 3.XI.1994, Øllgaard 15356 (AAU!, QCA). — Esmeraldas, San Miguel, Rio Cayapas, 20.XI.1992, Tipaz 2278 (AAU!, MO).

Acknowledgements

I hereby thank Michel Boudrie, Georges Cremers, Jacques Florence, Michael Kessler, Jefferson Prado, George R. Proctor, Eric Schuettpelz, and Alan R. Smith for their constructive comments. Special thanks go to Frédéric Tronchet, who helped me to solve some of the nomenclatural problems. Further, I am grateful to Sarah Bollendorff, Frank Katzer, Mikko Paajanen, Kalle Ruokolainen and Hanna Tuomisto for the great company and help during field work, and to

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the staff members of AAU, CAY, GOET, P, SP, and TUR for allowing study of their specimens. Financial support to the first author for the herbarium studies in P was provided by SYNTHESYS, and field work was funded by grants to Hanna Tuomisto from the Academy of Finland.

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Received: January 11th 2007 Revised: June 11th 2007 Accepted: June 11th 2007