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Three New Species of Scaly Tree Ferns (Cyathea-Cyatheaceae) from the Northern Andes

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Three New Species of Scaly Tree Ferns (Cyathea-Cyatheaceae) from the Northern Andes Phytotaxa 1: 43–56 (2009) ISSN 1179-3155 (print edition) www.mapress.com/phytotaxa/ Article PHYTOTAXA Copyright © 2009 • Magnolia Press ISSN 1179-3163 (online edition) Three new species of scaly tree ferns (Cyathea-Cyatheaceae) from the northern Andes MARCUS LEHNERT Staatliches Museum für Naturkunde Stuttgart, Abt. Botanik, Am Löwentor, Rosenstein 1, 70191 Stuttgart, Germany; email: [email protected], [email protected] Abstract Three species of the genus Cyathea (Cyatheaceae-Polypodiopsida) are described as new to science from the northern Andes: Cyathea aemula from southern Colombia and northern Ecuador, C. ars from southeastern Ecuador, and C. guentheriana from northeastern Ecuador. All species are illustrated and compared to their putative closest relatives. Key words: Andean cordillera, biodiversity, Colombia, Cyathea, Ecuador, ferns, pteridophytes Introduction The small Andean republic of Ecuador holds an outstanding position in the global species count, with 15,306 native species and an estimated endemism of 27.3 % in vascular plants (Jørgensen & León-Yánez 1999). The exceptionally high number of species compared to the small size of the country is due to its varied relief and the contact of several biogeographic units (Valencia et al. 1994), which are mainly caused by different precipitation regimes along the Andes (Killeen et al. 2007). The Andean cordillera effectively separates a western and an eastern lowland rainforest biome (Gentry 1982). Despite the relatively short distance, an exchange of species between the western and eastern Andean slopes is restricted, e.g. about 25% of the pteridophyte species found on the western side do not occur on the eastern side (Moran 1995a). The botanical explorations of this country still yield new discoveries of unknown species, including sizeable and conspicuous plants like the scaly tree ferns of the genus Cyathea (Moran 1991, 1995b, León & Moran 1996, Moran & Øllgaard 1998). The checklist of Ecuador lists 51 native species of Cyathea (Jørgensen & León-Yánez 1999) but several new discoveries increased the total count to 64 species in the meantime (Lehnert 2003, 2006a, 2006b, 2008, 2009). The family Cyatheaceae contains ca. 600 species with a pantropical-southern temperate distribution (Smith et al. 2006). Its members differ from other families of the tree fern alliance by having not only the common pluricellular hairs, but also various types of scales in their indument (Kramer & Green 1990). The family includes the tallest living ferns, whose erect trunk-like rhizomes can reach over 20 m tall. Due to their size they are less frequently and more fragmentarily collected than other fern groups (Janssen 2006). The taxonomic status, geographical range and ecological potential of many poorly understood species are just beginning to clarify (e.g. Lehnert 2005, 2008, Moran et al. 2008) thanks to an increased number of long-term field explorations. The generic subdivision of the family was long disputed (Holttum & Tryon 1977). Holttum (1963) recognized only one universal genus Cyathea with the two subgenera Cyathea and Sphaeropteris, including several sections. Tryon (1970) recognized six genera (Cyathea, Cnemidaria, Trichipteris, Alsophila, Nephelea, Sphaeropteris) based on studies focused on Neotropical species. Two of these genera, Trichipteris Accepted by M. Christenhusz: 7 Oct. 2009; published: 20 Oct. 2009 43 and Nephelea, were based on feeble morphological characters and were soon merged in Cyathea (Lellinger 1987) and Alsophila (Conant 1983), respectively. Holttum & Edwards (1983) acknowledged the remaining four genera of Tryon’s classification and treated Cyathea, Cnemidaria, and Alsophila as subsections of the section Cyathea within the subgenus Cyathea, which further included section Gymnosphaera. Phylogenetic investigations (Conant et al. 1995, 1996, Korall et al. 2006, 2007, Janssen et al. 2008, Moran et al. 2008) have helped with the interpretation of the importance of various characters that are used in the definition of genera and species. Most anatomy-based changes of the generic concepts (e.g. Lellinger 1987) were corroborated by the molecular data but some supported clades remain problematic in their taxomonic definition, e.g. the separation of the monophyletic genera Gymnosphaera and Alsophila (Korall et al. 2007). The genus Cyathea in its strict sense (Conant & Stein 2001) includes the genera Cnemidaria (Stolze 1974), Trichipteris (Barrington 1978, Lellinger 1987) and parts of the genus Sphaeropteris sensu Tryon (1970; Windisch 1977, 1978, Kramer 1978, Lellinger 1984). Evidently a small group of nine paleotropical species (Holttum 1982) stands basal to the rest of the genus Cyathea, which further contains 184 species currently recognized in the Neotropics. This number does not yet include the eight species of the genus Hymenophyllopsis, which apparently are just highly specialised species of Cyathea (Korall et al. 2006, 2007). This concept of Cyathea includes all species of Cyatheaceae that have a differentiated petiole scale margins consisting of cells smaller and of different orientation than those of the scale body and simultaneously lack dark marginal setae. The latter can be found in the other genera in combination with (Alsophila, Gymnosphaera) or without (Sphaeropteris) a differentiated scale margin. Besides their conspicuous cell wall thickening, setae can be distinguished from marginal teeth by their insertion into the supporting cell layers. Setae originate from the scale body and are each supported by one to many cells, which can be best seen in the enlarged apical setae of the genus Alsophila (Conant 1983). In the genus Sphaeropteris, the marginal setae sit directly on the scale body and can be recognized as the end points of simple parallel cell rows originating from the scale base. Opposed to this, marginal teeth, which may occur in Cyathea, are the end points of branching rows of cells that decrease in size. Consequently, several marginal teeth may sit on one supporting cell. This concept allows the true classification of species of Cyathea that previously had been erroneously placed in Sphaeropteris. Such species like Cyathea poeppigii (Hooker 1844: 43) Domin (1929: 263) have the petiole scale margins in most parts reduced to only one row of conspicuous marginal teeth that may be mistaken for setae. The new species described here were discovered during ongoing efforts towards a complete revision of the whole genus on the morphological and molecular level. Material studied for this part came mainly from the collections of GOET, MO, QCA and UC (following: Holmgren et al.1990). The terminology in this article follows principally Lellinger (2002). However, for some special tree fern characters established terms are preferably used. These include terms for the coloration of the petiole scales and shape of the indusia. “Discordantly bicolorous” refers to petiole scales with a differentiated margin of smaller cells in which the marginal color extends into the scale body, which has a different (usually darker) color. “Concordantly bicolorous” means that the structural border between scale margin and body is iterated by the coloration. The indusia may be absent (exindusiate) or cover the sori completely (sphaeropteroid, i.e. globose) or partially, and this by covering all sides of the receptacle equally (in descending order of extension: subsphaeropteroid, urceolate, cyatheoid, meniscoid, discoid) or from only one side (hemitelioid). For the relative position of the sori between the segment margins and the midveins, the terms “marginal”, “medial”, and “costal” are usually used. The term “costa” means midvein but in the pteridological terminology, it is generally applied to the main axis of the pinnae. Since the traditional term “costal” is not self-explaining, it is here replaced by the general term “proximal.” 44 • Phytotaxa 1 © 2009 Magnolia Press LEHNERT Systematic treatment Cyathea aemula Lehnert, sp. nov. (Figs. 1A, 2) Species generis Cyatheae exindusiata apicibus frondorum abrupte vel gradualiter terminantibus, squamis discordanter bicoloratis, paraphysibusque longis; a Cyathea ulei paraphysibus longioribus, furfure petiolorum albicante densiore plusque persitentiore (vs. Cyathea ulei furfure brunneo, fugaci vel absente), pinnulis fertilibus incisis plusquam medio spatio usque ad costulis (vs. non plusquam medio spatio) praestans; in pinnulis longe petiolatis similis Cyatheam kalbreyeri vel Cyatheam divergentem, sed ab hac in paraphyibus longis squamisque petiolorum bicoloratis (vs. paraphysibus brevibus squamisque concoloratis Cyatheae kalbreyeri), ab illa in absentia indusiorum (vs. Cyathea divergens cum indusiis sphaeropteroideis instructa) differt. Type:—ECUADOR: Pastaza: Mera Cantón, 2 km NW of Mera, Campamento Vacacional Evangelico Mangayacu, up the ridge behind the Campamento, 01°26'00"S, 78°07'30"W, 1350–1500 m, 29 Jul 1992, Fay & Fay 3781 (holotype UC, isotype MO). FIGURE 1. Habit of Cyathea species in the field. A. Cyathea aemula, trunk apex, Ecuador, Prov. Napo (Photo J. Homeier). B. Cyathea ulei, trunk apex, Ecuador, Prov. Loja (Photo M. Lehnert). Trunks slender, to 7 m tall (but commonly smaller), straight or only basally ascending, without adventitious buds or old petioles, fronds shed cleanly; trunk apices hidden in fascicles of petioles of green fronds, consisting of two to four pseudowhorls so that the fascicles appear stretched (Fig. 1A). Fronds to 235 cm long, arching to drooping, appearing spirally arranged
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