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A Synopsis of the Frullaniaceae (Marchantiophyta) from Colombia Sinopsis De La Familia Frullaniaceae (Marchantiophyta) Para Colombia

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A Synopsis of the Frullaniaceae (Marchantiophyta) from Colombia Sinopsis De La Familia Frullaniaceae (Marchantiophyta) Para Colombia BOTÁNICA-SISTEMÁTICA http://www.icn.unal.edu.co/ Caldasia 33(2):367-396.Gradstein & Uribe 2011 A SYNOPSIS OF THE FRULLANIACEAE (MARCHANTIOPHYTA) FROM COLOMBIA Sinopsis de la familia Frullaniaceae (Marchantiophyta) para Colombia ROBBERT GRADSTEIN Muséum National d’Histoire Naturelle, Dept. Systématique et Evolution, UMS 7205, Case Postale 39, 57 rue Cuvier, 75231 Paris cedex 05, France. JAIME URIBE-M. Instituto de Ciencias Naturales, Facultad de Ciencias, Universidad Nacional de Colombia, Apartado 7495, Bogotá D.C., Colombia. [email protected] ABSTRACT Frullania (Frullaniaceae) is the second-largest liverwort genus of Colombia (after Plagiochila), with 59 recorded species in six subgenera. In this paper keys are provided for the identification of 42 Frullania species reported from Colombia together with brief descriptions of their main morphological characters, distribution, habitat and elevation in the country, as well as worldwide distribution. Some species recorded from neighbouring countries, but not yet from Colombia, are also included in the keys. The remaining species reported from Colombia are considered doubtful taxa. Frullania apollinarii Steph. is a new synonym of F. lobato-hastata Steph. Key words. Colombia, elevational distribution, Frullania, identification keys, liverworts, Marchantiophyta, world distribution- RESUMEN Frullania (Frullaniaceae) es el segundo género en número de especies de Colombia (después de Plagiochila), con 59 especies en seis subgéneros. En este artículo se proveen claves para la identificación de 42 especies de Frullania registradas para Colombia, además se incluyen breves descripciones de los caracteres morfológicos diagnósticos, distribución, hábitat y gradiente altitudinal en el país, así como la distribución mundial. Algunas especies registradas de países vecinos, pero aún no registradas para Colombia son incluidas en la clave. Las demás especies registradas para Colombia se consideran taxones dudosos. Frullania apollinari Steph. es un nuevo sinónimo de F. lobato-hastata Steph. Palabras clave. Colombia, distribución altitudinal, Frullania, claves de identificación, hepáticas, Marchantiophyta, distribución mundial. INTRODUCTION et al., 2001, with updates). In Colombia, Frullania is the second-largest liverwort The liverwort family Frullaniaceae consists genus (after Plagiochila) with 59 species. of two genera, Frullania with about 300- The first records from Colombia were 350 species worldwide and Neohattoria by Alexander von Humboldt and Aimée with one species in eastern Asia (Gradstein Bonpland who collected Frullania atrata 367 Frullaniaceae from Colombia (Sw.) Nees (as Jungermannia atrata Sw.) and peruviana; Uribe, 2008) and F. mucronata F. riojaneirensis Raddi (as Jungermannia (Lehm. & Lindenb.) Lehm. & Lindenb. (= F. obscura Sw.) between Popayán and brasiliensis; Uribe & Gradstein, 2003). Almaguer, Dept. Cauca (Kunth, 1822). Gottsche (1864) reported 17 species from In this paper, keys are provided for the various localities based on specimens identification of 42 species of Frullania collected by Alexander Lindig and by José from Colombia which we have been able Jeronimo Triana. The majority of these to recognize with certainty. A few well- early collections from Colombia have not defined species known from neighbouring been re-examined critically; those studied countries but not yet from Colombia are also by Gottsche were presumably destroyed in included in the keys. For each species from 1944 with the bombing of the herbarium of Colombia a brief characterization is given of Berlin. A further 30+ species were reported its distribution, habitat and elevation in the as new to Colombia by Jack & Stephani country, its world distribution, as well the (1892), Stephani (1901-1905), Herzog (1942, main diagnostic morphological characters 1955), Bonner (1965), Robinson (1967) and separating the species from related ones. The others. Stotler (1969) in his revision of the remaining species recorded from Colombia neotropical species of Frullania subgenus are little known taxa that need more study and Thyopsiella Spruce (as subg. “Frullania”) may turn out to be synonyms; they are listed accepted 10 species for Colombia and under “Doubtful taxa” or mentioned briefly in reduced several others to synonymy. Yuzawa the discussions under the species. (1991) in his world monograph of subgenus Chonanthelia Spruce recorded 19 species The species are keyed according to the six from Colombia. subgenera to which they are traditionally assigned (Spruce, 1884; Uribe, 2008): The first edition catalogue of the liverworts subg. Chonanthelia with 17 species from and hornworts of Colombia by Gradstein Colombia, subg. Diastaloba with 2 species, & Hekking (1979) listed 58 species of subg. Frullania (= Trachycolea) with 1 Frullania for the country. This number has species, subg. Homotropantha with 1 species, remained almost unaltered in subsequent subg. Meteoriopsis with 4 species and editions (Uribe & Gradstein, 1998; Gradstein subg. Thyopsiella with 17 species. A first & Uribe, in press) in spite of intensive morphological-phylogenetic analysis of the recent collecting activities and discovery of genus Frullania (Uribe, 2008) recovered species new to the country, such as F. kunzei four of these subgenera as monophyletic s.l. (Pinzón et al., 2003) and F. dulimensis Uribe (incl. paraphyly; Hörandl & Stuessy, 2010): (Uribe, 2006). Thus, the second edition subg. Frullania, subg. Homotropantha, subg. of the catalogue lists 58 species (Uribe & Meteoriopsis and subg. Thyopsiella. The Gradstein, 1998) and the third edition 59 subgenera Chonanthelia and Diastaloba, species (Gradstein & Uribe, in press). The however, proved to be polyphyletic. Hentschel lack of increase in the number of species is et al. (2009) in a molecular-phylogenetic explained by the reduction of various species study based on a broader sampling confirmed to synonymy, thus counterbalancing the new the polyphyly of subg. Diastaloba and floristic records. Examples of recent new the monophyly of subg. Frullania and synonymy include F. atrosanguinea Tayl. subg. Homotropantha; in addition, subg. (= F. peruviana; Uribe, 2008), F. crinoidea Chonanthelia turned out to be monophyletic Spruce ex Steph. (= F. atrata; Uribe & in the latter study. Subgenus Thyopsiella, Gradstein, 2003), F. mathanii Steph. (= F. however, was recovered as two poorly 368 Gradstein & Uribe supported clades, a “temperate” clade cuencensis, dusenii, ecuadorensis, gibbosa, containing the type of subg. Thyopsiella intumenscens, kunzei, macrocephala, (F. tamarisci) and a few other species, montagnei, pittieri, setigera and a “tropical” clade which included the Andes, sometimes extending to Central majority of the Thyopsiella species and the America and Mexico: grandifolia, laxiflora, type of subg. Meteoriopsis (F. peruviana). paradoxa, peruviana, pluricarinata, Accordingly, the temperate clade was called sphaerocephala, standaertii, tetraptera, subg. Thyopsiella and the tropical clade was tunguraguana, winteri called subg. Meteoriopsis. In the present treatment, however, we have retained the Table 1. ���������Number of Frullania species recorded six subgenera of neotropical Frullania from the Departments of Colombia. as traditionally circumscribed. The new Department # Frullania species defintions of Thyopsiella and Meteoriopsis Amazonas 3 proposed by Hentschel et al. (2009) must be Antioquia 10 considered premature due to the lack of robust support for the two clades and the very limited Arauca 0 sampling of the subgenus Meteoriopsis. A Atlántico 0 reclassification of the polyphyletic subgenus Bolívar 0 Diastaloba, finally, should await the urgently Boyacá 21 needed systematic revision of this group, Caldas 4 especially of the neotropical species which Caquetá 0 have not been studied critically. Casanare 6 Cauca 11 DISTRIBUTION PATTERNS Cesar 8 Chocó 8 About half of the species of Frullania Córdoba 0 recorded from Colombia (22) have very wide Cundinamarca 26 geographical ranges and occur throughout Guainía 0 tropical America, from Mexico to southern Guaviare 0 Brazil (see below). Six of them also occur Huila 7 in the Old World tropics and are pantropical La Guajira 0 species. The remaining species (20) are largely restricted to the Andes; some of these Magdalena 10 extend to the mountains of Central America Meta 5 and Mexico and one species (F. standaertii) Nariño 4 occurs also in East Africa. Eight Andean Norte de Santander 2 species occur only in the northern Andes, from Putumayo 2 northern Peru to Venezuela (sometimes also in Quindío 13 Costa Rica) and two species (F. bogotensis, F. Risaralda 19 S a n A n d r é s y dulimensis) are only known from Colombia. 0 The geographical distribution patterns of Providencia Santander 17 the Colombian species of Frullania are as Sucre 0 follows: Pantropical: apiculata, arecae, ecklonii, Tolima 15 ericoides, nodulosa, riojaneirensis Valle del Cauca 7 Neotropical: atrata, beyrichiana, brasiliensis, Vaupés 0 caulisequa, confertiloba, convoluta, Vichada 0 369 Frullaniaceae from Colombia Northern Andes, sometimes extending Table 2. Altitudinal distribution of Colombian to Costa Rica: bicornistipula, clandestina, Frullania. formosa, jelskii, lobato-hastata, meridana, Altitudinal belt Number of species mirabilis, ringens 0-1000 m 15 Colombia only: bogotensis, dulimensis 1000-2000 m 30 2000-3000 m 25 The distribution of the species within 3000-4000 m 20 Colombia is still incompletely known. above 4000 m 8 The largest number of species is known from
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