Bischler etal Marchantiidae FN

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New York Botanical Garden Press Flora Neotropica Monograph 97: 1-262. 2005

Marchantiidae by

Helene Bischler-Causse, S. Robbert Gradstein, Suzanne Jovet-Ast, David G. Long and Noris Salazar Allen

Bischler et al. 2

CONTENTS Abstract.................................................................................................................................................. 3 Resumen................................................................................................................................................. 4 Introduction............................................................................................................................................ 6 Historical Survey ................................................................................................................................... 8 Morphology and Anatomy of Marchantiales and Monocleales............................................................. 9 Ploidy level, Population Structure, and Speciation.............................................................................. 14 Ecology, Life History Traits, and Dispersal ........................................................................................ 15 Distribution and Altitudinal Zonation.................................................................................................. 17 Phylogeny ............................................................................................................................................ 19 Systematic Treatment........................................................................................................................... 20 Conspectus ........................................................................................................................................... 20 Key 2. Key to the Neotropical Families of Marchantiales................................................................... 22 Key 3. Key to the Neotropical Genera of Marchantiales ..................................................................... 25 Marchantiales....................................................................................................................................... 29 Monocleales ....................................................................................................................................... 199 Sphaerocarpales ................................................................................................................................. 205 Doubtful , Excluded, or Erroneously reported taxa ........................................................................... 208 Acknowledgments.............................................................................................................................. 209 Literature cited ................................................................................................................................... 211 Index of Scientific Names.................................................................................................................. 227 (captions) ........................................................................................................................................... 232 Helène Bischler-Causse ..................................................................................................................... 252 Stephan Robbert Gradstein ................................................................................................................ 253 David Geoffrey Long......................................................................................................................... 255

Bischler et al. 3

ABSTRACT

Bischler-Causse, Helene (Laboratoire de Cryptogamie, Muséum National d’Histoire Naturelle, 12 rue Buffon, 75005 Paris, France), Gradstein, S. Robbert (Institute of Plant Sciences, University of Göttingen, Untere Karspüle 2, 37073 Göttingen, Germany), Jovet-Ast, Suzanne (Laboratoire de Cryptogamie, Muséum National d’Histoire Naturelle, 12, rue Buffon, 75005 Paris, France. Present address: Résidence Château d’Arcadie, 64200 Biarritz, France), Long, David G. (Royal Botanic Garden, Edinburgh EH3 5LR, U. K), and Salazar Allen, Noris (University of Panama, Department of Botany; Smithsonian Tropical Research Institute, Apartado 2072, Balboa, Ancón, Republic of Panama). Marchantiidae. Flora Neotropica Monograph xx: 1-xxx. 200x. — The liverworts or Hepaticae are traditionally subdivided into two subclasses, Jungermanniidae (4-5 orders) and Marchantiidae (3 orders). This monograph of the neotropical Marchantiidae treats 103 species, in two orders Marchantiales and Monocleales. Marchantiales are represented in the Neotropics by 102 species, in 18 genera, and 11 families. The majority of the genera have only few species: fourteen genera comprise less than 6 species, three genera 8-10 species. More than half of the neotropical species of Marchantiales belong to the genus Riccia (53 species). Monocleales are represented in the Neotropics by a single species. The third order of the Marchantiidae, Sphaerocarpales, is not yet known with certainty from the Neotropics but might be expected to occur in the region. All species of Marchantiidae are terrestrial or saxicolous and they often grow in harsh, disturbed biotopes. Many are able to withstand strong illumination and drastic diurnal temperature changes; drought-tolerance of gametophytes and spores is common. Gametophytes may be able to resume growth after months of drought and spores may take advantage of short favourable conditions to germinate, producing fast-maturing plants with sporophytes. Twenty-six species have wide, transoceanic ranges and occur in several continents. About half of these are widely distributed within the Neotropics, the others, preferring a Mediterranean-type of climate, are confined to the northern, or to the northern and the southern borders of the area and are absent from the tropical zone. The remaining species (76) and one genus of Marchantiidae, Cronisia, are restricted to tropical America. Twenty-six of these may qualify as endemics, having very restricted distributions within the Neotropics. The endemics are found in all areas except on the Guyana shield and on the smaller West Indian islands. More than half of the endemic species (18) belong to the genus Riccia. One new endemic species is proposed: Mannia hegewaldii. Highest species diversity is observed in areas with dry seasons, e.g., dry parts of Mexico and Central America (64 species) and the lowlands of Brazil and Argentina (65 species). Less droughttolerant species are distributed predominantly in the Andes (47 species). In contrast to leafy

Bischler et al. 4 liverworts and mosses, species diversity is higher in the lowlands than in the mountains. Eighty-three species are found in the lowland, 56 in the lower montane, 42 in the upper montane, and 15 in the subalpine and alpine belts. The classifications of the Marchantiidae, including the one in current use, are usually based primarily on gametangial characteristics. Two main evolutionary hypotheses were postulated in the past, 1) the progressive specialisation hypothesis, postulating the evolution of plants with archegoniophores from plants without them, and 2) the progressive reduction hypothesis, postulating evolution in the opposite direction (development of plants without archegoniophores from those having them). Interestingly, phylogenetic analysis using structural data (morphology, anatomy, development) lends support to the progressive specialisation hypothesis, with Monocleales and Sphaerocarpales being sister groups to the monophyletic Marchantiales, but molecular analysis supports the progressive reduction theory. Monocleales and Sphaerocarpales are unequivocally nested within the Marchantiales in the molecular tree. It appears that the traditionally recognised orders and families of the Marchantiidae as well as the delimitation of some of the genera in this group are not supported by molecular data.

RESUMEN

Bischler-Causse, Helène (Laboratoire de Cryptogamie, Muséum National d’Histoire Naturelle, 12 rue Buffon, 75005 Paris, France), Gradstein, S. Robbert (Institute of Plant Sciences, University of Göttingen, Untere Karspüle 2, 37073 Göttingen, Germany), Jovet-Ast, Suzanne (Laboratoire de Cryptogamie, Muséum National d’Histoire Naturelle, 12, rue Buffon, 75005 Paris, France, o Résidence Château d’Arcadie, 64200 Biarritz, France), Long, David G. (Royal Botanic Garden, Edinburgh EH3 5LR, U. K) y Salazar Allen, Noris (University of Panama, Department of Botany; Smithsonian Tropical Research Institute, Apartado 2072, Balboa, Ancón, Republic of Panama). Marchantiidae. Flora Neotropica Monograph xx: 1-xxx. 200x. — Las hepáticas o Hepaticae se subdividen, tradicionalmente, en dos subclases, Jungermaniidae (4-5 órdenes) y la Marchantiidae (3 órdenes). Esta monografía de la Marchantiidae neotropical trata 103 especies en dos órdenes, Marchantiales y Monocleales. Los Marchantiales están representados en el Neotrópico por 103 especies en 18 géneros, 11 familias y cuatro subórdenes. La mayoría de los géneros tienen pocas especies: catorce géneros incluyen menos de 6 especies, tres géneros, 8-10 especies. Más de la mitad de las especies neotropicales de Marchantiales pertenecen al género Riccia (53 especies). Los Monocleales están representados en el Neotrópico por una sola especie. El tercer orden de la Marchantiidae, Sphaerocarpales, no se conoce con certeza para el Neotrópico pero se puede esperar que esté presente en la región. Todas las especies de Marchantiidae son terrestres o saxícolas y a menudo crecen en

Bischler et al. 5 biotopos perturbados y difíciles. Muchas son capaces de soportar una fuerte iluminación y cambios diurnos drásticos de temperatura; es común la tolerancia a la desecación de los gametofitos y esporas. Sin embargo, ya que la fertilización requiere agua, todos los taxa son, al menos temporalmente, higrófitos. Los gametofitos pueden reasumir su crecimiento después de meses de sequía y las esporas pueden tomar ventaja de condiciones favorables de corto tiempo para germinar, produciendo plantas que maduran rápido con esporofitos. Veintiséis especies tienen rangos de distribución transoceánicos extensos, y se presentan en varios continentes. Cerca de la mitad de éstas se distribuyen ampliamente en el Neotrópico, las otras, que prefieren un tipo de clima mediterráneo, están confinadas al borde norte o norte y sur del área y están ausentes de la zona tropical. El resto de las especies (76) y un género de Marchantiidae, Cronisia, están restringidas a la América tropical. Veintiséis de éstas pueden calificar como endémicas con distribuciones muy restringidas en el Neotrópico. Las endémicas se encuentran en todas las áreas excepto en el macizo de la Guyana y en las pequeñas islas de las Indias Occidentales. Más de la mitad de las especies endémicas (18) pertenecen al genero Riccia. Se propone una nueva especie endémica: Mannnia hegewaldii. Se observa una alta diversidad de especies en áreas con estaciones secas, e.g., partes secas de América Central (64 especies) y las tierras bajas de Brasil y Argentina (65 especies). Las especies menos tolerantes a la desecación están distribuidas predominantemente en los Andes (47 especies). En contraste con las hepáticas foliosas y los musgos, la diversidad de especies es mayor en las tierras bajas que en las montañas. Ochenta y tres especies se encuentran en las tierras bajas, 56 en zonas montano bajas, 42 en las montano altas y 15 en las fajas subalpina y alpina. Las clasificaciones de la Marchantiidae, incluyendo la de uso actual, usualmente se basan en características gametangiales. En el pasado, se postularon dos hipótesis evolutivas principales, 1) la hipótesis de la especialización progresiva, que postula la evolución de las plantas con arquegonióforos a partir de plantas sin ellos, y 2) la hipótesis de la reducción progresiva, que postula la evolución en la dirección opuesta (el desarrollo de plantas sin arquegonióforos a partir de aquéllas que los tienen). Es interesante que el análisis filogenético utilizando datos estructurales (morfología, anatomía y desarrollo) apoya la hipótesis de la especialización progresiva, con los Monocleales y Sphaerocarpales como el grupo hermano de los Marchantiales monofiléticos, pero el análisis molecular apoya la hipótesis de reducción progresiva. Los Monocleales y Sphaerocarpales están inequívocamente anidados dentro de los Marchantiales en el árbol molecular. Parece que los órdenes, subórdenes y familias de la Marchantiidae reconocidos tradicionalmente, así como la delimitación de algunos géneros en este grupo no son apoyados por los datos moleculares.

Bischler et al. 6

INTRODUCTION

The liverworts are divided into seven orders: Jungermanniales Klinggr. and Calobryales Campb. (the leafy liverworts); Treubiales Schljak., Metzgeriales Chalaud, Sphaerocarpales Cav. and Monocleales Schust. (the simple thalloid forms); and Marchantiales Limpr. (comprising the forms with complex thalli). Based on morphological and ontogenetic characteristics, the orders Marchantiales, Monocleales and Sphaerocarpales have been grouped into the subclass Marchantiidae Schiffn., the Jungermanniales, Calobryales, Treubiales and Metzgeriales into the subclass Jungermanniidae Schiffn. The subdivision of the liverworts into two subclasses is also supported by molecular data (rRNA 18S and 26S genes, Capesius & Bopp, 1997; Hedderson et al., 1996; chloroplast gene rbcL, Lewis et al., 1997). Many morphological and, especially, ontognetic features separate the two subclasses. Marchantiidae are characterized by cuneate apical cells with four cutting faces, specialized cells containing a single, large oil-body, antheridial ontogeny with a four-celled pattern of division, ultrastructure of the mature spermatozoid, archegonial necks with six neck cell rows in transverse section, suppression of the function of the hypobasal cell in the embryo, placental organisation with coarse ingrowths in both generations, unistratose capsule walls, spherical spore mother cells, spore wall ontogeny with development of several to many multilamellar structures, and large, polar spores with both faces ornamented. Jungermanniidae normally have tetrahedral apical cells, several oil-bodies in each cell, antheridial ontogeny with a two-celled pattern of division, archegonial necks mostly with five cell rows in transverse section, the hypobasal cells differentiating into a haustorium in the embryo, placental organization with fine, dense labyrinths in the sporophyte placenta, capsule walls more than one cell-layer thick, quadrilobed spore mother cells, spore wall ontogeny with development of single tripartite structures, and small, inconspicuously polar spores with faint ornamentation (exception: Fossombronia). Additionally, the two subclasses differ in life history traits, dispersal mechanisms and habitat. Marchantiidae are characterized by good drought tolerance, rarity of asexual reproduction, high spore production, and restriction to terrestrial habitats. Jungermmaniidae are usually less drought tolerant than Marchantiidae, have frequent asexual reproduction and rather low spore production, and inhabit a wide diversity of habitats. The subclass Marchantiidae is subdivided into three orders, Marchantiales, Sphaerocarpales, and Monocleales; the majority of the species belong to the first group. Most genera of the Marchantiales are characterized by structurally complex thalli, with epidermal pores and assimilatory layer on the dorsal side and scales and pegged rhizoids on the ventral side. In many, the gametangia develop on stalked gametangiophores. The Sphaerocarpales and Monocleales lack epidermal pores, discrete

Bischler et al. 7 assimilatory layers, scales, pegged rhizoids, and gametangiophores. The calyptra becomes 2-4-layered only after fertilization in the Marchantiales and Sphaerocarpales, but differentiates before fertilization in the Monocleales. The seta is short, necrotic, or lacking in most Marchantiales and Sphaerocarpales, but conspicuously elongates at sporophyte maturity in the Monocleales. In recent classifications based on morphology, anatomy and ontogeny (Grolle, 1983; Schuster, 1984, 1992b; Bischler, 1998; Crandall-Stotler & Stotler, 2000), Sphaerocarpales and Monocleales are sister groups of the monophyletic Marchantiales and are believed to represent remnants of early diversification in liverworts. Instead, most molecular data indicate that the Sphaerocarpales and Monocleales are nested within the Marchantiales (rbcL data, Lewis et al., 1997; 26S rRNA data, Wheeler, 2000; Boisselier et al., 2002; plastid trnL data, Wheeler, 2000). The close relationship between Monocleales and Marchantiales is also supported by morphological characteristics (e.g., antheridia grouped in receptacles) as well as phytochemical data (Gradstein et al., 1992). The order Marchantiales is comprised of 14 families, 28 genera, and about 355 species (Bischler, 1998). Eleven families, 18 genera and 102 species occur in the Neotropics and one genus, Cronisia, is exclusive to this region. Most of the genera are represented by only few species (14 genera comprise 1-5 species, three genera 8-10 species) and more than half of the neotropical species of Marchantiales belong to the genus Riccia (53 spp.). The order Monocleales comprises a single genus with two species, one of them occurring in montane regions throughout the Neotropics. The Sphaerocarpales are comprised of 2 families, 3 genera and about 25 species worldwide. They occur mainly in subtropical and warm-temperate regions of southern South America and southern U. S. A., and are not yet known with certainty from the Neotropics. The systematic revision presented here is based based on morphology and the families are arranged according to the recent cladistic analysis, based on morphological characters, published by the first author (Bischler, 1998). Genera within families and species within genera or subgenera are arranged here alphabetically as usual in FLORA NEOTROPICA. Nomenclature of families and genera follows Grolle (1983) and terminology follows the Glossarium polyglottum bryologiae (Magill, 1990). The descriptions include the entire range of morphological variation as far as is known and are based on study of living material and herbarium specimens. Data on worldwide and neotropical distributions are synthesised and references are given for neotropical areas from which material has not been seen. The distribution maps are based on specimens examined and reliable bibliographic records. Figure captions include citation of herbaria of vouchers specimens only for specimens not cited in the text. The systematic treatment of the genus Riccia is based on the monographs by Jovet-Ast (1991, 1993) and was translated from the French and shortened by the first author; the final text was checked by S. Jovet-Ast. The treatments of the genera Plagiochasma and Marchantia are updated, shortened

Bischler et al. 8 versions of the revisions by Bischler (1979, 1984). Additional information, figures, and SEM pictures can be found in these monographs. The treatment of the genus Asterella in this volume is a preliminary one; the monograph of this large and difficult genus is in progress (Long, in prep.).

HISTORICAL SURVEY

The Marchantiidae as a subclass was first established by Schiffner (1893a: p. 1, 6), and was later on recognised by Müller (1940, as Marchantiales), Schuster (1980) and other authors. The subclass is sometimes referred to as “Marchantiidae Schust.” but this is erroneous, the group having already been recognised as a separate subclass in the late 19th century by Schiffner. Most of the genera of this subclass were described before 1880 and their taxonomic delimitation has not changed substantially since the fundamental morphological studies of Leitgeb (1879-1881). In contrast, half of the family names date only from the last 30 years, and subfamily subdivisions are even more recent. These recent developments in classification are not due to changes of taxonomic definitions but result from the application of more restrictive concepts in the delimitation of suprageneric taxa. Since Micheli (1729) many different classifications have been proposed for the genera of the Marchantiidae, the majority of them (including the current one) being based on characteristics associated with the sex organs. The main systematic treatments dealing with neotropical Marchantiidae are those of Bischler (1979, 1984), Evans (1915, 1917, 1919a, b), Evans, Howe & Haynes (1923), Gradstein et al. (1992, 2001), Hässel de Menéndez (1963), Jovet-Ast (1991, 1993), Spruce (1884-1885), Vianna (1985), and Yano (1995). Many species with wide geographical ranges are also treated in Schuster’s North American Flora (Schuster, 1992b). Many species of Marchantiidae were decribed from single or few specimens and from limited geographical areas, and have now been shown to be synonyms. Elimination of these “geographical” species started with the work of A. W. Evans, between 1914 and 1930, and is still in progress. Many species were shown to have wide distributions, disjunct between arid and semi-arid areas on several continents. Large numbers of collections made in the Neotropics in recent years have permitted us to gain a better understanding of the morphological variation at the species level. In spite of this, many neotropical species are still known from few specimens only, precluding critical evaluation of their taxonomic status.

Bischler et al. 9 MORPHOLOGY AND ANATOMY OF MARCHANTIALES AND MONOCLEALES

Since Sphaerocarpales are still unknown from the Neotropics, the present chapter deals with Marchantiales and Monocleales only. Thallus (Figs. 1: A-M, 2: A-C, E-F, 97: C, 158: C, 159: A). The thallus is usually structurally complex, differentiated into the epidermis, assimilatory layer and basal tissue. The epidermis is cut off from the distalmost epithelial cells and is usually unistratose, with few or no chloroplasts. The epidermal pores are 1) simple openings among epidermal cells, 2) simple openings bounded by one or several concentric rings of differentiated cells, or 3) compound openings with several concentric rings of cells, some above the epidermis, some bulging into the air chambers (e.g. Fig. 147: E). Often, a faint hyaline inner ring surrounds the opening. Epidermal pores are absent or vestigial in Dumortiera, and absent in Monoclea The assimilatory layer (s) develops by internal, schizogenous cell separation at thallus apex. The air chambers are in one layer, with or without chlorophyllose filaments, each chamber opening by a pore, or in several layers, with only some chambers opening by a pore. The assimilatory layer is compact in many Riccia species, without enlarged air chambers. In Dumortiera it is vestigial or missing and in Monoclea always absent. The ventral tissue is usually thick in the costal part of the thallus and thinner or absent in the thallus wings. In Ricciocarpos, Cyathodium and some species of Riccia, however, the ventral tissue may be thin (2-6 cell layers) in the costal part of the thallus as well. Oil-cells are usually numerous, except in Oxymitra and Riccia. In Monoclea, they may contain chloroplasts. Yellow- or red-walled sclerotic cells with strongly thickened walls are seen in several Marchantia species and mucilage cavities are observed in many unrelated taxa. Other idioblasts are known in Riccia, Cronisia, and Cyathodium. All taxa have unistratose scales on the ventral side of the thallus, except in a few species of Riccia. In Monoclea, they are replaced by one- to several-celled mucilage papillae. The scales are either in two rows, one on each side of the midrib, in several ill-defined rows, in 4-8 rows, each pair of rows of different size and shape, or in single, central row in a few Riccia. Often they have oil cells, one or several appendages, and marginal papillae. The unicellular rhizoids are partly smooth, partly pegged, i.e., with intracellular wall projections. In Monoclea, they are usually smooth and often branched. However, some isolated pegs have been observed in the latter genus.

Bischler et al. 10 Purplish or dark red pigmentation is frequent on thalli and/or scales; its chemical nature is unknown (Whittemore, 1991). The pigments are located in the cell walls, not in vacuoles like the anthocyanins (= oxidized flavonoids) of higher plants. Some species such as Monoclea gottschei, Riccia spp., Cyathodiumi and shade forms of other Marchantialean species lack purplish or red pigmentation. Dichotomous or pseudodichotomous branching exist in all taxa; dichotomies may be unequal, with one branch much larger than the other (i.e., in Lunularia, Marchantia). Additionally, ventral and apical adventitious branches produced after cessation of growth or initiation of terminal gametangiophores, are frequently observed. The development of several branching types presumably allows the plants to increase more rapidly in size under favourable conditions. In many Marchantiales the thalli roll up laterally when dry, only the blackish ventral side and the scales remaining visible. Some perennate during the dry seasons by tuberous, geotropical shoot apices or apical (rarely ventral) “ stalked ” tubers. Asexual Reproduction (Fig. 2: G-I). Asexual reproduction by specialized diaspores is much less frequent in the Marchantiidae than in the Jungermanniidae and is present in about 10 % of the neotropical species. Pluricellular gemmae develop either in thallus outgrowths or gemma-cups, which function as splash cups (in Lunularia, Marchantia), or as thallus outgrowth from thallus margins (in Cyathodium). Tubers, which are sometimes copiously produced, may also function as propagules. Sexual Reproduction (Figs. 2: D, K-M, 3: A, C-F, H, 97: B, F-G, 158: A-B, F-G, 159: B). Male and female gametangia develop either on the same thallus (monoecious), or on different thalli (dioecious). Monoecious taxa are more frequent in the Marchantiales (almost 70 % of neotropical taxa) than in other liverwort orders. Both monoecy and dioecy are found in all groups, and many genera include both monoecious and dioecious species. Dioecy may be linked to heterothallism, the male gametophyte being smaller than the female one (e.g., in some Riccia species). Among the species endemic to the Neotropics 77 % are monoecious, among those with wider geographical ranges 66 %. Apparently, monoecy versus dioecy does not play an important role in dispersal and migration in the Marchantiidae. Sex distribution is not always easy to assess because individuals may be protandrous or protogynous, male and female gametangia being induced by different environmental factors. Many species, stated in the past to be dioecious, might be monoecious, with delayed development of either male or female gametangia. Gametangia develop in acropetal sequence. A wide range of gametangial disposition is observed. A simple acropetal sequence of female gametangia, without aggregation into

Bischler et al. 11 receptacles or development of protective devices, allows rapid sequential sporophyte and spore production, and rapid completion of the life cycle. It is found mainly in taxa of transient or of permanent habitats with short rainy season (e.g., Riccia). Location of reproductive activity, with the tendency to group gametangia in receptacles (grooves, cushions, gametangiophores) is common in taxa with longer life cycles. Antheridia and archegonia may develop from superficial initials at the thallus apex or behind it, either dorsally or terminal (associated with loss of apical cell function) on the thallus or ventral branches. The antheridia may be embedded in thallus tissue, scattered in an open groove, irregularly and loosely aggregated in groups, in cushions, or, rarely, on stalked receptacles. The antheridial clusters can be naked or bounded by cilia, scales, or a membrane. The antheridia are cylindricalellipsoidal, with a unistratose jacket except distally where two layers may be present. The antheridial stalk is short, of 2-4 tiers of 4 cells (as far it is known). One- or two-celled paraphyses may be intermixed with the antheridia (Aytoniaceae). In Monoclea, mucilage hairs are present in the antheridial cavities, not only on the floor but also on the side walls of the cavities. The ostioles through which the spermatozoids are released are formed by protruding thallus tissue. The archegonia are embedded in the thallus, scattered or in 2-3 rows along thallus midline, loosely aggregated in groups, located in thallus depressions or cavities, in a single group at the thallus apex and subsequently shifted to the ventral side of thallus, in cushions, or initiated underneath the lobes of stalked receptacles. The dorsal position of archegonia, found in the Ricciaceae, Oxymitraceae, Corsiniaceae, Athalamia, and Plagiochasma, allows continuous initiation of female gametangia and sporophyte production. Unistratose scales bounding the gynoecia are present in the genera with stalked archegoniophores, and are found in the mature archegoniophores at the top and the bottom of the stalks, sometimes also along the stalks. The archegonia are bottle-shaped, with a neck of 6 cell rows in transverse section. Neck canal cells are 4-8 in the Marchantiales, 14-20 in Monoclea. Paraphyses (Corsiniaceae, Exormothecaceae) or mucilage hairs (Monoclea) can be intermixed with the archegonia in the archegonial clusters. Several envelopes of gametophytic origin develop around the fertilised archegonia. The archegonial wall (= calyptra) is initially one-layered and becomes 2-3-layered after fertilization, except in Monoclea where it divides prior to fertilization (2-3 layers), becoming subsequently 6-7-layered, or in Corsinia where it becomes thick and warty. It remains two-layered, with the inner layer collapsing before spore maturity, in the Ricciaceae (Ligrone et al., 1993) and probably in Oxymitra. In addition, pseudoperianths develop from the archegonial stalk and bound each sporophyte in some genera. They are campanulate in Marchantia and split at maturity into linear segments in Asterella. Groups of several archegonia and subsequently one or several sporophytes are bounded by involucres (of

Bischler et al. 12 thalline origin), except in the Ricciaceae. The involucre consists of a posterior, pluristratose scale in Corsiniaceae or a pyriform structure with epidermal pores and air chambers in Oxymitra. In the Marchantiales with stalked archegoniophores it is usually cup-shaped, bilabiate, bivalved or tubular, in Monoclea tubular and fused to thallus margins. Each archegonial cluster comprises several archegonia, except in the Ricciaceae (and Sphaerocarpales). Fertilization of more than one archegonium in a cluster or development of more than one sporophyte per cluster is rare and is observed in Corsiniaceae, Cyathodium, Lunularia, Marchantiaceae, and Monoclea. Gametangiophores (Figs. 3: B, G, I-K, 160: M). Gametangiophores are not found in all families. Gametangiophores are branch systems with thallus dichotomies separeated by very short internodes resulting in receptacles which become elevated on constricted thallus segments (= stalks). The gametangiophore stalks may have assimilatory strips on the anterior side and rhizoids on the posterior side, resembling closely contracted thallus segments. Male gametangiophores or antheridiophores are present only in the Marchantiaceae. Disk-shaped receptacles characterize all genera except Marchantia in which the receptacles are lobed. In the female gametangiophores or archegoniophores, secondary cell proliferation of the dorsal receptacular tissue displaces the archegonia onto the ventral side of the receptacles. The dorsal side of the receptacles frequently has epidermal pores and an assimilatory layer, like the thalli. The structure of the epidermal pores is not necessarily the same in thallus and receptacles; for instance, pores are simple in the thallus and compound in the receptacles of the Aytoniaceae. In Lunularia and some species of Athalamia and Sauteria the receptacular tissue is so restricted that epidermal pores and assimilatory layer fail to develop or are very reduced. The stalks of the gametangiophores often show channel-like rhizoid furrows. Early involvement of the apical cell, dividing before development of the receptacle, results in stalks without rhizoid furrows and the persistence of apical thallus growth, e.g., in Plagiochasma and Athalamia. Fertilization takes place before stalk elongation, except in the family Marchantiaceae where the apical cells of the receptacular branches remain active and produce archegonia after stalk elongation. Stalk elongation is delayed until spore maturity in Lunularia. Sporophytes (Fig. 159: C-D; 160: L). Two patterns of embryonic development are known in the Marchantiales: 1) development of a four-

Bischler et al. 13 celled filamentous embryo (filamentous type), or 2) formation of an “ octant ” embryo (octant type), by subdivision of the daughter cells by means of successive vertical walls at right angles. Both types have been observed in closely related taxa within a single genus, or even within the same species. A very different early ontogeny, including development of several free nuclei not separated by cell walls, has been observed in Monoclea (Ligrone et al., 1993). Placental organization is characterized by one layer of well differentiated transfer cells, with highly branched and anastomosing ingrowths in the sporophyte, and by several layers of transfer cells in the gametophyte, with extensive wall labyrinths or coarse wall ingrowths (Ligrone et al., 1993). However, no transfer cells have been found at the sporophyte-gametophyte junction in Riccia. The mature sporophyte is radially symmetric and comprised of foot, seta, and capsule. It is of small size, chlorophyllose when immature but usually non-chlorophyllose at maturity. The foot is spherical, conical, or cup-shaped; it is missing in the Ricciaceae and Oxymitraceae. The seta usually remains very short except in Lunularia, where it is massive and shortly elongating, and Monoclea, in which a very long and thick seta (20-40 cells across diameter) is produced. In taxa with cleistocarpous capsules the seta is particularly short and thin, being necrotic at the time of spore maturity (Corsiniacae) or missing (Ricciaceae, Oxymitraceae). The capsule is usually globose, with a unistratose wall and a pluristratose apical cap or lid. Annular or semi-annular thickenings are often present in the capsule wall. Absence of thickenings is observed in unrelated taxa (Corsinia, Lunularia, Aytoniaceae). Upon maturity of the spores, the capsule wall splits into irregular valves, into four valves after having shed a small apical cap, or opens by a lid. Cleistocarpous capsules are known in the Corsiniacae, Ricciaceae and Oxymitraceae. The spores are released after decay of thallus tissue, on the dorsal or the ventral side of the thallus. In the Ricciacae and Oxymitraceae, the wall disintegrates before spore maturity. Capsule shape, structure, and dehiscence are very different in Monoclea: the capsule is ellipsoidal, opens by a single, longitudinal slit, and the wall is unistratose and provided with internal and external spiral thickenings. Meiosis is usually preceded by individualization of spore mother cells and elaterocytes in the capsules. Insertion of mitotic divisions into either cell type (spore mother cells or elaterocytes) results in three types of spore/elater ratios. The sporocyte-elaterocyte separation can be followed directly by meiosis of the spore mother cells, resulting in a spore/elater ratio of 4 : 1. The sporogonous cells can also undergo several mitotic divisions before meiosis, the elaterocytes remaining undivided. The spore/elater ratio will then be more than 4 : 1 (as e.g., in Lunularia, Marchantia, Monoclea). Finally, the elaterocytes may undergo mitotic divisions before meiosis, and the spore/elater ratio will be nearly 1 : 1 (as in Exormotheca). In the Corsiniaceae, the elaterocytes do not or imperfectly differentiate into elaters and persist as sterile cells or reduced elaters intermixed with the spores. In the Ricciaceae and Oxymitraceae, no sporocyte-elaterocyte individualization occurs; all cells develop into spore mother

Bischler et al. 14 cells. Elaters and Spores (Elaters, Figs. 136: L-M, 158: D-E, 160: K. Spores, Figs. 78-81, 97: A, D-E, H, 104, 116-118, 136: A-K, 160: A-I). The elaters are unicellular, usually long and hyaline cells with 1-4 pigmented helical thickenings. The spores are polar through compression, the proximal face has a triradiate scar, and both faces ornamented. Size of mature spores varies from 8 µm to 200 µm diameter and is largest in genera with short-lived gametophytes but smaller in long-lived taxa. Mature spores are smallest in Lunularia, most of the species of the Marchantiaceae, and Monoclea. Spore number varies from 25 to over 10, 000 per capsule and is fairly constant within each genus. Spore ornamentation patterns often vary within the same genus and can be used for species distinction (i.e., in Riccia, Asterella, Marchantia), whereas in other genera they are quite uniform (i.e., Plagiochasma). Some have strikingly different ornamentation patterns on distal and proximal faces (i.e., Oxymitra, Corsiniaceae, some Exormothecaceae, some Asterella and Marchantia species). Spore viability is usually long and seems uncorrelated to spore size. Spore germination is exosporic, with the development of a germ tube and a germ rhizoid. The spore wall ruptures irregularly, tangentially, or on the proximal or distal face. The germ rhizoid originates in front, laterally, or obliquely to the germ tube, separated or not by a septum. As far as is known germination patterns are constant within each genus (Inoue, 1961). In Monoclea, spores swell at the onset of germination (exosporic), but a germ tube and germ rhizoid fail to develop.

PLOIDY LEVEL, POPULATION STRUCTURE, AND SPECIATION

The basic haploid chromosome numbers in Marchantiales and Monocleales are x = 8 or 9. Unrelated genera may have 8 chromosomes (Riccia, Corsinia, Exormotheca) and 8 or 9 chromosomes occur within the Corsiniaceae and Ricciaceae. Meiotic failures can result in unreduced spores that germinate into functionally haploid gametophytes with a duplicated chromosome set. Polyploids are frequent in the Marchantiales; commonly, the haploid and polyploid populations are morphologically identical. Among species occurring in the Neotropics haploid and polyploid populations are known from Reboulia hemisphaerica, Plagiochasma rupestre, Corsinia coriandrina, Targionia hypophylla, Dumortiera

Bischler et al. 15 hirsuta, Riccia atromarginata, R. lamellosa, and R. trichocarpa; only polyploid ones in Plagiochasma crenulatum, Exormotheca pustulosa, and Marchantia breviloba. Polyploids of Reboulia hemisphaerica, Corsinia coriandrina, and Targionia hypophylla have been shown to be of allopolyploid origin (Boisselier-Dubayle et al., 1998; Boisselier-Dubayle & Bischler, 1998, 1999). A sharp genetic difference between European and Texan populations with the same ploidy level was found in Corsinia coriandrina and Targionia hypophylla, whereas European and North American populations of Reboulia hemisphaerica were genetically similar. It should be noted, however, that almost all genetic data on Marchantiidae are from populations from temperate regions. Very few chromosome counts and no genetic analyses at all are available for neotropical populations. In species with dioecious haploids, the polyploids are usually monoecious, though dioecious polyploids exist also (Bischler & Boisselier-Dubayle, 1993). The monoecious polyploids of the Marchantiaceae often have archegoniophores with male branchlets sprouting from the female receptacle, e.g., in Marchantia breviloba. Population studies in European and Mediterranean Marchantialean species usually recorded low levels of inter-colony genetic variability. Sexual reproduction occurred frequently but gene flow seemed to be restricted. Intra-colony fertilisations seemed to prevail, inter-colony exchanges being severely restricted by habitat specificity and the constraints of aquatic fertilization. Most colonies examined consisted of a single genotype. Despite reproductive isolation, no morphological divergence was observed between widely separated, conspecific populations. Self-fertilization was shown to result in viable and healthy offspring in Plagiochasma rupestre (Boisselier-Dubayle et al., 1996). It thus appears that active speciation in Marchantiidae is lacking in the present day. Although data are lacking, we expect that population structure and genetic behaviour of the neotropical species of Marchantiidae are similar to those observed in Europe and elsewhere.

ECOLOGY, LIFE HISTORY TRAITS, AND DISPERSAL

The species of Marchantiales and Monocleales are terrestrial or saxicolous and often grow in harsh, disturbed habitats. Drought- and cold-tolerance are common. Many are able to withstand strong illumination and drastic diurnal temperature changes. Microtopography, microclimate and edaphic conditions are of primary importance for their establishment and geographical distribution. Drought-resistance in the Marchantiales involves gametophytes having a metabolism that permits the resumption of growth after months or even years of drought and desiccation. The capacity of cells

Bischler et al. 16 to resume growth after prolonged water stress is seen in several lineages, i.e., in Ricciaceae, Corsiniaceae, Targioniaceae, Exormothecaceae, most Aytoniaceae and some taxa of the Cleveaceae. Others, i.e., many Riccia species, escape dry periods by the rapid development of gametophytes and the continuous initiation of fast-maturing sporophytes whilst water is available. These species persist as spores left behind once the favourable growing season is over, which may be as short as 2-3 weeks. However, fertilization requires water; all taxa need water at some time in their lives. Four groups of species with similar life history patterns have been proposed (Bischler, 1998). Each group shows specific life history traits, associated not only with ecological factors corresponding to specific habitats, but also with morphological characteristics and, as a consequence, with generic and familial membership. The first group is characterized by short life expectancy, high reproductive effort, dorsal gametangia, the archegonial clusters not developing into archegoniophores, and cleistocarpous capsules, containing few, large spores. The gametophytes are usually small, drought-tolerant, and asexual propagules are missing. The representatives of this group colonize shortly available, open habitats with intermittent moist conditions, as seepage areas, intermittent rivers, or temporary ponds. In undisturbed habitats, some may have higher life expectancies and perennate in dry condition. The group includes Oxymitraceae, Ricciaceae, Corsiniaceae, Riella and Sphaerocarpos. It is represented in the Neotropics by five genera and 57 taxa. The second group comprises species with large, drought-intolerant gametophytes of long life expectancy, with male and female gametes produced in terminal receptacles, the archegonial clusters developing into archegoniophores with long stalks. The spores are small and numerous. Sexual reproductive effort is moderate, asexual propagules frequent. This group colonizes mesic to wet, sometimes open and disturbed habitats, available for more or less long periods in pioneer situations, without marked water stress. In the Neotropics it is comprised of four genera (Lunularia, Dumortiera, Marchantia, Monoclea) and 11 taxa. Even though Monoclea does not develop archegoniophores, the genus fits in this group by its life history traits. The third group is small and closely related to the following group. It includes species with perennating tubers or tuberous thallus apices. The gametophytes are small, with dorsal androecia and terminal gynoecia. Spores are few, medium-sized or large. Reproductive effort is moderate. The group is represented in the Neotropics by two genera (Exormotheca, Cyathodium) and 6 species. Cyathodium is the sole genus of the Marchantiales able to survive under low light intensities, e.g., on lowland rainforest floors. The fourth group is characterized by small to large gametophytes, with dorsal androecia and dorsal

Bischler et al. 17 or terminal gynoecia, short archegoniophore stalks, and medium sized, fairly numerous spores. Life expectancy is moderate, as is reproductive effort. No asexual propagules are produced. Habitats are usually somewhat exposed and characterized by more or less long periods of water stress, as are rock crevices, or rock outcrops. Some species are drought tolerant, others hardly stand prolonged water stress, especially some Asterella species and Athalamia andina. The group includes the Targioniaceae, Cleveaceae, and Aytoniaceae. It is represented in the Neotropics by eight genera and 28 taxa. A striking feature of life history patterns is the differential investment in spores as a function of life expectancy. In habitats which undergo long periods of stress, life span is reduced. High growth rates are required, with early production of spores with sufficient reserves to permit rapid development of new gametophytes. Increased investment in spores, few spores of large size with prolonged germination capacity might be more efficient for reproduction in habitats of short availability but of periodical recurrence. Spore inefficiency, from failure to find suitable habitats, seems to be low. Riccia is the best example of this pattern, and represents also the genus that has known the highest diversification in the area. In wet to mesic or disturbed habitats which last longer, opportunities for spore establishment are scantier. Individuals with a longer life span, producing numerous, small spores might be favoured in these conditions, for instance Marchantia, Dumortiera, and Monoclea. In longer-lasting habitats subject to prolonged periods of water stress such as rock crevices and rock outcrops, intermediate investment in spore production is found. The species are stress-tolerant and produce fairly numerous spores of moderate size, which might find suitable conditions for germination in the vicinity of the mother colony. Plagiochasma and Asterella are representatives of this pattern. Familial and generic membership have previously been shown to be linked to life history patterns (Bischler, 1998). Major relationships within groups sharing similar life history traits are most likely due to common ancestry and not to convergence resulting from particular habitat conditions. Diversification in life history traits seems to have occurred at either the generic or the familial level, and to have been conserved in the course of evolution.

DISTRIBUTION AND ALTITUDINAL ZONATION

Marchantialean species are widespread, from the Arctic (Svalbard) to the Antarctic (Antarctic peninsula and Antarctic islands). Many species show outstanding adaptive potentialities and ranges including several continents, without showing morphological differentiation.

Bischler et al. 18 Wide geographical ranges, including more than one continent, are found in 26 neotropical taxa. Fifteen of these are widespread in tropical America, the 11 others are Mediterranean species occurring in the Neotropics only at the northern (4 species), or northern and southern limits of the region (7 species). All these widespread species occur also in the southern U. S. A. except Cyathodium cavernarum and C. foetidissimum (tropical ranges), Marchantia berteroana (southern hemisphere range), and Exormotheca pustulosa, recorded in the New World solely from Mexico where it might have been introduced. The southern U. S. A. is also the northernmost occurrence of 13 droughttolerant neotropical species, six of which occur only in Mexico whereas seven are widespread in the Neotropics. One genus (Cronisia) and 76 species of Marchantiidae are only known from the Neotropics. Among the strictly neotropical species, 31 have wide distributions within the region. The distribution of the majority of these neotropical species, however, is rather narrow and 26 may be considered endemics, being restricted to a very limited area. The endemics are found in all regions except on the Guyana shield and on the small West Indian islands. Four are known from Central America, two from Greater Antilles, two from the Galapagos islands, seven from the Andes, and 11 from eastern Brazil. Eighteen belong to the genus Riccia, including all endemic species from the Antilles, Galapagos islands, and Brazil. Analysis of the elevational distribution of the species, using the altitudinal belts defined by Gradstein (1995) (0-1200 m = lowland, 1200-2200 m = lower montane zone, 2200-3500 m = upper montane zone, 3500 m to more than 4000 m = subalpine), shows that 51 species occur in a single zone, 26 in two zones, 23 in three zones, and six in the four zones. In contrast to Jungermanniidae (Gradstein, 1995) more than half of the taxa are found in more than one altitudinal belt and 18 out of the 26 endemics are lowland species. Species diversity is highest in the lowlands (83 species), somewhat lower in the lower montane (56 ) and the upper montane (42) zones, and lowest in the subalpine belt (15). The elevational distribution of the Marchantiidae thus clearly differs from that of the Jungermanniidae, which are most diverse in the upper montane forests, between 2500 and 3000 m. These differences are presumably due to the very different ecological requirements of the two groups. Marchantiidae are drought-resistant plants that do not need permanent humidity, and they require high light intensities so that they are absent from the dense forest. Jungermanniidae, however, are much less drought-resistant, need more constantly high humidity levels, favour shaded habitats and abound in the dense forest. A clear preference for higher, montane elevations is also found in the mosses (Churchill et al., 1995).

Bischler et al. 19

PHYLOGENY

The classifications of the Marchantiidae, including the one in current use, are usually based primarily on gametangial characteristics. Two main evolutionary hypotheses were postulated in the past, 1) the progressive specialisation hypothesis (Leitgeb, 1879-1881; Mishler & Churchill, 1985), postulating the evolution of plants with archegoniophores from plants without them, and 2) the progressive reduction hypothesis (Wettstein, 1903-1908; Schuster, 1984, 1992b), postulating evolution in the opposite direction (development of plants without archegoniophores from those having them). Phylogenetic analysis using structural data (morphology, anatomy, development) clearly fails to support either hypothesis, with bootstrap values below 50 % for all deep branches (BoisselierDubayle et al., 2002). Interestingly, all molecular trees published thus far disclose different relationships and are in clear support of the progressive reduction theory (18S rRNA data, Capesius & Bopp, 1997; rbcL data, Lewis et al., 1997; 26S rDNA data, Wheeler, 2000, Boisselier-Dubayle et al., 2002; plastid trnL-F genes, Wheeler, 2000; plastid cpDNA matK and trnL intron, Long et al., 2001). The data show that extant genera without archegoniophores are derived from archegoniophore-bearing forms and that Monocleales and Sphaerocarpales are nested within the Marchantiales (Fig. 5). The position of Monoclea within the Marchantiales comes as a surprise in view of the numerous morphological autapomorphies of the genus and its presumed isolated systematic position. However, all studies including Monoclea (LSU rDNA data (Wheeler, 2000; Boisselier-Dubayle et al., 2002), plastid trnL-F data (Wheeler, 2000), rbcL data (Lewis et al., 1997) show this genus to be firmly nested within the Marchantiales. At the generic level, the single large genus that has been analysed, Asterella, also proved to be paraphyletic (plastid trnL intron and matK data, Long et al., 2001) and the delimitation of other genera was cast into doubt. It thus appears that the traditionally recognised orders and families of the Marchantiales as well as the delimitation of some of the genera are not supported by the molecular data and that the classification of the Marchantiales as adopted here might have to be revised in the future. It should be kept in mind, however, that only a rather limited number of genera and species was included in the molecular trees published thus far and that many taxa have not yet been analysed genetically. In contrast, the recent morphological trees were based on a very complete analysis of the group. For this reason, the recently published morphological trees (Bischler, 1998; Crandall-Stotler & Stotler, 2000) are chosen as the basis for the classification of the Marchantiidae in this treatment.

Bischler et al. 20

SYSTEMATIC TREATMENT

CONSPECTUS

1. Marchantiales 1. Oxymitraceae 2. Ricciaceae

1. Oxymitra (1 species) 2. Riccia (53) 3. Ricciocarpos (1)

3. Corsiniaceae

4. Corsinia (1) 5. Cronisia (2)

4. Cyathodiaceae 5. Targioniaceae

6. Cyathodium (5) 7. Targionia (2)

6. Exormothecaceae

8. Exormotheca (1)

7. Cleveaceae

9. Athalamia (2)

`

10. Sauteria (1) 8. Aytoniaceae

11. Asterella (10) 12. Cryptomitrium (1) 13. Mannia (2) 14. Plagiochasma (9) 15. Reboulia (1)

9. Lunulariaceae

16. Lunularia (1)

10. Marchantiaceae

17. Dumortiera (1) 18. Marchantia (8)

2. Monocleales 11. Monocleaceae

19. Monoclea (1)

12. Riellaceae

20. Riella (0)

13. Sphaerocarpaceae

21. Sphaerocarpos (0?)

3. Sphaerocarpales

INTRODUCTORY KEY TO THALLOID LIVERWORTS OF TROPICAL AMERICA

Bischler et al. 21 Modified after Gradstein et al. (2001). Bold face: this treatment.

1. Plants consisting of a filamentous protonema or a tiny stem without leaves. Gametoecia on leafy branches ..................................................................................... (Jungermanniales: Lepidoziaceae) 1. Plants thalloid, not consisting of a filamentous protonema or a tiny stem. Leafy branches absent ........................................................................................................................................................ 2

2. Plants aquatic, erect, consisting of a narrow axis and a broad, undulate or spirally coiled wing, growing in alkaline ponds or lakes, not yet known from the Neotropics ....................................... ...................................................................................................... 3. Sphaerocarpales: 20. Riella 2. Plants terrestrial or epiphytic, not consisting of a narrow axis and a broad, undulate or spirally coiled wing .................................................................................................................................. 3

3. Dorsal thallus surface of fertile plants covered by numerous obovate to cylindrical involucres, the involucres to 3 mm long. Thallus delicate, very thin, growing on naked earth in subtropical regions at low elevations...............................................3. Sphaerocarpales: 21. Sphaerocarpos 3. Dorsal thallus surface of fertile plants not covered by numerous involucres ....................... 4

4. Thallus unistratose throughout, without midrib .......................................... fern prothallium 4. Thallus more than one cell thick, or unistratose and with a midrib (midrib simple, not branched; when branched: Hymenophyllaceae!) ................................................................... 5

5. Epidermal cells with only 1 large chloroplast (2-4 in Megaceros). Capsules long-cylindrical to linear, green (turning ± black after dehiscence), without seta; usually numerous capsules projecting from the thallus surface. Thallus growing in rosettes............................................. ..................................................................................................... Anthocerotae or hornworts 5. Epidermal cells with numerous small chloroplasts. Capsules globose to ellipsoidal, black, immersed in the thallus or hanging down from stalked receptacles (then without or with reduced seta), or projecting upwards from the thallus on a long, colourless seta ................ 6

6. Thallus with air chambers (cross section) and numerous small pores on the dorsal surface. Thallus underside with scales, at least near the tip. Antheridia and archegonia usually born in receptacles............................................................................. 1. Marchantiales (Keys 2, 3) 6. Thallus lacking air chambers and pores. Thallus underside with or without scales. Receptacles absent (except Dumortiera, couplet 13, which has stalked receptacles, and Monocleales, couplet 12, which have sessile male receptacles) ....................................... 7

Bischler et al. 22

7. Plants growing in complete or incomplete rosettes (loosely forked in aquatic plants). Upper surface of the thallus with a median groove. Sporophytes embedded in the thallus ....................................... 1. Marchantiales: 1. Oxymitraceae, 2. Ricciaceae (Keys 2, 3) 7. Plants not growing in rosettes. Upper surface of the thallus without median groove. Sporophytes not embedded in the thallus ....................................................................... 8

8. Thallus surface in fresh material densely spotted with numerous small, whitish or darkish dots. Thallus usually without midrib and large (3-20 cm long), forked, with crispate-undulate margins. Rhizoids smooth. Thallus with mucilage papillae at apex. Capsules opening along a single, longitudinal slit. Seta long .......................................... ....................................................................................... 2. Monocleales: 19. Monoclea 8. Thallus surface not densely spotted in fresh material. Thallus with or without midrib. Rhizoids smooth or (Dumortiera) pegged. Thallus without mucilage papillae at apex (along margins in Pteropsiella). Capsules opening along 2-4 slits or irregularly. Seta short or long .......................................................................................................................... 9

9. Thallus large, 1-3 cm wide, forked, with a ventral midrib made of rhizoids. Rhizoids pegged (papillose). Archegonia and antheridia on stalked receptacles ...................... .............................................................................1. Marchantiales: 17. Dumortiera 9. Thallus smaller, less than 1 cm wide, simple, forked or pinnate, without ventral midrib made of rhizoids. Rhizoids smooth. Stalked receptacles lacking .............. 10

10. Thallus unbranched or with a few short, ventral-intercalary branches. Thallus margins, at least near the apex, with long, sausage-shaped slime papillae. Midrib present, narrow, on the ventral side with rhizoids in interrupted tufts. Sexual organs on leafy branches.......................... (Jungermanniales, Lepidoziaceae: Pteropsiella) 10. Thallus variously branched. Thallus margins without sausage-shaped slime papillae. Midrib present or absent, rhizoids when present not in interrupted tufts. Sexual organs on the thallus surface or under the surface in cavities, not on leafy branches ........................................................................................... (Metzgeriales)

KEY 2. KEY TO THE NEOTROPICAL FAMILIES OF MARCHANTIALES

Bischler et al. 23 1. Archegonia scattered or loosely aggregated in a dorsal groove, a cavity or a thallus depression, or in an involucre below and beyond the thallus apex........................................................................ 2 2. Archegonia and sporophytes in a dorsal groove, a cavity or in a thallus depression. Sporophyte foot small or absent; seta short and thin (4-6 cells across diam.) or absent. Capsule cleistocarpous. Elaters no more than twice as long as wide, with single helical band, or absent.. 3 3. Thalli often rosette-forming. Air chambers in 1-3 layers, without chlorophyllose filaments, sometimes narrower than the cells and inconspicuous. Calyptra thin. Sporophyte lacking foot and seta. Capsule wall disintegrating prior to spore maturity. Elaters lacking. Spores on distal face with ornamented or pitted areoles, with spines or nearly smooth........ ...4 4. Radial walls of epidermal pores often thickened. Ventral scales in 2 rows. Antheridia in a dorsal groove bordered by cilia. Archegonia and sporophytes in dorsal thallus depression, in pyriform involucres with air chambers and pores, bounded by scales. Proximal and distal faces of spores with strikingly different ornamentation ............................................................................................................... 1. Oxymitraceae 4. Epidermal pores, if differentiated, with thin radial walls. Ventral scales lacking, in 1-2 rows or in several ill-defined rows. Antheridia scattered or in a dorsal groove, without scales or cilia. Archegonia and sporophytes scattered or in 2-3 rows in dorsal groove, involucres and scales lacking. Proximal and distal faces of spores with similar or nearly similar ornamentation, or spores remaining in tetrads .................................2. Ricciaceae 3. Thalli not rosette-forming. Air chambers in 1(-2) layers, with chlorophyllose filaments. Calyptra thin, or thick and warty. Sporophyte with a small foot and a short seta. Capsule wall persistent, with or without annular thickenings. Elaters present, with or without helical band. Spores on distal face with rounded protuberances, or with plates.. ............................ ................................................................................................................... ...3. Corsiniaceae 2. Archegonia and sporophytes in an involucre below and beyond thallus apex. Sporophyte with a well developed foot and seta, 6-20 cells in diam. Capsule not cleistocarpous, opening by a lid, then splitting into irregular valves to 1/3-1/2 of capsule length. Elaters long, at least ten times as long as wide, with 1-4 helical bands...........................................................................5 5. Thallus without purplish pigmentation, with tubers. Epidermal pores a simple opening among epidermal cells, without hyaline inner ring. Air chambers empty, in one layer. Basal tissue restricted to ventral epidermis or to 1-2 cell layers. Ventral scales small, without or with filiform appendage. Asexual reproduction by fragmenting thallus segments, or ventral

Bischler et al. 24 tubers. Capsule wall with annular thickenings in upper part only. Elaters and spores few.......................... ............................................................................. ......4. Cyathodiaceae 5. Thallus with purplish pigmentation, without tubers. Epidermal pores bounded by 1-3 concentric rings of cells, with hyaline inner ring. Air chambers in one layer, with chlorophyllose filaments. Basal thallus tissue thick. Ventral scales large, with conspicuous, lanceolate-triangular appendage. No asexual reproduction. Capsule wall with annular thickenings throughout. Elaters and spores numerous......... ...................... 5. Targioniaceae 1. Archegonia in well delimited cushions with several archegonial cavities, the cushions developing after fertilization into stalked archegoniophores, or archegonia initiated on ventral side of stalked archegoniophores ........……………………..................................................................6 6. Epidermal pores compound (absent or vestigial in Dumortiera). Both archegoniophores and antheridiophores produced . .........................................................................10. Marchantiaceae 6. Epidermal pores simple, with one or several concentric rings of cells. Archegonia in well delimited cushions with several archegonial cavities, the cushions developing after fertilization into archegoniophores. Antheridiophores never produced . .....................................................7 7. Epidermal pores with one ring of differentiated cells. Epidermal pores of female receptacle simple or absent. Capsule wall with annular thickenings. Spores with rounded protuberances on distal face .................................................................................................8 8. Epidermal pores strongly elevated above epidermis. Air chambers in a single layer, with chlorophyllose filaments. Ventral scales in 2 rows. Involucres tubular ................... ..........................................................................................................6. Exormothecaceae 8. Epidermal pores not strongly elevated above epidermis. Air chambers in 1-4 layers, without chlorophyllose filaments. Ventral scales in several ill-defined rows. Involucres bilabiate ......................................................................................................7. Cleveaceae 7. Epidermal pores with several concentric rings of cells (with a single ring of hardly differentiated cells in Plagiochasma rupestre). Epidermal pores of female receptacle compound (absent in Lunularia). Capsule wall without annular thickenings. Spores without rounded protuberances on distal face (except in Mannia californica).................................9 9. Air chambers in several layers, without chlorophyllose filaments. Ventral scales with 1-4 lanceolate or filiform appendages, usually not constricted basally, acute apically, or single, rounded to ovate. Gemma cups never produced. Antheridia in irregular groups or

Bischler et al. 25 in cushions, not bounded by a membrane. Archegoniophore stalk elongating after fertilization; receptacle with compound epidermal pores. Involucres cup-shaped or bilabiate. Capsule urn-shaped, opening by a lid. Spores 35-120 µm in diameter ............ ..................................................................................................................8. Aytoniaceae 9. Air chambers in a single layer, with chlorophyllose filaments. Ventral scales with single, large, reniform appendage, constricted basally, rounded apically. Gemmae in crescent-shaped gemma-cups. Antheridia in cushions bounded by a membrane. Archegoniophore stalk elongating at time of spore maturity; receptacle without epidermal pores. Involucres tubular. Capsule splitting to the base into 4 valves. Spores 14-22 µm in diameter ....... ......................................................................................... 9. Lunulariaceae

KEY 3. KEY TO THE NEOTROPICAL GENERA OF MARCHANTIALES

1. Archegonia and sporophytes scattered on the thallus, in a dorsal groove, depression or cavity, or in an involucre below and beyond the thallus apex, never on stalked receptacles .......................2 2. Thallus branching dichotomous. Archegonia and sporophytes scattered or in a dorsal cavity, groove, or depression. Capsule cleistocarpous. Elaters absent or short, at the most twice as long as wide, without or with a single helical band ..........................................................................3 3. Thalli often rosette-forming. Air chambers in 1-3 layers, without chlorophyllose filaments, sometimes narrower than the cells and inconspicuous. Archegonia scattered or in dorsal groove or thallus depression. Capsule wall disintegrating before spore maturity. Elaters lacking. Spores on distal face with areoles, spines, or nearly smooth ....................4 4. Radial walls of epidermal pores thickened. Ventral scales in 2 rows. Antheridia in dorsal groove bordered by cilia. Archegonia and sporophytes in dorsal thallus depression, in pyriform involucres, with air chambers and pores, bounded by scales. Proximal and distal faces of spores with strikingly different ornamentation.......................1. Oxymitra 4. Epidermal pores, if present, with thin radial walls. Ventral scales, if present, in 1-2 or in several ill-defined rows. Antheridia scattered or in dorsal groove, without scales or cilia. Archegonia and sporophytes scattered, or in 2-3 rows in dorsal groove, involucres and scales lacking. Proximal and distal faces of spores with similar or nearly similar ornamentation, or spores remaining in tetrads................................................................5

Bischler et al. 26 5. Epidermal pores absent, or a simple opening among epidermal cells, rarely bounded by a ring of differentiated cells. Thallus and scales without oil-cells. Ventral scales in 2 rows, short, margins not serrulate, rarely in single row or absent. Antheridia and archegonia scattered ......................................................................................2. Riccia 5. Epidermal pores bounded by a ring of differentiated cells. Thallus and scales with oil-cells. Ventral scales in several ill-defined rows, very long in aquatic forms, with serrulate margins. Antheridia and archegonia in dorsal groove ..............3. Ricciocarpos 3. Thalli not rosette-forming. Air chambers in 1 (2) layers, with chlorophyllose filaments. Archegonia in thallus cavities. Involucre a posterior, pluristratose scale, incompletely enclosing or involute around archegonia. Capsule wall persistent, with or without annular thickenings. Elaters short, with single helical band, or sterile green cells mixed with spores. Spores on distal face with plates or rounded protuberances ...............................................6 6. Thallus without purplish pigmentation. Ventral scales in several ill-defined rows. Calyptra thick, warty. Involucre a thick, posterior, pluristratose scale. Capsule wall without annular thickenings............................................................................4. Corsinia 6. Thallus often with purplish pigmentation. Ventral scales in two rows. Calyptra thin, not warty. Involucre a thin, unistratose scale, involute around archegonia. Capsule wall with or without annular thickenings ...............................................................5. Cronisia 2. Thallus branching dichotomous and ventral. Archegonia and sporophyte in an involucre below and beyond the thallus apex. Capsule opening at spore maturity by a lid, then splitting into irregular valves. Elaters long, at least ten times longer than wide, with 1-4 helical bands7 7. Thallus with purplish pigmentation. Epidermal pores bounded by 1-3 concentric rings of cells, with hyaline inner ring. Air chambers with chlorophyllose filaments. Ventral tissue more than 2 cell layers thick. Ventral scales large, with conspicuous, lanceolate-triangular appendage. Asexual reproduction lacking. Elaters numerous (spore/elater ratio 4/1). Spores numerous (approx. 3000 per capsule) ...............................................................7. Targionia 7. Thallus without purplish pigmentation. Epidermal pores simple openings among epidermal cells, not bounded by concentric rings of cells and without hyaline inner ring. Air chambers without chlorophyllose filaments. Ventral tissue thin, of 0-2 cell layers. Ventral scales small, with filiform appendage or appendage lacking. Asexual reproduction by fragmenting thallus segments or ventral tubers. Elaters few (spore/elater ratio higher than 4/1). Spores few (20-100 per capsule) ........................................................... 6. Cyathodium

Bischler et al. 27 1. Archegonia in cushions with several archegonial cavities, the cushions developing after fertilization into stalked archegoniophores, or archegonia initiated on the ventral side of stalked archegoniophores.. .........................................................................................................................8 8. Epidermal pores with one ring of differentiated cells, with or without a hyaline inner ring, never compound. Epidermal pores on female receptacle simple or absent. Capsule wall with annular thickenings. Spores with rounded protuberances on distal face ..................................9 9. Epidermal pores strongly elevated above epidermis. Air chambers in one layer, with chlorophyllose filaments. Ventral scales in two rows. Involucres tubular... 8. Exormotheca 9. Epidermal pores not strongly elevated above epidermis. Air chambers in 1-4 layers, without chlorophyllose filaments. Ventral scales in several ill-defined rows. Involucres bilabiate .............................................................................................................................10 10. Ventral scales without oil-cells. Archegoniophores not terminal, stalk without rhizoid furrow. ......................................................................................................... 9. Athalamia 10. Ventral scales with oil-cells. Archegoniophores terminal, stalk with a single rhizoid furrow. ......................................................................................................... 10. Sauteria 8. Epidermal pores with several concentric rings of cells and a hyaline inner ring, or compound, rarely vestigial or absent (with a single ring of hardly differentiated cells and without hyaline inner ring in Plagiochasma rupestre). Epidermal pores on female receptacle compound (absent in Lunularia). Capsule wall with or without annular thickenings. Spores on distal face variously ornamented but without rounded protuberances (except in Mannia californica) ..................11 11. Epidermal pores with several concentric rings of cells (except in Plagiochasma rupestre). Antheridiophores not produced. Pseudoperianth lacking or split into linear segments. Capsule wall without annular thickenings ........................................................12 12. Air chambers in one layer, with chlorophyllose filaments. Ventral scales with a single, reniform appendage. Gemmae in crescent-shaped gemma cups. Capsule splitting to the base into 4 regular valves. Spores 14-22 µm diameter ....................16. Lunularia 12. Air chambers in several layers, without chlorophyllose filaments. Ventral scales with 1-4 ovate, lanceolate or filiform appendages. Gemmae lacking. Capsule opening by a lid. Spores 35-120 µm diam................................................................................................13

Bischler et al. 28 13. Antheridia in dorsal cushions bounded by conspicuous scales. Archegoniophores dorsal, stalk without rhizoid furrow................................................ 14. Plagiochasma 13. Antheridia dorsal, terminal or on small ventral branches, in cushions or in irregular groups, scales few or absent. Archegoniophores terminal, stalk with single rhizoid furrow.......................................................................................................................14 14. Ventral scales with wide appendages. Involucres cup-shaped or nearly flat .. ............................................................................................................................15 15. Radial walls of epidermal pores often thickened. Female receptacle subglobose when young, becoming conical, ovoid or inversely saucer-shaped when mature. Each archegonium enclosed in a large pseudoperianth splitting into linear segments ........................................................................................... ..........11. Asterella 15. Radial walls of epidermal pores thin. Female receptacle hemispherical. Archegonia not bounded by a pseudoperianth .............................. 13. Mannia 14. Ventral scales with filiform appendages. Involucres bilabiate.....................16 16. Epidermal cells without trigones. Radial walls of epidermal pores thin. Female receptacle hardly lobed. Spores with smooth areoles or with irregularly branching ridges on distal face .......................................... 12. Cryptomitrium 16. Epidermal cells with bulging trigones. Radial walls of epidermal pores usually strongly thickened. Female receptacle distinctly lobed. Spores with tuberculate areoles on distal face ................................................. 15. Reboulia 11. Epidermal pores compound (absent or vestigial in Dumortiera). Antheridiophores produced. Pseudoperianth campanulate (absent in Dumortiera). Capsule wall with annular thickenings.........................................................................................................................17 17. Epidermis, pores and air chambers absent or vestigial. Ventral scales small, without marginal papillae; no laminal scales. Bristles on ventral side of thallus and gametangiophores. Gemma-cups not produced. Archegoniophore stalks without assimilatory strips. Involucres tubular. Pseudoperianth lacking ............. 17. Dumortiera 17. Epidermis, pores and air chambers well developed, the pores compound. Ventral scales large, with marginal papillae, laminal scales conspicuous. Bristles lacking.

Bischler et al. 29 Gemma-cups produced. Archegoniophore stalk with assimilatory strip(s). Involucres bilabiate. Pseudoperianth campanulate................................................... .18. Marchantia

MARCHANTIALES

1. MARCHANTIALES Limpr., in F. Cohn, Krypt. Fl. Schlesien 1: 239, 336. 1877. Type. Marchantiaceae (Bisch.) Lindl. Gametophyte thalloid, growing prostrate, upper side often areolate. Apical cell cuneate, with four cutting faces. Branching dichotomous, ventral or by apical apical adventitious branches, especially after initiation of gametangial receptacles or gametangiophores. Thallus structurally complex, with epidermis and epidermal pores, assimilatory layer with air chambers, compact basal tissue often with mucilage cavities, and with oil cells without chloroplasts, containing a single, large oil-body. Ventral scales present, unistratose, in two or several rows (rarely in one row). Rhizoids of two types, pegged and smooth. Asexual multiplication by specialized diaspores infrequent, by pluricellular, discoid gemmae or by modified, fragmenting thallus segments. Monoecious or dioecious. Gametangia nonaggregated or clustered in grooves, cushions, or on receptacles that become elevated on stalks, the gametangiophores. Antheridia embedded in thallus or receptacular tissue, cylindrical-ellipsoidal, jacket unistratose except distally where 2 layers may be present, stalk usually short; ostioles formed by protruding thallus tissue. Archegonia embedded in thallus or receptacular tissue, with 4-8 neck canal cells, in cross section with 6 neck cell rows. Sporophytes radially symmetric, protected until maturity by several gametophytic envelopes: a) the calyptra (= archegonial wall), becoming usually 24-layered after fertilization, b) the involucre (of thalline origin) and, sometimes, c) a pseudoperianth (from archegonial stalk tissue). Seta short, sometimes necrotic, or lacking. Capsules globose, rarely ellipsoidal, wall unistratose except the apical cap or lid, with or without annular or semi-annular thickenings, or wall disintegrating prior to spore maturity; capsules opening irregularly or with apical lid, or cleistocarpous. Spores usually large, polar, the proximal face with triradiate scar, both faces ornamented. Elaters with helical thickenings, or absent.

1. OXYMITRACEAE Müll. Frib. ex Grolle, J. Bryol. 7: 215. 1972. Type. Oxymitra Bisch. ex Lindenb. The family is monogeneric.

1. OXYMITRA Bisch. ex Lindenb., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 14,

Bischler et al. 30 Suppl. 1: 124. 1829. Pycnoscenus Lindb., Oefvers. Kongl. Svenska Vet.-Akad. Förhandl. 19: 606. 1863, nom. illeg. Tessellina Dumort., Bull. Soc. Roy. Bot. Belgique 15: 164. 1874, nom. illeg. Lectotype (Howe 1923). O. paleacea Bisch. ex Lindenb. (= O. incrassata (Brot.) Sergio & Sim-Sim. Name from oxys = acute, mitra = helmet, referring to the pyriform involucres. Thallus 2.5-7 mm wide, often rosette-forming, dark green, often tinged with purple or dark red, with deep median groove apically; branching dichotomous. Epidermis delicate. Epidermal pores with single ring of 4-6 cells, without hyaline inner ring, radial walls often thickened. Air chambers narrow, in one layer, occupying 1/3-1/2 thallus height, without chlorophyllose filaments. Basal tissue without mucilage cavities. Oil-cells lacking. Ventral scales in 2 rows, acute apically, long, acuminate, projecting beyond thallus margins, in two rows, without oil-cells, with some marginal papillae. No asexual reproduction by specialized propagules. Dioecious or monoecious. Antheridia in dorsal groove along thallus midline, bounded by crests with cilia. Archegonia single per cavity, in ill-defined groups in dorsal thallus depression, delimited by crests with cilia. Calyptrae 2-3-layered after fertilisation. Involucres pyriform (in the South African species several involucres fused to form a crest) with epidermal pores and air chambers. Sporophytes one per involucre, lacking foot and seta. Capsules cleistocarpous, wall disintegrated at time of spore maturity. Elaters lacking. Spores 80-200 per capsule, 100-175 µm diameter, proximal and distal faces dissimilar (less so in the South African species), trilete scar distinct. Gametophytic chromosome number n = 9 (n = 18 in the South African species). A genus comprising two species growing on soil or rocky soil, forming soil crusts in intermittently dry, temperate to warm, Mediterranean-type climates. The plants are drought-tolerant and able to recover growth after long periods of drought. Oxymitra is disjunct between the Mediterranean and warmer parts of Europe, South Africa (the endemic O. cristata Garside ex Perold), SE United States, central and northern Mexico, and southern South America. The genus is infrequent in the Neotropics and has been recorded only from its northern and southern borders.

1. Oxymitra incrassata (Brot.) Sergio & Sim Sim, J. Bryol. 15: 662. 1989. Riccia incrassata Brot., Fl. Lusit. 2: 428. 1804. Type. Portugal. Near Coimbra (n.v.). Figs. 1: C, 3: D-E, 6, 7, 104: A-B Riccia pyramidata Raddi, Opusc. Sci. 2: 350. 1818. Oxymitra paleacea Bisch. ex Lindenb., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 14, Suppl.: 124. 1829. Oxymitra pyramidata (Raddi) Huebener, Hep. Germ. 24. 1834. Pycnoscenus pyramidatus (Raddi) Lindb., Oefv. Svenska Vetensk.- Akad. Förh. 19: 606. 1863. Tessellina pyramidata (Raddi) Dumort., Bull. Soc. Roy. Bot. Belgique 15: 164. 1874. Type. Italy, Florence, Raddi s.n. (n.v). Oxymitra androgyna M.Howe, Bryologist 17: 93. 1914. Type. U. S. A. Texas: Austin, Feb 1914, Young s.n. (NY).

Bischler et al. 31 Thallus greyish-green or dark green, often tinged with purple, dark red on margins and at base, 1-3 times branched, often forming hemi-rosettes; lobes up to 10 mm long, 2.5-7 mm wide, in cross section somewhat higher than wide; median groove deep. Epidermal cells thin-walled, pores bounded by 5-7 cells. Ventral scales white apically, large, extending beyond thallus margins. Dioecious. Archegonia enclosed in pyriform involucres with a short beak, bounded by white, filiform scales. Spores black, 80-200 per capsule, 100-175 µm diameter, distal face with 4-5 shallow, faintlytuberculate areoles across diameter, proximal face faintly tuberculate, triradiate mark distinct. Distribution and ecology. Common in the Mediterranean area, central Europe to 50° N, SW Asia, and Macaronesia, disjunct in southern U. S. A. (Texas, Oklahoma, Kansas) and South America (35°36°S). In the Neotropics rare, known from Mexico (D. F., Puebla (Schuster, 1992b), San Luis Potosí (Schuster, 1992b), Brazil (Rio Grande do Sul (Vianna, 1976)), Paraguay, Argentina (Buenos Aires, Catamarca, Córdoba, Tucumán (Hässel, 1963)), Uruguay. The species grows on periodically moist, compact, sandy soil or clay at base of rocks, on soil between calcareous or granitic blocks, exposed, in grassland or under open vegetation (shrubs, open forest), often associated with Riccia species and Sphaerocarpos, from sea level to 3000 m. Specimens examined. MEXICO. D. F.: Mexico city, Botanical Garden, 1975, Düll s.n. (PC); Pedregal de San Angel, University, Delgadillo 1646 (MEXU); Desierto de los Leones, Delgadillo 1345 (MEXU). PARAGUAY. Asunción, Balansa 1281 (G, NY); Cerro Yaguaron, Balansa 3709 (G, NY, PC). ARGENTINA. BUENOS AIRES: Sierra de la Ventana, Kühnemann 4157 b (BA). CATAMARCA: S. Ambato, Volk 28019 (BA). CORDOBA: Totoral, Las Peñas, Estancia la Maraña, Hosseus 16 (NY). URUGUAY. Montevideo, Cervio, Herter 99763 (G). The thallus of Oxymitra incrassata resembles Riccia but differs by the large, white, acute ventral scales that extend beyond the thallus margins and by the pyriform involucres that bound each sporophyte.

2. RICCIACEAE Reichenb., Bot. Damen 255. 1828. Type. Riccia L.

Gametophytes forming rosettes or gregarious patches. Thallus forked, often tinged with purple, red or violet, usually with deep apical groove; branching dichotomous. Epidermis well developed, or disintegrating, or absent. Epidermal pores with 1 (-2) rings of 4-9 cells, without hyaline inner ring, radial walls thin, or epidermal pores absent. Air chambers narrow and channel-like, or in 1-3 layers and wide, empty. Basal tissue thick, or reduced to 1-2 cell layers, without mucilage cavities. Oil-cells

Bischler et al. 32 present or absent. Ventral scales usually in 2 rows, or in single, median row, or in several ill-defined, transverse rows, or absent. Asexual reproduction sometimes by tubers. Monoecious or dioecious. Antheridia scattered or in dorsal groove, singly embedded in thallus, without scales or cilia. Archegonia scattered, or in 2-3 rows in dorsal groove, singly embedded in thallus. Calyptrae 2layered after fertilization. No involucres. Sporophytes without foot and seta. Capsules globose, cleistocarpous, dorsally or ventrally protruding, wall disintegrated at time of spore maturity. Spores large or medium sized (50-200 µm diameter), few per capsule, sometimes remaining in tetrads. No elaters. Gametophytic chromosome number n = 8, 16, or 24, or n = 9. The family includes two genera, one monospecific (Ricciocarpos), the other comprising at least 150 species (Riccia) and being the largest genus of the Marchantiales. Both genera occur in the Neotropics (54 species). Ricciaceae are widely distributed from cold to tropical areas on all continents. Most species are terrestrial, growing on soil and soil over rocks, often forming soil crusts, in open seepage areas, on margins of flooding zones of temporary ponds, in arable fields and other disturbed areas, some floating on stagnant water.

KEY TO THE NEOTROPICAL GENERA OF RICCIACEAE

1. Epidermal pores bounded by a ring of differentiated cells. Thallus and scales with oil-cells. Ventral scales in several ill-defined rows, short to very long, with serrulate margins. Antheridia and archegonia in dorsal groove. Gametophytic chromosome number n=9 ......................3. Ricciocarpos 1. Epidermal pores absent, or a simple opening among epidermal cells, rarely bounded by a ring of somewhat differentiated cells. Thallus and scales without oil-cells. Ventral scales absent or (usually) in 2 rows, short, margins not serrulate. Antheridia and archegonia scattered. Gametophytic chromosome number n=8, 16, or 24 ......................................................................................2. Riccia

RICCIA

by S. Jovet-Ast (based on the monograph of neotropical Riccia by S. Jovet-Ast (1991, 1993), adapted and translated from the French by H. Bischler).

Bischler et al. 33 2. RICCIA L., Sp. pl., ed. 1, 1138. 1753. Lectotype (Hässel de Menéndez, 1963). Riccia glauca L. Genus dedicated to the Italian senator and member of the Botanical Society of Florence, Pietro Francesco Ricci. Thallus forked, forming rosettes or gregarious patches; lobes 0.5-4 mm wide, with deep median groove apically, green, greyish green, yellow green of glaucous, often tinged with purple, red or violet; flanks sometimes with cilia. Epidermis in Neotropical species present, or disintegrating. Epidermal pores not differentiated or with 1(-2) rings of 4-9 cells with thin radial walls. Air chambers in one layer, narrow and channel-like, or in 1-3 layers and wide. Basal tissue thick or reduced to 1-2 cell layers. Oil-cells lacking. Ventral scales usually in 2 rows, without oil-cells or marginal papillae, without or with small appendage, not constricted basally, or scales in single, median row, or absent. Asexual reproduction by perennating tubers (several tubers may develop from the same thallus). Monoecious or dioecious. Antheridia and archegonia scattered. Sporangia dorsally or ventrally protruding. Spores 32-1350 per capsule, 40-200 µm diameter, sometimes remaining in tetrads, when free (not in tetrads) proximal and distal faces with more or less complete areoles, or with ridges and depressions, with or without tubercles on ridges and at corners, or spiny, or nearly smooth, trilete scar more or less distinct. Gametophytic chromosome number n = 8, 16, or 24. A genus of at least 150 species in 7 subgenera, drought-tolerant, sometimes perennating with tubers or tuberous apices, forming soil crusts on compact soil in open seepage areas in semi-arid to arid regions, but also in arable fields, gardens, along trails, on river borders and other disturbed habitats. Most species are terrestrial, some are floating on stagnant water. The genus is distributed worldwide, from the Arctic to the Antarctic, but the bulk of species occurs in areas with dry seasons. In the Neotropics, 54 species from four of the seven subgenera have been recorded (Riccia, 38 species; Ricciella, 14 species; Leptoriccia, one species; Thallocarpus, one species), from sea level up to 4500 m. Species diversity is highest in Brazil (32 species) and Mexico (21 species), somewhat lower in Paraguay, northern Argentina and Uruguay (20), Peru (16), and Central America south of Mexico (10), below 10 species in the other areas, lowest in Bolivia (1) and the Guianas (2). The data on frequency and regional distribution of the species are still very incomplete due to strong seasonality of the species, disappearing during the dry seasons and sometimes during several years, and the many undercollected areas. Species delimitation remains controversial in Riccia, precluding the use of unchecked bibliographic records. Only the most reliable references have been included in the data on geographical distribution.

KEY TO THE NEOTROPICAL SUBGENERA OF RICCIA

Bischler et al. 34 1. Thallus only 2-3 cell layers thick throughout. Thallus segments widened to the tips ................... ..................................................................................................................... 2b. subgen. Leptoriccia 1. Thallus thicker, at least in the middle ......................................................................................... 2 2. Dorsal surface of thallus without or with inconspicuous pores not surrounded by a ring of cells. Thallus tissue compact, discrete air chambers lacking (but narrow air channels present) ........... .........................................................................................................................2a..subgen. Riccia 2. Dorsal surface of thallus with distinct pores surrounded by a ring of cells, or with irregular openings. Thallus of loose tissue, air chambers present ............................................................ 3 3. Mature spores united in tetrads .................................................. 2c. subgen. Thallocarpus 3. Mature spores not united in tetrads ....................................................2d. subgen. Ricciella

2a. RICCIA subgen. RICCIA

Dorsal tissue with air channels. Epidermal pores a simple opening among partitions of air channels. Basal tissue thick. Epidermal cells usually disintegrating. Ventral scales in two rows. Sporangia bulging dorsally. Spore tetrads disintegrating at maturity of spores. Riccia subgen. Riccia is distributed over all continents. In the Neotropics this subgenus comprises 37 species, all of them members of the sect. Riccia (Jovet-Ast, 1991).

KEY TO THE NEOTROPICAL SPECIES OF SUBGEN. RICCIA 1. Lobes with papillae .............................................................................................................................2 2. Papillae numerous. Spores with 12-18 areoles across diam. of distal face....................................3 3. Papillae finger-like, present on dorsal side of lobes, margins and flanks, 60-210 µm long, smooth. Ventral scales without papilliform cells on margins ......................... 3. R. atromarginata 3. Papillae present only on margins and flanks of lobes, absent on dorsal side, 25-130 µm long, smooth or granulose. Ventral scales with papilliform cells on margins ......................................... ................................................................................................................................35. R. violacea 2. Papillae few. Spores with 16-25 areoles across diam. of distal face.................... 20. R. iodocheila 1. Lobes without papillae........................................................................................................................4 4. Lobes ciliate...................................................................................................................................5 5. Lobes densely ciliate, from lobe apex to base ..........................................................................6

Bischler et al. 35 6. Cilia smooth, to 100 µm long, or granulose, to 240 µm long. Spores 74-86 µm in diam., with 8-9 areoles across diam. of distal face, and smooth, triangular area around the pores on the proximal face............................................................................................ .....17. R. horrida 6. Cilia smooth, 400-1000 µm long. Spores 60-120 µm in diam. with 8-14 areoles across diam. of distal face; without differentiated area around the pores on the proximal face ......33. R. trichocarpa 5. Lobes with few, scattered cilia .................................................................................................7 7. Cilia rare, mainly on lobe margins. Proximal face of spores with smooth, triangular area surrounding the pores ...................................................................................30. R. subdepilata 7. Cilia in upper half of lobes, or from apex to base. Proximal face of spores without smooth triangular area surrounding the pores ......................................................................................8 8. Cross section of lobes with obliquely ascending, concave flanks. Spores 72-92 µm diam. ...................................................................................................................................9 9. Lobes deeply divided, not broadened apically.........................................13. R. enyae 9. Lobes shallowly divided above the basal fork, broadened apically............................. ................................................................................................................ 10. R. cubensis 8. Cross section of lobes with erect, convex flanks. Spores 84-120 µm diam..................10 10. Lobes with cilia in upper part, from top to bottom, flanks violet. Spores 84-108 µm diam., with 7-10(-11) areoles across diam. of distal face.......................................... ..............................................................................................................6. R. brasiliensis 10. Lobes with cilia apically and on margins, flanks hyaline, sometimes with violaceous spots. Spores 100-120 µm diam., with (10-)12-16 areoles across diam. of distal face.............................................................................................. 22. R. lindmanii 4. Lobes without cilia.......................................................................................................................11 11. Dorsal side of lobes calcified. Spores nearly smooth ........................ ....................1. R. albida 11. Dorsal side of lobes not calcified. Spores ornamented .........................................................12 12. Lobes with idioblasts (scattered cells with dense, white or coloured contents) ..............13 13. Distal face of spores areolate. Idioblasts with white contents, distinctly visible through epidermis (appearing as whitish dots) ..................................... ..2. R. albopunctata 13. Distal face of spores vermiculate. Idioblasts with orange-red contents, not visible through epidermis .................................................................................. 9. R. campbelliana 12. Lobes without idioblasts ..................................................................................................14

Bischler et al. 36 14. Ventral scales black, glossy........................................................................................15 15. Lobes 1-1.2 mm wide, lateral flanks black and glossy. Spores 60-80 µm diam., winged, with irregular or incomplete areoles............................................24. R. nigrella 15. Lobes 1.1-2.5 mm wide, lateral flanks dark red-violet. Spores 96-125 µm diam., winged or not winged, distal face tuberculate..............................................................16 16. Spores winged, tuberculate on both faces .................................. 5. R. boliviensis 16. Spores wingless, distal face areolate and with thick tubercles, proximal face with shallow and ill-defined tubercles .............................................29. R. squamata 14. Ventral scales hyaline or violaceous...........................................................................17 17. Cells of dorsal tissue thickened or with longitudinal, thickened strips..................18 18. Cells of dorsal tissue with thick, branched strips or with thickened walls........... ......................................................................................................... 23. R. mauryana 18. Cells of dorsal tissue with two longitudinal, thickened strips..........................19 19. Spores subspherical, distal face areolate at pole, proximal face granulose36. R. vitalii 19. Spores tetrahedral, areolate on both faces, areoles on proximal face often incomplete or ill-delimited ...............................................................................20 20. Lobe margins violet. Distal face of spores with 10-15 areoles across diam.............................................................................................26. R. ridleyi 20. Lobe margins light green or hyaline, dotted with pink-violet. Distal face of spores with 6-9 areoles across diam. .......................................................21 21. Flanks of lobes and scales violet. Proximal face of spores often with 10-15 incomplete areoles per facet ....................... ............37. R. weinionis 21. Flanks of lobes hyaline apically; scales with violet spots. Proximal face of spores with numerous ill-delimited areoles..............11. R. ekmanii 17. Cells of dorsal tissue without thickened longitudinal strips (sometimes with one or several thin or pitted strips)..........................................................................................22 22. Basal wall of epidermal cells granulose.................................... 23. R. mauryana 22. Basal wall of epidermal cells smooth.............................................................. 23 23. Cells of epidermal and/or hypodermal cells and/or flanks thickened.........24 24. Two cell layers below epidermis hyaline...............................................25

Bischler et al. 37 25. Epidermis and the two cell layers below epidermis hyaline, basal walls of epidermal cells and walls of the 4-5 hypodermal cell layers strongly thickened; flanks often with the external layer of cells thickened; cells of dorsal tissue without thin, longitudinal strips ................................. ................................................................................. 27. R. sanguineisporis 25. Only the two cell layers below epidermis hyaline, the upper one with thickened walls; cells of dorsal tissue with thin, longitudinal strips. 34. R. viannae 24. No hyaline cell layers below epidermis .................................................26 26. Lobes with hyaline margins. Basal walls of epidermal and hypodermal cells thickened.....................................................................27 27. Walls of hypodermal cells often yellow-tinged; flanks bordered by a strip of cells with thickened walls. Spores 100-140 µm diam., wingless......................................................................... ...8. R. brittonii 27. Walls of hypodermal cells not yellow-tinged; cells of flanks thinwalled. Spores 70-98 µm diam., winged, ...................28. R. sorocarpa 26. Lobes with light brown margins. Only basal walls of epidermal cells strongly thickened....................................................... 14. R. erythrocarpa 23. All cells of epidermis, flanks and dorsal tissue thin-walled .......................28 28. Scales white, extending beyond lobe margins .................21. R. lamellosa 28. Scales hyaline or tinged with violet, not extending beyond lobe margins, or absent .......................................................................................................29 29. Spores wingless, subspherical ...........................................................30 30. Lobes 1.5-3 times as wide as high. Tubercles not projecting on spore periphery; distal face with 12-16 areoles across diam................. .......................................................................................19. R. ianthina 30. Lobes 2-6 times as wide as high. Tubercles projecting on spore periphery; distal face with fewer than 12 areoles across diam. ........31 31. Lobes 2.5-3.8 mm wide. Spores with 4-6(-8) areoles across diam. of distal face ......... Cells of dorsal tissue with narrow, pitted longitudinal strip ......................................................31. R. subpana 31. Lobes 1.5-2.3(-3) mm wide. Cells of dorsal tissue Spores with 8-11 areoles across diam. of distal face. Cells of dorsal tissue

Bischler et al. 38 without pitted longitudinal strip....................................................... .....................................................................25. R. plano-biconvexa 29. Spores tetrahedral or subtetrahedral, with or without wing .............32 32. Lobes without hyaline margins. Spores winged...........................33 33. Thalli rosette-forming; lobes 2-4 mm wide. Spores 80-95 µm diam.......................................................................... ..16. R. grandis 33. Thalli in gregarious patches, or forming incomplete rosettes; lobes 1-2.2 mm wide. Spores (84-)96-125 µm diam.....................34 34. Spore wing brown, weakly or not crenulate.......................... ........................................................................ .....4. R. australis 34. Spore wing yellow, strongly crenulate.................................. ................................................................. ..32. R. taeniaeformis 32. Lobes with hyaline margins. Spores winged or wingless ............35 35. Spore wing (4-)7-9 µm wide ........................ .....15. R. fruchartii 35. Spore wing 4 µm wide or absent ............................................36 36. Spores 80-90 µm diam., winged, wing 4 µm wide, distal face with 4-6 areoles across diam. ........................12. R. elliottii 36. Spores 96-140 µm diam., wingless, distal face with 6-10 areoles across diam..................................................................37 37. Flanks of lobes subhorizontal .....................7. R. breutelii 37. Flanks of lobes obliquely ascending or subvertical .......... ......................................................................... 18. R. howellii

1. Riccia albida Sull. ex Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 231. 1870. Type. U. S. A. Texas: 1849, Wright s.n. (holotype, FH).

Figs. 8, 11

Thalli in partial rosettes, or in gregarious patches; lobes 2-3 times forked, 3-5 mm long, 0.8-1.8 mm wide, rounded apically or shallowly emarginate; dorsal side white, roughened by projecting epidermal cells covered with chalk, light green in groove, margins rounded, not inflexed when dry. Median groove deep, persistent to the lobe base. Epidermal cells bulging, thickened, conical or pyriform. Pores rounded, 30 µm diameter, or elongate, 25-35 x 50-60 µm. Ventral scales rare, small, or absent. Cross section of lobes 0.3-0.7 mm high, 1.5-2.5 times as wide as high; dorsal edge rounded laterally, flanks rounded, ventral edge flat or slightly convex; dorsal tissue in 2-4 layers, hyaline,

Bischler et al. 39 sometimes with pink strip basally; ventral tissue chlorophyllose, of smaller cells. Dioecious. Sporangia not bulging. Spores subspherical, 65-85 µm diameter, yellow-brown or red-brown, wingless, nearly smooth or weakly plicate and rugulose on both faces, triradiate scar indistinct. Distribution and ecology. Riccia albida is known from southern United States (Texas, Louisiana), Australia, and Mexico (Nuevo León). The single neotropical collection has been gathered on bare, calcareous, exposed soil, on a river border, at 880 m elevation. Specimen examined. MEXICO. NUEVO LEÓN: Junction of Monterrey bypass and Hwy. 40, ca. 16 mi. W of Monterrey, Whittemore et al. 1599 (Hb. Whittemore). Riccia albida is allied to the Mediterranean and Central Asiatic R. crustata Trab. The two species can be distinguished by their epidermal cells; those of R. albida are more distinctly bulging, and are separated by pores of different shape and size.

2. Riccia albopunctata Jovet-Ast, Cryptog. Bryol. Lichénol. 12: 237. 1991. Type. Ecuador. Galapagos Is.: Isabela, Volcano Alcedo, 1100 m, Gradstein H 218 (holotype, U).

Figs. 9, 11

Thalli in gregarious patches; lobes 2-3 times forked, 6-15 mm long, 1.3-2.5 mm wide, rounded apically; dorsal side pale green, or bright green with white dots, corresponding to the idioblasts distinct through epidermis; flanks black-violet, nearly completely inflexed when dry; ventral side green, in median portion with narrow strips dotted with pink-violet. Median groove deep apically, vanishing below. Epidermal cells rounded or pyriform, disintegrating except in groove. Ventral scales nearly black and glossy, or brown-violet, rarely only tinged with violet, not extending beyond lobe margins. Cross section of lobes 2-6 times as wide as high, dorsal edge acute laterally with violet, short wings; flanks obliquely ascending or nearly erect; ventral edge flat; dorsal tissue in 3-5 cell layers, each cell with a longitudinal, thin and sometimes pitted strip. Idioblasts scattered, large, 36-40 x 4876 µm, with white contents. Monoecious. Spores tetrahedral, 100-125 µm diameter, orange-brown or light red-brown; wing thin, light yellow, sometimes incomplete, with entire or crenulate margin; distal face with 8-10 areoles across diameter, with thin ridges and large, simple or furcate tubercles, to 9 µm high; proximal face with 20-30 tubercles per facet, triradiate scar well marked, consisting of irregular, often interrupted thickenings. Distribution and ecology. Riccia albopunctata has been recorded from the Galapagos Islands (Isabela), Brazil (Bahía, Mato Grosso, Paraná, Rio Grande do Sul, Santa Catarina, São Paulo), and Argentina (Misiones). The species is infrequent and has been collected on sandy or rocky soil, on river borders, sometimes on roadsides, under open vegetation, in caatinga, or in pampa, from sea level to 1100 m. Specimens examined. ECUADOR. GALAPAGOS: Isabela, Volcano Cerro Azul, Gradstein 440 (U);Volcano Alcedo, Gradstein 218 (U).

Bischler et al. 40 BRAZIL. BAHÍA: 14 km E of Morro do Chapen, Vital 6067 (SP); Jaguarari, Vital 8098 (SP). MATO GROSSO: Miranda, Vital 2366 (SP). PARANÁ: Barrancos de Painha, near Cachoeiras de Sete Quedas, Guaira, Vital 8364, 8372 (SP). RIO GRANDE DO SUL: Vacaria, Vital 2048 (SP); Tenente Portela, R. F. do Turvo, Rio Uruguai, Bueno 4255 , 4595 (ICN); Nova Preta, Rio Branco, Comada de Agro, Vianna 3306 (ICN); Arrio dos Ratos, Bueno 4603 (ICN); Sta Maria, Boca de Monte, Vianna 4560 (ICN); Ivoti, Cascata S. Miguel, Vianna 545 (ICN). SANTA CATARINA: Curitibanos, Vital 9415 (SP). SÃO PAULO: Pirajú, Vital 2181 (SP). ARGENTINA. MISIONES: Parque Nacional Iguazú, Grassi 119 (record based on Hässel, 1963, fig. 96). Riccia albopunctata is easily distinguished from the other neotropical Riccia species by its white idioblasts, visible through the epidermis as whitish dots. R. campbelliana, the other species with idioblasts occurring in the Neotropics, has different scales and a different spore wall ornamentation.

3. Riccia atromarginata Levier, in Martelli, Nuovo Giorn. Bot. Ital. 21: 291. 1889. Type. Italy. Sicily: Villa Belmonte near Palermo, Martelli & Ross s.n. (holotype, FI; isotype, BM). Figs. 10, 11, 78: A Thalli in partial rosettes or in crowded mats; lobes 2-3 times forked, 3-6 mm long, 1-1.8 mm wide, rounded apically, margins obtuse; dorsal side bright green or bluish green, sometimes with violaceous spots, margins dark violet; flanks black-violet, completely inflexed when dry. Median groove shallow, often wide. Epidermal cells disintegrating except in groove, rounded or pyriform. Papillae numerous, smooth, hyaline, finger-like, rounded apically, 60-210 µm long, always present on margins, often on flanks, and on dorsal side everywhere or near apex of lobes only. Ventral scales small, violet, margins entire. Cross section of lobes 0.6-0.7 mm high, as wide as high near apex of lobes, 1.5 times as wide as high below; dorsal edge rounded laterally; flanks erect, rounded apically. Dioecious. Archegonial necks protruding, dark violet. Spores tetrahedral, (75-)80-122 µm diameter, dark red-brown or nearly black, wingless; distal face with 12-18 areoles of irregular shape across diameter, with thick, nearly black ridges and large, black, sometimes anastomosed tubercles on corners; proximal face with similar ornamentation, triradiate scar weak. Gametophytic chromosome number n = 8 or 16. Distribution and ecology. Riccia atromarginata is widely distributed in the Mediterranean area, the Canary Islands, SW Asia, the Arabian Peninsula, and Iran. It has been recorded also from the southern United States (Texas). A single neotropical collection is known (Mexico, Nuevo León). The species colonizes usually sandy or clayey, exposed soil and soil over calcareous rocks in semi-arid and arid areas. Neotropical specimen examined. MEXICO. NUEVO LEÓN: 10 mi. S of Linares, Mc Gregor 8001 (NY).

Bischler et al. 41 The Mexican collection belongs to var. atromarginata. The species can be distinguished from the other neotropical Riccia species by its finger-like papillae, 60-210 µm long. Number, size and distribution of papillae vary. The species is closely related to R. violacea and R. iodocheila from which it differs by its numerous, long and smooth papillae. Schuster (1992b) treated R. iodocheila as a subsp. of R. atromarginata and R. violacea as a synonym of subsp. atromarginata. In our opinion, Riccia atromarginata, R. iodocheila and R. violacea are three distinct species.

4. Riccia australis Steph., Sp. hepat. 1: 29. 1898. Type. Uruguay. Montevideo, Fruchart 65 (holotype, G).

Figs. 12, 13

Thalli forming partial rosettes, or gregarious mats; lobes 2-3 times forked, 6-8 mm long, 1-2 mm wide, rounded, thick and weakly emarginate apically, thin below; dorsal side light green or greyish, sometimes with violet strip or violet veinules on margins, and violaceous spots above sporangia. Median groove shallow, distinct only apically. Epidermal cells rounded or pyriform. Ventral scales small, hyaline, violet, or violaceous. Cross section of lobes 3 times as wide as high near apex of lobes, 5-6 times as wide as high below, of 10-15 cell layers; dorsal edge rounded laterally or acute, with thin wings, often tinged with violet; flanks obliquely ascending; ventral edge slightly convex; dorsal tissue in two layers. Dioecious. Spores tetrahedral, (84-)96-120 µm diameter, light yellow-brown with brown ornamentation; wing 3-7 µm wide, irregular, papillose; distal face with 8-9(-10) areoles across diameter, limited by low and narrow ridges with large tubercles; proximal face similarly ornamented, with smaller tubercles, triradiate scar weak. Distribution and ecology. Riccia australis is distributed on the South American continent in Venezuela (Falcón), Ecuador (Pichincha), Peru (Loreto), Brazil (Paraná, Pernambuco, Rio Grande do Sul, São Paulo), Argentina (Misiones ?), and Uruguay. The species seems to be common in southeastern Brazil, rare elsewhere. It has been collected on sandy soil, or soil over sandstone, in exposed places, among paving stones, in gardens, open woodlands, on borders of paths and on water edges, from sea level to 2000 m. Specimens examined. VENEZUELA. FALCÓN: Sierra San Luís, 1.5 km W of Carrizalito, Wingfield 13987 (Hb. Wingfield). ECUADOR. PICHINCHA: road Quito to Sto Domingo de los Colorados, Gradstein et al. 6986 (U). PERU. LORETO: Maynas, E of Rio Sucasari, Timme 12917 (KSP). BRAZIL. PARANÁ: Guaira, Barrancos da Prainha, near Cachoeiras de Sete Quedas, Vital 8366 (SP). PERNAMBUCO: Saltinho Preserve, Porto 1375 (PC). RIO GRANDE DO SUL: Guaiba, Vital 2037 (SP); Vacaria, Vital 2047 (SP); Montenegro, Polotroquimico, Homrich 4605 (ICN); Sta Cruz do Sul,

Bischler et al. 42 Trombudo, Wechter 4606 (ICN); Gravatoi, Vianna 4473 (ICN); Taquara, Vianna 344 (ICN); Hamburgo Velho, Vianna 6 (ICN); Gramato, Vianna 1250 (ICN); Tenente Portela, R. F. do Turvo, Rio Murinci, Lindeman 6588 (U), Oliveira 1313, 1318 (ICN); Tenente Portela, Parque do Turvo, Bueno 4264 (ICN); Porto Alegre, Fac. de Direito, Vianna 12 (ICN); Porto Alegre, Av. Paulo Gama, Vianna 58723, 58733 (ICN); Porto Alegre, Ponta Grossa, Vianna 184, 3889, 6393 (ICN); Porto Alegre, Belem Novo, Praia do Ciego, Vianna 2651 (ICN); Porto Alegre, Jardim Schönwald, Homrich 2670 (ICN); Porto Alegre, Parque Farroupilha, Vianna 11 (ICN); Porto Alegre, Tristesa, Vianna 9 (ICN); Uruguaiana, 1978, Volk s.n. (Hb. Volk). SÃO PAULO: Serra do Mar, between São Paulo and Santos, Paranapiacaba, Schäfer-Verwimp & Verwimp 9004 (Hb. Schäfer-Verwimp); Restinga, Schäfer-Verwimp & Verwimp 7746 (Hb. Schäfer-Verwimp). ARGENTINA. Ignocii (Misiones?), Vianna 237 (ICN). URUGUAY. Type; Dep. Canelones, Santa Lucia, Herter 78902 (PC); S. José, Herter 86056 (PC). Riccia australis is allied to R. grandis. However, the thalli form complete rosettes in R. grandis, with lobes to 4 mm wide, and the spores are smaller, less than 95 µm in diameter.

5. Riccia boliviensis Jovet-Ast, Cryptog. Bryol. Lichénol. 12: 242. 1991. Type. Bolivia. Cochabamba: 3 mi. W Quillacolla-Cochabamba, 2560 m, Hermann 24590 (holotype, PC; isotype, US).

Figs. 14: A-K, 16, 78: B

Thalli in gregarious patches; lobes 2-3 times forked, 7-10 mm long, 1.5-2.5 mm wide, rounded apically; dorsal side pale greyish green, often with brown spots above sporangia; flanks violet, inflexed when dry; ventral side light green, dotted with violet, at least at apex of lobes. Median groove only apically distinct. Epidermal cells convex or pyriform. Ventral scales black-violet, glossy, imbricate, extending beyond lobe margins, often bordered by small, hyaline cells. Cross section of lobes 0.6-0.8 mm high, 3-4 times as wide as high; dorsal edge acute laterally, with short, black-violet wings; flanks obliquely ascending, ventral edge slightly convex; cells of dorsal tissue thin-walled. Monoecious. Antheridial necks hardly protruding, hyaline. Sporangia hardly bulging. Spores tetrahedral, 96-120 µm diameter, orange-brown or red-brown; wing 2.5-6 µm wide, light yellow, margin crenulate; distal face with rounded, glossy, orange tubercles or with sinuose ridges; proximal face with similar ornamentation, triradiate scar weak. Distribution and ecology. Riccia boliviensis has been recorded from Venezuela (Falcón, 140 m) and Bolivia (Cochabamba, 2560 m). The species seems to be rare; it has been collected on clay and sandstone, on borders of intermittent rivers, under shrubby vegetation and in thorn-woodland (JovetAst, 1991). Specimens examined. VENEZUELA. FALCÓN: El Bejuquero, 19 km SSW of Coro, at N foot of Sierra San Luís, Wingfield 13911 (Hb. Wingfield).

Bischler et al. 43 BOLIVIA. COCHABAMBA: Type. The nearly black, glossy scales of Riccia boliviensis are similar to those of R. albopunctata and R. squamata. However, in R. boliviensis idioblasts are lacking and spores have a crenulate wing.

6. Riccia brasiliensis Schiffn. & S.W.Arnell, Oesterr. Akad. Wiss., Math.-Naturwiss. Kl., Denkschr. 111: 6. 1964. Type. Brazil. Paraná: River Paranapanema, Salto Grande, 500 m, Schiffner 2395 (holotype, FH).

Figs. 14: L-S, 16, 78: C

Thalli in partial rosettes, or in gregarious mats, or 2-3 lobes grouped; lobes 2-3 times forked, 3-7 mm long, 0.6-0.9 mm wide, rounded, or obtuse, or emarginate apically, sometimes broadened behind apex; dorsal side light bluish green, margins obtuse, inflexed; flanks pink-violet; ventral side pale green. Median groove indistinct. Cilia few, hyaline, narrowly triangular, in upper part of flanks from top to bottom of lobes, more numerous apically, some short, 25-75 µm long, smooth, obtuse or rounded apically, others 80-240 µm long, 20-30 µm wide, subacute or obtuse apically, thin-walled, granulose in the upper 2/3. Ventral scales short, inserted below the cilia, pink-violet, cells 36-55 x 2430 µm. Cross section of lobes 0.4-0.5 mm high, 2-3 times as wide as high; dorsal edge slightly ascending and rounded laterally; flanks rounded, erect; ventral edge convex. Monoecious. Sporangia bulging. Spores 84-108 µm diameter, light red-brown; wing irregular, narrow, 2.5-3 µm wide, wider near the pores where it reaches 5-7 µm, granulose, or wing nearly absent; distal face with 7-10(-11) areoles across diameter, limited by conspicuous ridges with large tubercles, the areoles 8-10(-12) µm diameter; proximal face with smaller areoles but with similar ridges and tubercles, triradiate scar weak. Distribution and ecology. Riccia brasiliensis has been recorded from south-eastern Brazil only (Paraná, Pernambuco (Porto & Bezerra, 1996), Rio Grande do Sul, Santa Catarina, São Paulo). The species is infrequent and has been collected on exposed or partly shaded soil over rocks, on river borders, at low elevations. Specimens examined. BRAZIL. PARANÁ: Type; Guaira, Barrancos da Prainha, near Cachoeiras de Sede Quedas, Vital 8377 (SP). RIO GRANDE DO SUL: Porto Alegre, Morro Teresopolis, Vianna 2714 (ICN); Porto Alegre, Escola Agronomia, Vianna 4483 (ICN). SANTA CATARINA: From Lauro Müller to São Joaquim, near Bom Jardim da Serra, Vital 2004 (SP); Lajes, 19 km S of the city, along BR-116, Vital 5649 (SP). SÃO PAULO: Piraju, left margin of river Paranapanema, Vital 2176 (SP). Riccia brasiliensis is related to R. lindmanii but can be distinguished from the latter species by the presence of smooth as well as granulose cilia, and by the spores less than 108 µm in diameter, with no more than 11 areoles across the diameter of the distal face.

Bischler et al. 44 7. Riccia breutelii Hampe ex Steph., Sp. hepat. 1: 17. 1898. Type. Leeward Is. St. Kitts: Breutel s.n. (syntype, G).

Figs. 15, 16, 78: D

Riccia gaumeri Underw. ex M.Howe, in E.Britton et al., N. Amer. fl. 14 (1): 22, 1923, nom. inval. Type. Mexico. Yucatan: Without locality, Gaumer 1152 (holotype, NY). Thalli in gregarious patches; lobes 2-3 times forked, 5-10 mm long, 1-2.2 mm wide, thin and flattened, rounded and slightly emarginate apically; dorsal side light brown or brownish when dry, with narrow, hyaline, slightly undulate margins; flanks and ventral side pale. Median groove deep apically, vanishing below. Epidermal cells disintegrating. Ventral scales short, hyaline. Cross section of lobes 0.3-0.4 mm high, 2-5 times as wide as high; dorsal edge acute laterally, ending in a wing with 2-3 hyaline cells apically; flanks obliquely ascending, becoming horizontal in the wings; ventral edge convex; dorsal tissue in 2-4 cell layers, cells identical to those of ventral tissue. Dioecious? Spores subtetrahedral, 100-140 µm diameter, red-brown or dark brown, wingless or with incomplete, narrow, yellow strip on equator; distal face with 6-7 areoles across diameter in the smallest spores, with 7-10 in the largest spores, with thick, dark red or nearly black ridges and big tubercles; proximal face with similar or smaller areoles and narrower ridges, 20-25 per facet, triradiate scar weak. Distribution and ecology. Riccia breutelii is a neotropical species distributed in Mexico (Jalisco, Yucatan), Costa Rica, Cuba (Howe, 1923), Virgin Islands (San Juan (Howe, 1923)), Leeward Islands (St. Kitts), Venezuela (Falcón), Trinidad (Howe, 1923), Peru (Junín, Loreto), and Brazil (Amazonas). It has been collected on sandy soil and soil over limestone rocks, in deciduous woodlands and flood forest, from 50 to 4175 m. The species seems to be rare. Specimens examined. MEXICO. JALISCO: Puerto Vallarta, Howell 236 (NY). YUCATÁN: Type of Riccia gaumeri. COSTA RICA. Braxilito Bay, Howell 238, 239 (NY); Murcielago Bay, Howell 240 (NY). LEEWARD ISLANDS. ST. KITTS: Type. VENEZUELA. FALCÓN: 118 km WSW of Coro, Wingfield 14482 (Hb. Wingfield); Paraguara peninsula, Fila Monte Cano, above El Recreo, 2 km W of Pueblo Nuevo, Wingfield 14730 (Hb. Wingfield). PERU. JUNÍN: Jauja, La Oroya, Uchcupampa near Canchayllo, P. & E. Hegewald 5449 (Hb. Hegewald). LORETO: Maynas, along quebrada Yanomono, Timme 12351, 12954, 13833, 14289 (KSP). BRAZIL. PARÁ: Belém, Museu Paraense Emilio Goeldi, Vital 201-934 (SP). The Brazilian specimen has spores with 4-7 areoles only across diameter of distal face. Riccia breutelii, R. brittonii and R. elliottii are sometimes considered to be synonyms (Howe, 1923). Several characteristics keep the three species apart, however. In R. elliottii, the spores are relatively small (80-90 µm diameter) and the proximal face has only 4-5 areoles per facet. In R. brittonii, the

Bischler et al. 45 epidermal cells have thickened walls, and the flanks of the lobes show a violaceous strip of cells with thickened walls.

8. Riccia brittonii M.Howe, in E.Britton, Ann. Missouri Bot. Gard. 2: 50. 1915. Type. Puerto Rico. Mona Is, between Sardinero and Ubero, Britton et al. 1749a (holotype, NY). Figs. 117, 20, 78: E Thalli in gregarious patches; lobes simple or once forked, 2-5 mm long, 1-2 mm wide; dorsal side light green with narrow, hyaline margins; flanks brownish, or pink-violet, or violet. Median groove deep apically, vanishing towards base of lobes. Epidermal cells disintegrating, basal walls thickened, hypodermal cells with thickened, often yellow-tinged walls. Ventral scales very small, hyaline, not extending beyond lobe margins. Cross section of lobes twice as wide as high; dorsal edge laterally ascending and acute; flanks bordered by a strip of cells with thickened walls, brownish or violet in upper part, below hyaline; ventral edge flat or slightly convex; dorsal tissue in 4 layers. Monoecious. Spores subtetrahedral, 100-140 µm diameter, red-brown to dark brown, wingless but sometimes with an incomplete, yellow, narrow and granulose strip on equator; distal face with 6-9 areoles across diameter, granulose inside, with thick, dark red or black-red ridges and big tubercles as wide as the ridges; proximal face with areoles of same size than those of distal face, triradiate scar well marked. Distribution and ecology. Riccia brittonii has a limited range. It is known from Puerto Rico and was cited from the Virgin Islands (St. Thomas (Evans, 1918)). It has been collected on exposed soil, at low elevations. Specimen examined. PUERTO RICO. MONA ISLAND: Type. For the characteristics distinguishing Riccia brittonii from the allied R. breutelii and R. elliottii, see under R. breutelii. Riccia brittonii has a hyaline lobe margin and thickened epidermal and hypodermal cells as in R. sorocarpa. However, the wall thickenings are less conspicuous in R. brittonii, and the spore wall ornamentation is different.

9. Riccia campbelliana M.Howe, Mem. Torrey Bot. Club 7: 26. 1899. Type. U. S. A. California: Near Stanford University, 1896, Campbell s.n. (syntype, NY).

Figs. 18, 20, 78: F

Thalli in gregarious patches; lobes 1-3 times forked, 5-8 mm long, 1-2.5 mm wide, obtuse apically; dorsal side light green or brownish green, margins thin, membranaceous; ventral face brown, sometimes black. Median groove narrow apically, vanishing below. Epidermal cells convex. Idioblasts in epidermis and dorsal and ventral tissue, with glossy, dense, orange-red body, filling nearly the cell. Ventral scales imbricate, entire, lunate, extending beyond lobe margins, orange-red,

Bischler et al. 46 purplish or violet, cells 60-95 x 32-48 µm, marginal cells partly hyaline, partly orange-red. Cross section of lobes 2-4(-5) times as wide as high; dorsal edge convex laterally, acute, with short, thin wings ending in single cell; flanks ascending vertically or obliquely, abruptly narrowed into the wings; ventral edge convex. Monoecious. Antheridial necks protruding, colourless. Archegonial necks protruding, red or nearly black. Sporangia large, numerous, bulging. Spores tetrahedral, 75-120 µm diameter, yellow-brown or orange-brown; wing 4-5 µm wide, 6-7 µm at level of pores, light yellow, translucent, papillose; distal face vermiculate rather than areolate, with 8-15 ill-defined areoles across diameter, with inconspicuous ridges and numerous, rounded, contracted or branched tubercles or, sometimes, some slender tubercles only near equator; proximal face with similar ornamentation, but with smaller tubercles, triradiate scar narrow, conspicuous. Gametophytic chromosome number n = 8. Distribution and ecology. Riccia campbelliana has a very wide geographical distribution. It is known from South Africa, central Asia (Kazakstan), western Siberia, and has been recorded from the United States (Arkansas, California, Georgia, Kansas, Louisiana, Nebraska, Oklahoma, Texas). In the Neotropics its range extends from Mexico (Dept. Mexico) to Brazil (Rio Grande do Sul). The species is infrequent in the Neotropics; it has been collected on soil over cliffs and on roadsides. Neotropical specimens examined. MEXICO. MEXICO: 32.5 mi. W of Toluca, Whittemore 1839 (Hb. Whittemore). BRAZIL. RIO GRANDE DO SUL: Encruzilhada do Sul, Homrich 3148 (ICN). Riccia campbelliana is closely related to the Mediterranean Riccia macrocarpa Lev. Both species have idioblasts, but the lobes of R. macrocarpa are narrower (1 mm wide), nearly filiform, and only 7-8 areoles are seen across the distal face of the spores. One specimen has been recorded from Argentina (Tucumán: Tafi, La Cienaga, Sleumer 1755, reported by Hässel, 1963, n.v.). This specimen was subsequently described as Riccia campbelliana subsp. austrigena (Schuster, 1992a). Whether it really belongs to R. campbelliana remains questionable.

10. Riccia cubensis S.W.Arnell, Bryologist 61: 142. 1958. Type. Cuba. Prov. Oriente: Sierra Maestra, Daiquiri, Ekman 8348 (holotype, S).

Figs. 19, 20

Thalli forming partial rosettes or gregarious mats; lobes 2 mm long, 0.7-1 mm wide, thin, flattened when dry, rounded and broadened apically, 2-3 times forked, basal dichotomy deep, terminal shallow; dorsal side light green; ventral side pale, with narrow strips dotted with light pink-violet. Median groove deep apically, vanishing towards base of lobes. Epidermal cells disintegrating except in groove, subspherical. Ventral scales violet, imbricate, entire. Cilia few, on margins of dorsal side and upper part of flanks, numerous in the apical half of lobes, rare below, short, more or less arched, rounded, sometimes acute apically, 50-120 µm long, granulose to 3/4 of length or entirely granulose.

Bischler et al. 47 Cross section of lobes 1.5 times as wide as high near lobe apex, 2-3.5 times below; dorsal edge rounded or obtuse laterally; flanks obliquely ascending, with pink-violet spots apically. Dioecious. Spores (not seen, description after Arnell, 1958): 80-90 µm diameter, dark brown, wing incomplete, irregularly papillose, distal face with irregular lamellae. Distribution and ecology. The species is known from a single specimen, collected on bare soil in thickets on a coral reef. Specimen examined. CUBA. Type. The two other specimens cited by Arnell (1958) in the original description (Cuba, Ekman 8701 and 1155 (S)) belong to Riccia violacea var. violacea.

11. Riccia ekmanii S.W.Arnell, Bryologist 61: 143. 1958. Type. Cuba. Prov. Oriente, Daiquiri, Ekman 8432 (holotype, S).

Figs. 20: A-K, 23, 78: G

Thalli in gregarious patches; lobes 2-3 times forked, 8-10 mm long, 1.5-3 mm wide; dorsal side greenish, flattened and slightly concave when dry, rounded apically, margins acute, slightly undulate, hyaline and dotted with pink-violet. Median groove deep apically, shallower and vanishing below. Epidermal cells disintegrating except apically, convex. Ventral scales imbricate, entire, hyaline, dotted with pink-violet near margins, or entirely pink-violet, reaching or extending beyond lobe margins. Cross section of lobes 4-4.5 times as wide as high; dorsal edge acute laterally; flanks obliquely ascending, outer layer of larger cells than those of ventral tissue, tinged with pink-violet, with 1-2 hyaline cells apically; ventral edge convex; dorsal tissue in 3-5 cell layers, cells with two thickened, longitudinal strips. Monoecious. Antheridial necks colourless. Archegonial necks long, violaceous. Sporangia bulging mainly on ventral side. Spores tetrahedral, 90-120 µm diameter, redbrown, wingless, margins weakly crenulate; distal face with (6-)8-9 areoles across diameter, with thick, red-black ridges and large tubercles; proximal face with short ridges and tubercles of varying size and shape, ridges of triradiate scar discontinuous. Distribution and ecology. The sole specimen known of this species was collected on a coral reef exposed to salt water spray. Specimen examined. CUBA. Type. Riccia ekmanii is close to R. weinionis and R. ridleyi. It is characterized by hyaline lobe margins, dotted with violet at level of insertion of the scales. Its spores have 6-9 areoles across the diameter on the distal face, as in R. weinionis (not 10-12 as in R. ridleyi). The facets of the proximal face show 1012 areoles in R. weinionis, whereas in R. ekmanii they are more numerous and ill-delimited, with short, discontinuous ridges and tubercles of variable size and shape.

Bischler et al. 48 12. Riccia elliottii Steph., Sp. hepat. 1: 16. 1898. Type. Windward Islands. Dominica. Elliott 984 (holotype, G; isotype, BM).

Figs. 21: L-Q, 23, 78: H

Thalli in irregular, gregarious mats; lobes 2-3 times forked, 8 mm long, 1.5-2 mm wide, thin, rounded and slightly emarginate apically; dorsal side light brown or greenish brown when dry, light green when moist, with narrow, hyaline, slightly undulate margins; ventral side brown, violaceousbrown or reddish. Median groove deep apically, vanishing rapidly below. Epidermal cells disintegrating except in groove, rounded. Ventral scales not seen. Cross section of lobes 2-6 times as wide as high; dorsal edge acute laterally with hyaline wings; flanks obliquely ascending; ventral edge convex; dorsal tissue in 3-4 layers. Monoecious. Spores tetrahedral, 80-90 µm diameter, red-brown; wing 4 µm wide, light brown, granulose; distal face with 4-5(-6) areoles across diameter, granulose inside, with high, red-brown or nearly black ridges and large tubercles; proximal face with 3-6 areoles per facet, larger than those of distal face, with high, narrow ridges at pole, with lower ridges and weak tubercles, progressively vanishing towards equator, triradiate scar weak. Distribution and ecology. Riccia elliottii has been recorded from Mexico (Jalisco (Howe, 1934)), Costa Rica (Howe, 1934), and the Windward Islands (Dominica). Whether the records from the Virgin Islands (St. Thomas (Pagán, 1939)) and Puerto Rico (Mona Is. (Pagán, 1939)) belong to this species remains questionable. Specimen examined. WINDWARD ISLANDS. DOMINICA: Type. Riccia elliottii is allied to R. breutelii and R. brittonii. However, it differs from these two species by its thallus characteristics (see description) and by the spore wall ornamentation, with a smaller number of areoles across the distal and proximal faces.

13. Riccia enyae Jovet-Ast, Cryptog. Bryol. Lichénol. 12: 230. 1991. Type. Brazil. Rio Grande do Sul: Gravatoi, Itacolomi, Vianna 3075 (holotype, PC; isotype, ICN).

Figs. 22: A-N, 23, 78: I

Thalli forming partial rosettes or gregarious patches; lobes 10 mm long, 0.9-1.6 (-2.2) mm wide, sometimes slightly emarginate apically, 2-3 times deeply forked, the basal dichotomy reaching 7-9 mm; dorsal side pale green, margins rounded. Epidermal cells partly disintegrating, conical, thinwalled. Median groove shallow. Ventral scales hyaline, not extending beyond lobe margins, with slightly thickened walls, cells 30-38 x 24-25 µm. Cilia few, from top to bottom of lobes, on margins and in upper part of flanks, erect or slightly arched, finely verrucose in the upper 2/3, some acute, to 240 µm long, others, especially on the flanks, shorter, 60-112 µm. Cross section of lobes 0.4-0.5 mm high, 3-4 times as wide as high; dorsal edge obtuse laterally; flanks ascending obliquely and concave; ventral edge convex; cells of dorsal tissue of the same size as those of ventral tissue. Monoecious. Antheridial necks weakly protruding, hyaline. Archegonial necks weakly protruding, the young hyaline, the older red-violet. Sporangia weakly bulging on dorsal side. Spores 72-84(-92) µm

Bischler et al. 49 diameter, light red-brown; wing irregular, 4-5(-7) µm wide; distal face with (5-)6-8 areoles across diameter, with thin ridges and low, sometimes anastomosed tubercles; proximal face with numerous areoles, smaller than those of distal face, with thin ridges and small but distinct tubercles, triradiate scar weak. Distribution and ecology. Riccia enyae is known from north-eastern and southern Brazil (Paraíba (Gradstein & Pinheiro da Costa, 2003), Rio Grande do Sul) and seems to be rather common in Rio Grande do Sul. It has been collected on moist soil, in gardens and parks, at low elevation (0-200m). Specimens examined. BRAZIL. RIO GRANDE DO SUL: Type; Gravatoi, Itacolomi, Vianna 3160 (ICN); Tenete Portela, Parque do Turvo, Bueno 4263 (ICN); Santa Cruz do Sul, Trombudo, Wechter 7440 (ICN); Uruguaiana, 44 km from Porto Alegre, 1978, Volk s.n. (Hb. Volk); Porto Alegre, Av. Paulo Gama n° 40, Vianna 7566, 7567 (ICN); Viamão, Parque Saint-Hilaire, Vianna 2801 (ICN); Porto Alegre, Jardím Fac. de Direito, Vianna 7, 12 (ICN); Parque Fattoupi, Vianna 7571 (ICN). Riccia enyae is allied to R. horrida and R. cubensis. In R. enyae, cilia are less numerous than in R. horrida. The lobe dichotomies are always deep in R. enyae, whereas in R. cubensis only the basal fork is deep.

14. Riccia erythrocarpa Jovet-Ast, Cryptog. Bryol. Lichénol. 12: 257. 1991. Type. Brazil. Bahía: Mun. Urandi, Vital 7933 (holotype, PC; isotype, SP).

Figs. 24, 26, 78: K

Thalli in gregarious patches, with unpleasant smell; lobes 1-2 times forked, 4-6 mm long, 0.7-1.1 mm wide, obtuse apically; dorsal side light green with narrow, light brown margins; flanks inflexed when dry, violet; ventral side violet. Median groove shallow, only apically distinct. Epidermal cells disintegrating, basal walls strongly thickened. Ventral scales reaching lobe margins, hyaline, or violaceous with hyaline margins. Rhizoids pinkish. Cross section of lobes 0.9 mm high apically, 0.550.6 mm high in older parts, 1-1.5 times as wide as high; dorsal edge ending laterally in a peak of 2 hyaline uniseriate cells with thickened walls; flanks ascending and convex, with wide, red-violet strip in inner layers and narrow, pink-violet strip in outer layer; ventral edge convex; cells of dorsal and ventral tissue thin-walled. Dioecious? Spores subspherical, 100-136 µm diameter, red-brown or red, with dark black-red ornamentation, wingless; distal face with 10-12(-14) complete areoles across diameter, with thick, sometimes anastomosing ridges, or areoles incomplete with thick, sinuose ridges and large tubercles; facets of proximal face with subhexagonal areoles with thick ridges and more or less anastomosing tubercles, or with incomplete areoles and vermiculate thickenings, or with strong tubercles, triradiate scar weak. Distribution and ecology. The species is known from a single Brazilian specimen, collected in partial shade on the borders of a small pond, in caatinga vegetation. Specimen examined. BRAZIL. BAHÍA: Type.

Bischler et al. 50 Riccia erythrocarpa is distinguished by the cross section of lobes, which are nearly as wide as high, and the strong thickening of the basal walls of the epidermal cells. The species shares this latter characteristic with R. sorocarpa; however, in R. sorocapra all walls of the epidermal and hypodermal cells are thickened (in R. erythrocarpa only the basal wall of the epidermal cells). Additionally, the spores of the two species are different, having a conspicuous wing in R. sorocarpa but lacking one in R. erythrocarpa.

15. Riccia fruchartii Steph., Sp. hepat. 1: 22. 1898. Type. Uruguay. Montevideo, Fruchart 718 (holotype, G; isotype, PC).

Figs. 25: A-G, 26, 78: L

Riccia weberbaueri Steph. ex Weberb., Mundo Veg. Andes Peruanos 238. 1945, nom. inval. Type. Peru. Arequipa: Mollendo, 600 m, Weberbauer 1477 (holotype, G). Thalli in semi-rosettes or in crowded mats; lobes emarginate apically, 7-9 mm long, 2-3 mm wide, three times forked, the basal dichotomy deep, more than 1/2 of total length of lobes, younger dichotomies shallow; dorsal side pale green with wide, hyaline margins; flanks often pink. Median groove shallow, only apically distinct. Epidermal cells disintegrating, sometimes preserved in groove, rounded. Ventral scales hyaline, reaching the margins of lobes, cells 70-90 x 50-60 µm. Cross section of lobes with 15-20 cell layers in median part; dorsal edge slightly ascending laterally into the hyaline wings; flanks obliquely ascending or erect, abruptly extending into the wings, sometimes with a pinkviolaceous strip; ventral edge convex; cells thin-walled. Monoecious. Archegonial necks pink. Sporangia large, bulging, often 2-3 grouped. Spores tetrahedral, 95-108(-120) µm diameter, redbrown with dark brown ornamentation; wing (4-)7-9 µm wide, lighter in colour, granulose; distal face with 8-10(-12) areoles across diameter, with thick ridges and large, sometimes anastomosed tubercles, rarely near equator, ridges vanishing and tubercles alone persisting; proximal face with numerous areoles smaller than those of distal face, with thin ridges and small tubercles, triradiate scar thin, often not reaching the equator. Distribution and ecology. Riccia fruchartii is known from Peru (Arequipa, La Libertad), Brazil (Paraná, Rio Grande do Sul, Santa Catarina, São Paulo), Argentina (Buenos Aires, Hässel, 1963), and Uruguay. The species seems to be quite common in south-eastern Brazil; it has been collected on bare, exposed, sandy, granitic or basaltic soil, in gardens, along paths and roadsides, on drainage borders, from sea level to 2690 m in Peru. Specimens examined. PERU. AREQUIPA: Type of Riccia weberbaueri. LA LIBERTAD: Otuzco, Huancamarca, Quebrada Hornillo, P. & E. Hegewald 5147 (Hb. Hegewald); Otuzco, Coina, P. & E. Hegewald 5264 (Hb. Hegewald). BRAZIL. PARANÁ: Barrancos da Prainha, Cachoeiras de Sete Quedas, Guaira, Vital 8366 (SP). RIO GRANDE DO SUL: Cachoeira do Sul, Vital 9229, 9240 (SP); Canela, Vital 9287 (SP); São

Bischler et al. 51 Lourenço, Vital 8956 (SP); Xui, Vital 9018 (SP); Pedro Osorio, Vital 9124, 9125 (SP); Baje, Vital 9161 (SP); Canela, Vianna 3 (ICN); Porto Alegre, Ponta Grossa, Vianna 183 (ICN); Torres, Vianna 635 (ICN); Rosario do Sul, Dardano de A. Lima 2763 (ICN); Porto Alegre, Escola de Agronomía, Vianna 3085, 7442 (ICN); Osorio, Estrada Praia Presidente, Vianna 3994 (ICN); Porto Alegre, Belém Novo, Praia do Cego, Vianna 176 (ICN); Nonoai, Vianna 4607 (ICN). SANTA CATARINA: Bom Jardím da Serra, Lauro Müller, Vital 2006a (SP). SÃO PAULO: São Paulo, Cidade Universitaria, Vital 8499 (SP); São Paulo, IBT, around the greenhouses, Vital 11326 (SP). URUGUAY. Type. Riccia fruchartii is related to R. glauca L. but in the latter species the margins of the thallus lobes are not hyaline.

16. Riccia grandis Nees, in Martius, Fl. bras. 1: 300. 1833. Type. Brazil. Rio de Janeiro: near Rio de Janeiro, St. Hilaire s.n. (holotype, PC).

Figs. 25: H-L, 26, 78: M

Riccia flavispora Steph., Sp. hepat. 1: 17. 1898. Brazil. Without locality, Ule 219 (holotype, G) Thalli in rosettes of 10-15 mm diameter; lobes thin, flat, 2-3 times forked, with deep dichotomies, 6 mm long, 2-4 mm wide, obtuse and shortly bilobed apically; dorsal side greenish; ventral side pale green. Median groove shallow, only apically distinct. Epidermal cells disintegrating. Ventral scales hyaline or violet, extending beyond lobe margins. Cross section of lobes 3-6 times as wide as high, with 12 thin-walled cell layers of which 4 are dorsal tissue; dorsal edge acute or obtuse laterally; flanks indistinct, the hardly convex ventral edge extending progressively into the flanks. Monoecious. Antheridial necks protruding, white. Sporangia hardly bulging, light yellow. Spores tetrahedral, 80-95 µm diameter, reddish brown; wing light brown, irregular, 5 µm wide, granulose; distal face with 8-10 areoles across diameter, with narrow ridges and thin, high tubercles; proximal face with numerous complete areoles with low, wide ridges and tubercles, triradiate scar with wide ridges not reaching the equator. Distribution and ecology. Riccia grandis has been recorded only from south-eastern Brazil (Rio Grande do Sul, Rio de Janeiro, São Paulo). It has been collected on sandy soil, in cultivation, at low elevations. Specimens examined. BRAZIL. RIO DE JANEIRO: Type. RIO GRANDE DO SUL: Viamão, Vianna s.n. (ICN). SÃO PAULO: Litoral, Itanhaem, Fazenda Aurea on Rio Branco, Schäfer-Verwimp & Verwimp 11137 (Hb. Schäfer-Verwimp). Riccia grandis is somewhat related to R. sorocarpa but the spores in R. grandis have complete areoles on the proximal face, whereas in R. sorocarpa, the proximal face is granulose-tuberculate, not areolate.

Bischler et al. 52

17. Riccia horrida Jovet-Ast, Cryptog. Bryol. Lichénol. 12: 226. 1991. Type. Brazil. Espíritu Santo: Linhares, Rio Doce, Vital 1969 (holotype, PC; isotype, SP).

Figs. 27: A-H, 28

Thalli forming rosettes of 12 mm diameter; lobes 2-3 times forked, 3-6 mm long, 0.8 mm wide, rounded apically; dorsal side light green or light brown; flanks violet. Median groove wide. Epidermal cells disintegrating. Ventral scales violaceous, not extending beyond lobe margins. Cilia numerous, inserted on top of flanks and on lobe margins, sometimes in groove, some short (less than 100 µm long), smooth, most granulose, to 240 µm long, both types acute, stiff, erect or oblique, some intercrossing over the lobes. Cross section of lobes 0.4 mm high at apex, 0.2-0.3 mm near base; dorsal edge rounded laterally; flanks ascending; ventral edge convex. Dioecious? Spores spherical or tetrahedral, 74-86 µm diameter, red-brown with red-brown or dark red-brown ornamentation; wing 7 µm wide, 12-15 µm wide near the pores, slightly papillose and crenulate on margin, pores bounded by a triangular, smooth area; distal face with 8-9 areoles across diameter, each of 6-8 µm diameter, with rather thin ridges and weak tubercles; proximal face with numerous, smaller areoles, with thin ridges and small tubercles, triradiate scar weak. Distribution and ecology. Riccia horrida has been collected twice, in Venezuela (Infante-Sanchez & Heras Pérez, 2002) and Brazil (Espíritu Santo). The type was found on sandy soil on islets in a river bed, at low elevation. Specimen examined. BRAZIL. ESPÍRITU SANTO: Type. The abundance of cilia distinguishes Riccia horrida from all other neotropical Riccia species. The cilia are of two types, smooth ones to 100 µm long and granulose ones to 240 µm long. The scales are violaceous. The spores are rather small, 74-86 µm in diameter, and have a characteristic, smooth triangular area bounding the pores on the proximal face. The latter characteristic is shared with R. subdepilata but this species has less abundant and exclusively granulose, 120-380 µm long cilia, colourless scales, and larger spores (84-96 µm).

18. Riccia howellii M.Howe, Proc. Calif. Acad Sci. ser. 4, 21: 202. 1934. Type. Ecuador. Galapagos Is.: Isabela, Iguana Cove, Howell 209 (isotype, NY).

Figs. 22: O-W, 28, 79: A

Thalli in gregarious patches; lobes 3-4 times forked, 5-10 mm long, 0.9-1 mm wide apically and rounded, progressively narrowed and 0.7 mm wide basally; dorsal side pale green, greyish or whitish when dry, bright green when moist, with narrow, hyaline or light brown margins; flanks violaceous or violet; ventral side brown or violaceous. Median groove deep, vanishing towards base of lobes. Epidermal cells disintegrating. Scales hyaline or violeaceous-brown, not extending beyond lobe margins. Cross section of lobes 1-3 times as wide as high; dorsal edge laterally acute, with short wings; flanks obliquely ascending or erect; ventral edge convex; dorsal tissue in 2-4 layers.

Bischler et al. 53 Monoecious. Spores subtetrahedral, 96-120(-132) µm diameter, red-brown to violet-brown, wingless but sometimes with an incomplete, light brown, narrow, granulose strip at equator; distal face with 68(-9-10) complete areoles across diameter, with thick ridges and conspicuous tubercles; proximal face with similar areoles and ridges, triradiate scar weak. Gametophytic chromosome number n = 8. Distribution and ecology. Riccia howellii is known only from the Galapagos Islands (Española, Floreana, Isabela, Pinta, San Cristóbal, Santa Maria), where it seems to be fairly common. It has been collected on sandy or clayey soil among rocks, shaded by open forests, or along trails, at low elevations (50-250 m). Specimens examined. ECUADOR. GALÁPAGOS IS. ESPAÑOLA: Howell 184 (NY). FLOREANA (=SANTA MARIA): above Black Beach, Gradstein H 132 (U); Howell 247 (NY). ISABELA: Type; W of Tagus Cove Mt., Howell 219, 220 (NY); James Bay, Howell 187, 188 (NY); Volcano Cerro Azul, above Iguana Cove, Gradstein H 459a, b (U). PINTA: Gradstein H 489 (U). SAN CRISTÓBAL: Howell 203, 204, 205 (NY). Riccia howellii belongs to the species group with lobes bordered by a hyaline margin. The spores are wingless, as are those of R. subplana, but sometimes a light brown belt around the spore connecting the protruding tubercles is present.

19. Riccia ianthina Jovet-Ast, Rev. Bryol. Lichénol. 44: 418. 1978. Type. Ecuador. Galapagos Is.: San Salvador, 660 m, Pike 2762 (holotype, U).

Figs. 27: I-W, 30, 79: B

Thalli in gregarious patches; lobes 2-3 times forked, 6 mm long, 2 mm wide, rounded or truncate apically, 1.5-1.7 mm wide beyond the apical dichotomy; dorsal side bright green when moist, pale bluish-green when dry, violaceous in older parts and on both sides of median groove, with light green strips on margins; flanks inflexed, violet; ventral side violet. Median groove narrow and deep apically, flattened towards base. Epidermal cells slightly concave, sometimes with median mamilla, persistent in the groove and on the flanks of the groove. Ventral scales large, not extending beyond lobe margins, pink or pink-violet, margins entire, with 2 rows of hyaline cells. Cross section of lobes 1.5-3 times as wide as high; dorsal edge acute laterally; flanks obliquely ascending; ventral edge convex; dorsal tissue 1/3-1/2 of lobe height, in 6-9 layers, cells with thin, pitted, longitudinal strips. Monoecious. Necks of antheridia and archegonia hardly protruding. Spores nearly spherical, 96-108 µm diameter, red-brown or tinged with pink-violet, wingless; distal face with (12-)16 well delimited areoles across diameter, each 6-8 µm diameter, smaller near pole than at periphery, with conspicuous tubercles; proximal face with similar ornamentation, triradiate scar indistinct. Distribution and ecology. Riccia ianthina is known from a single specimen collected on the Galápagos Islands, on flat slabs of volcanic rocks, near a water pool, at 650 m. Specimen examined. ECUADOR. GALAPAGOS ISLANDS: San Salvador, type.

Bischler et al. 54 Riccia ianthina is related to R. iodocheila and R. violacea. However, in R. iodocheila and R. violacea the lobe margins, flanks and, in R. iodocheila, lobe apex have papillae and the margins of the scales are provided with violaceous papillae and short, hyaline setae. Moreover, in the latter two species tubercles are lacking at the corners of the areoles on the distal face of spores (present in R. ianthina).

20. Riccia iodocheila M.Howe, Proc. Calif. Acad. Sci. ser. 4, 21: 200. 1934. Type. Ecuador. Galapagos Is.: San Cristóbal, Wreck Bay, Howell 201 (holotype, NY). Riccia atromarginatar Levier subsp. iodocheila (M.Howe) R.M. Schust., The Hepaticae and Anthocerotae of North America 6: 623. 1992

Figs. 29, 30

Thalli in gregarious patches, not forming rosettes; lobes 2-3 times forked, 3-5 mm long, 0.5-1.4 mm wide, rounded or obtuse apically, margins acute; dorsal side bluish green; flanks black-violet, inflexed; ventral side light greyish-green. Median groove shallow or absent. Epidermal cells rounded or pyriform, disintegrating except in groove. Papillae sometimes present on margins and near lobe apex. Ventral scales small, black-violet, of few cells, crenulate by violet projecting marginal cells, and with papilliform cells 60-65 µm long, violet or violaceous basally, becoming hyaline towards apex. Cross section of lobes 0.4-0.7 mm high, as wide as high in apical part of lobe to twice as wide as high beyond; dorsal edge rounded laterally; flanks ascending vertically; ventral edge convex. Monoecious. Spores tetrahedral, 70-107 µm diameter, dark brown or nearly black, wingless, crenulate on margin; distal face with (16-)20-25 areoles across diameter, each of 3-6 µm diameter, with thick ridges; proximal face with similar ornamentation, triradiate scar weak. Distribution and ecology. Riccia iodocheila is known from southern United States (California, Texas) and in the Neotropics from Mexico (Baja California, Oaxaca, Socorro Is., Sonora (Howe, 1934)), the Galapagos Islands (San Cristóbal, San Salvador (Gradstein & Weber, 1982)), and Argentina (Tucumán, Salta, La Rioja, Córdoba (Hässel, 1963)). The species seems to be rare; it has been collected on silty soil and on rocks, at the borders of intermittent rivers, from sea level to 380 m. Specimens examined. MEXICO. BAJA CALIFORNIA: Bahia de los Muertos, Golfo de California, Steere 17562 (NY); Golfo de California, Steere 17563 (NY); Golfo de California, near SW corner of Ceralvo Is., Steere 17567 (NY). OAXACA: between Teotitlan and Cuitlapan, 14 km before Cuitlapan, Düll 2/230, 2/232 (Hb. Düll). SOCORRO ISLAND: Bosque de Guayabillo, Colima, 1958, Herrera s.n. (JE). ECUADOR. GALAPAGOS IS.: San Cristóbal, type. Riccia iodocheila is closely related to the Mediterranean Riccia trabutiana Steph. The presence of papillae on the scale margins in R. iodocheila, however, separates the species from R. trabutianaa as well as from R. atromarginata.

Bischler et al. 55

21. Riccia lamellosa Raddi, Opusc. Sci. 2: 351. 1818. Type. Italy. Florence, without collector (holotype, PI; isotype, PC).

Figs. 2: E, 31, 32, 79: C

Riccia grandisquama Steph., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 23 (III, 2): 28. 1897. Type. Brazil. Rio Grande do Sul: Cachoeira, Lindman 216 (holotype, G, without spores). Riccia lamellosa americana M.Howe, Bull. Torrey Bot. Club 25: 189. 1898; Riccia americana (M.Howe) M.Howe, Mem. Torrey Bot. Club 7: 24, 1899 p.p. Type. U. S. A. California: San Francisco, Bolander s.n., Howe s.n., etc. (syntypes, NY). Riccia austinii Steph., Sp. hepat. 1: 28. 1898. Type. U. S. A. New Jersey: Closter, Austin s.n. (holotype, G). Riccia indusiata S. Winkler, Rev. Bryol. Lichénol. 42: 789. 1976. Type. Colombia. Magdalena: Sierra Nevada de Santa Marta, Pico Bolivar, 4200 m, Winkler C 295 (holotype, ULM, slide), syn. fide Gradstein (1998). Thallus forming complete or partial rosettes, or gregarious patches; lobes 20 mm long, 2-4 mm wide, rounded apically, 2-4 times forked, basal dichotomy deep, reaching 3/4 of lobe length; dorsal side pale green or bluish, with acute, pale margins; flanks inflexed, often with purplish spots. Median groove shallow and wide, persistent to lobe base. Epidermal cells convex, disintegrating, except in groove; usually two layers of hyaline cells beyond the epidermis. Ventral scales white, extending beyond lobe margins, semi-lunate, imbricate, of large cells (135-145 x 52-80 µm), margin entire, of smaller cells (100 x 72 µm). Cross section of lobes as wide as high near thallus apex, 1.5-2 times as wide as high below; dorsal edge laterally ascending and forming short, thin wings on both sides; flanks erect, extending into the short wings; ventral edge convex or plano-convex. Monoecious. Antheridial necks hardly protruding, pale. Sporangia large. Spores tetrahedral to nearly spherical, 76130 µm diameter, red-brown or dark brown; wing 2-4 µm wide, yellow or light brown with thickenings, margin crenulate, or incomplete, or absent; distal face with 8-15 complete areoles across diameter, with thick ridges and tubercles; sometimes, areoles replaced near equator by straight ridges, or areoles only near pole, with tubercles near equator; proximal face with numerous and smaller areoles with low ridges and dark tubercles, with weak triradiate scar. Gametophytic chromosome number n = 8, 16, 24. Distribution and ecology. Riccia lamellosa is widely distributed in central Europe and the Mediterranean area, Macaronesia (Canary Is., Madeira), dry tropical and subtropical Africa (Ethiopia, Central Africa, Tanzania, Kenya, South Africa), central and SW Asia (Syria, Kazakhstan, Tajikistan, Pakistan), and Australia. It is known from Canada, and from the United States from most states. In the Neotropics, it has been recorded from Mexico (Baja California (Schuster, 1992b), D. F., Mexico, Puebla, Querétaro, Zacatecas), Colombia (Winkler, 1976, as R. indusiata; Gradstein, 1998), Peru (Pasco (Bryan, 1929)), Brazil (Rio Grande do Sul), Argentina (Catamarca (Müller, 1955), Tucumán

Bischler et al. 56 (Hässel, 1958), La Rioja (Hässel, 1958), Buenos Aires (Hässel, 1958)), and Uruguay. The species is rare in the Neotropics; it has been collected in semi-arid zones on calcareous or siliceous, sandy soil or clay, among rocks and lava, exposed in grasslands or along paths, or sheltered in open, scrubby vegetation or woodland, from sea level to 4200 m. Specimens examined. MEXICO. D. F.: Mexico city, Botanical Garden, Düll 2/3A (Hb. Düll). MEXICO: 8 km E Rio Frio Pass, Düll 2/98 (Hb. Düll); Sombrereto, Volk 6/0728 (Hb. Volk). PUEBLA: Road Tehuacan-Alcuntzinga, 25 km E Tehuacan, Düll A 71 (Hb. Düll). QUERÉTARO: N of Cadereyta, Düll 4 (Hb. Düll). ZACATECAS: Mexico Hwy., 49.6 mi. W of Zacatecas, Reveal 2663 (NY); 19.5 mi. N of San Tiburcio, Whittemore 1600 (Hb. Whittemore). COLOMBIA. CESAR: Type of Riccia indusiata S. Winkler. BRAZIL. RIO GRANDE DO SUL: Cambara do Sul, Vital 2026 (SP); Uruguaiana, 44 km from Porto Alegre, 1978, Volk s.n. (Hb. Volk). URUGUAY. Tacuarembo, Tambores, Herter 9, 8444 (PC); Durazno, Blanquillo, Herter 99729 (PC). Riccia lamellosa is easily recognised by its pale green or bluish colour and by its white scales covering the flanks and extending beyond the lobe margins. Variation is observed in the degree of imbrication of the scales, scale colour (usually white but sometimes with pink, violaceous or brown spots), spore diameter (80-84 to 98-120 µm), the spore wing, which may be wide (3-6 µm) or incomplete to absent, and the number of areoles across distal spore faces (8-9 to 10-15). The morphological variation observed is not geographical and but may correlate with ecology as the species occurs in a large variety of habitat conditions, and also with polyploidy (n = 8, 16 or 24). The genetic structure of R. lamellosa needs investigation.

22. Riccia lindmanii Steph., Bih. Kongl. Svenska Ventensk.-Akad. Handl. 23 (III, 2): 29. 1897. Type (Jovet-Ast, 1991). Paraguay. Asunción, Lindman 220a (lectotype, S).

Figs. 33, 36, 79: D

Thalli forming rosettes 6-10 mm in diameter, or in gregarious patches; lobes 2-4 times deeply forked, 2.5-3 mm long, 0.5-1(-1.5) mm wide, flat and thin basally, rounded or slightly emarginate apically; dorsal side light brown or greyish when dry; flanks hyaline, often with pink-violaceous spots. Median groove shallow, wide. Ventral scales hyaline or pink-violaceous, short, not reaching lobe margins. Cilia few, on lateral borders of dorsal side and in upper part of flanks, numerous near apex of lobes, isolated or absent below, 60-215 µm long, 30-35 µm wide basally, acute or obtuse apically, granulose to 1/2-2/3 of length. Cross section of lobes 0.6 mm high, 7-8 cells thick; dorsal edge ascending and rounded laterally; flanks rounded apically; ventral edge convex; dorsal and ventral tissues hardly distinct. Monoecious. Spores (85-)100-120 µm diameter, orange-brown or light red-brown; wing 4-5 µm wide, irregular, sometimes narrow or nearly absent, translucent, margins

Bischler et al. 57 crenulate-mamillose; distal face with (10-)12-16 areoles across diameter, each of 10-15 µm diameter, with dark ridges often thickened near pole and large tubercles; proximal face with numerous areoles of 10 µm diameter, with low ridges, the ridges ill-defined, papillose and tuberculate, triradiate scar weak. Distribution and ecology. Riccia lindmanii is a neotropical species known from Peru (La Libertad), Brazil (Mato Grosso do Sul, Rio Grande do Sul), Paraguay, Argentina (Buenos Aires, Catamarca, Tucumán (Hässel, 1963)), and Uruguay (Hässel, 1963). It seems to be infrequent and has been collected on exposed, clayey or sandy soil, on river borders, in parks and on street borders, from sea level to 2050 m. Specimens examined. PERU. LA LIBERTAD: Otuzco, Coina, P. & E. Hegewald 5230 (Hb. Hegewald). BRAZIL. MATO GROSSO DO SUL: Corumbo, Pantanal, Fazenda Santa Clara, Schäfer-Verwimp 8455 (Hb. Schäfer-Verwimp). RIO GRANDE DO SUL: Pedro Osorio, Vital 9126 (SP); Tenente Portela, Parque do Turvo, Bueno 4263 (ICN); Caixas do Sul, Vianna 7575 (ICN); Ivoti, Cascata S. Miguel, Vianna 544 (ICN). PARAGUAY. Type; El Chaco, near Asunción, Lindman 250 (G), Lindman 251 (syntypes, S). Riccia lindmanii is related to R. brasiliensis but the spores of the two species are different, those of R. brasiliensis being larger, to 120 µm in diameter, with up to 16 areoles across diameter of distal face. The cilia in R. lindmannii are rather abundant and their length varies. The specimens recorded by Vianna (1985) have smaller spores (77-92 µm) and the number of areoles across the diameter of the distal face is only 8-11.

23. Riccia mauryana Steph., Sp. hepat. 1: 19. 1898. Type. D. F.: Cerro de Guadalupe, Maury (3228) (holotype, G).

Figs. 34, 36, 79: E

Riccia andina Müll. Frib., Feddes Repert. 58: 65. 1955. Type (Hässel, 1963). Argentina. Catamarca: dep. Andalgala, Aconquija (Sierra de las Canas), 2600 m, Sleumer 2141 (lectotype, S; isolectotype, LIL). Riccia dorsiverrucosa Hässel, Opera Lilloana 7: 261. 1963. Type. Argentina. Catamarca: dep. Andalgala, Alto de las Juntas-Las Rosas, ca. 1600 m, 25 Dec 1951, Sleumer 2099 (holotype, LIL). Thalli in gregarious patches; lobes 2-3 times forked, 3-10 mm long, 1-2.5(-3) mm wide, rounded apically; dorsal side light green to greyish green, with narrow, hyaline margins; ventral side pale green or brownish. Median groove deep. Epidermal cells disintegrating, ovate-rounded where intact, with basal and lateral walls somewhat thickened, the basal granulose; hypodermal cells with more or less thickened walls. Ventral scales hyaline, imbricate, lunulate, entire, of large cells (60-72 x 36-50

Bischler et al. 58 µm), bordered by smaller cells. Cross section of lobes 2-6 times as wide as high, 0.5-0.7 mm high; dorsal edge acute laterally; flanks obliquely ascending, or slightly concave, recurved and acute apically; ventral edge convex; dorsal tissue in 6-8 cell layers of large cells with two thickened walls or with thick, branched strips and a hyaline, thin, longitudinal strip. Dioecious? Spores subspherical, 67108 µm diameter, red-brown with darker ornamentation, wingless, but sometimes with a narrow, incomplete and irregular, hyaline or yellowish strip; distal face with 7-14 areoles across diameter, ridges thick, more or less hidden by conspicuous tubercles; proximal face with incomplete areoles with low ridges and small tubercles, triradiate scar weak. Distribution and ecology. Riccia mauryana is a species known from southern United States (Texas (Schuster, 1992b)), Mexico (D. F., Guadalupe Is. (Clark, 1953), Hidalgo, Mexico, Oaxaca, Puebla, San Luis Potosí), Brazil (Pernambuco), and Argentina (Catamarca, Salta). The species is infrequent and has been collected on soil and soil over rocks (granitic or calcareous soil, lava), exposed in seepage areas, on water edges and along paths, or sheltered around tree bases in open forests, or under scrub, from 80 m to 2800 m. Specimens examined. MEXICO. D. F.: Type; Botanical Graden, Düll A 72, 75 (Hb. Düll), 1979, Jovet-Ast s.n. (PC). HIDALGO: Above La Trinidad, 10 mi. S of Pachuca, Whittemore 1961 (Hb. Whittemore). MEXICO: Lecheria, Pringle 15327 (G). OAXACA: Pan American Road, km 817, 22 km S Juchitlan near Tahuantepec, Düll A 70 (Hb. Düll). PUEBLA: Hwy. 190, 3.2 mi. E of junction Hwy. 160, Whittemore 2279 (Hb. Whittemore). SAN LUIS POTOSÍ: 40 km S San Luis Potosí, Volk 6/0723, 6/0725 (Hb. Volk); 40 km E Sombrereto, Volk 6/0727, 6/0730 (Hb. Volk); Sombrereto, Volk 6/0729 (Hb. Volk). BRAZIL. PERNAMBUCO: Belém de São Francisco, Vital 8184 (SP). ARGENTINA. CATAMARCA: Type of R. dorsiverrucosa; type of R. andina; Cumbres de Suncho, Sleumer 2140 (LIL); Las Rosas, Sleumer 2142 (LIL); Belén, Las Bayas, Portenzuelo del Rio Blanco de las Granadillas, Sleumer 2153 (LIL). SALTA: Cafayate, Vervoost 5054 b (LIL); Cerro Teresita, Vervoost 5052 b (LIL). Riccia mauryana belongs to the species group comprising R. vitalii, R. ridleyi, R. ekmanii and R. weinionis, all with thickened cell walls in dorsal tissue. However, in these species the thickened longitudinal bands never extend over the entire walls whereas in R. mauryana two walls are entirely or nearly entirely thickened, and the other walls show a narrow, longitudinal, hyaline strip. Hässel de Menéndez (1963) treated R. andina and R. dorsiverrucosa as distinct species but did not recognize R. mauryana.

24. Riccia nigrella DC., in DeCandolle & Lamarck, Fl. franç. ed. 3, 6 : 193. 1815. Type. France. Grammont near Montpellier, 1807, Bouchet s.n. (holotype, PC; isotype, G).

Figs. 35, 36, 79: F

Bischler et al. 59 Riccia aggregata Underw., Bot. Gaz. 19: 275. 1894. Type. U. S. A. California: Pasadena, McClatchie 24 (holotype, NY). Thalli forming rosettes, or gregarious, crowded mats; lobes 1-1.2 mm wide, acute or obtuse apically, 3 times forked with lobes divaricate at 40°, the basal dichotomy deep (2.5 mm), the following at 1.5 mm and the last short (0.2-0.6 mm); dorsal side dark blue-green, red-brown basally, on ventral side and on the narrow margins, margins inflexed when dry and lobes showing only the black flanks, appearing as black threads. Median groove deep, narrow. Epidermal cells persistent, hyaline, convex, 1.5 times higher than hypodermal cells. Ventral scales large, imbricate, semi-lunate, reaching or extending beyond lobe margins, dark violet or nearly black and glossy, with entire, pink margins. Cross section of lobes 0.8 mm high, 1-1.5 times as wide as high; dorsal edge acute laterally; flanks ascending, recurved apically and acute; ventral edge convex; dorsal tissue reaching half of lobe height. Monoecious. Archegonial necks long, black. Sporangia hardly bulging, dorsal tissue above the sporangia usually red-brown. Spores 60-80 µm diameter, light brown; wing 5-7 µm wide, faintly crenulate with a few dark, vermiculate ridges; distal face with 8-9(-10) areoles across diameter, incompletely delimited by irregular, thick, sinuose and anastomosed, dark brown ridges, some areoles with partly thin or vanishing ridges, more complete and smaller areoles with tubercles on the ridges near equator; proximal face with complete areoles and large tubercles, or not areolate, with irregular, thick and sinuose, bifurcate ridges, triradiate scar thin, sometimes interrupted. Gametophytic chromosome number n = 8. Distribution and ecology. Riccia nigrella is widely distributed in central and Atlantic Europe, reaching northwards to Great Britain, in the Mediterranean area, Macaronesia, South Africa, China, and Australia. It is known from southern United States (Arizona, California, Georgia, Texas), and has been recorded in the Neotropics from Mexico (Baja California (Howe, 1934), D. F., Guadalupe Is. (Howe, 1934), Oaxaca). The species is infrequent in the Neotropics. It has been collected on soil with lava fragments or in seepage areas, from sea level to 2300 m. Neotropical specimens examined. MEXICO. D. F. : Botanical Garden, 1979, Jovet-Ast s.n. (PC). OAXACA: Road Teotitlan-Cuitlapan, 14 km of Cuitlapan, Düll 2/230, 2/232 (Hb. Düll). Riccia nigrella is easily recognised by its black and glossy lateral flanks, and its narrow, bluishgreen lobes tinged with red-brown basally and on the margins.

25. Riccia plano-biconvexa Steph., Bih. Kongl. Svenska Ventensk.-Akad. Handl. 23 (III, 2): 29. 1897. Type (Jovet-Ast, 1991). Brazil. Rio Grande do Sul: Cachoeira, Lindman 184 (lectotype, G) Figs. 37, 40, 79: G Riccia tenuilimbata Steph., Bih. Kongl. Svenska Ventensk.-Akad. Handl. 23 (III, 2): 30. 1897. Type. Paraguay. Colonia Presidente Gonzales, Lindman 300 (holotype, S).

Bischler et al. 60 Thalli in gregarious patches; lobes 7-12 mm long, 1.5-2.3(-3) mm wide, thin, rounded or slightly emarginate apically; dorsal side light green or greenish with hyaline, margins 3-4 cells wide; flanks and ventral side light green, or brownish, or violeaceous. Median groove distinct apically, flattened and vanishing towards base. Epidermal cells disintegrating. Ventral scales hyaline, usually not extending beyond lobe margins. Cross section of lobes 3-5 times as wide as high; dorsal edge abruptly extending into thin wings 3-4 cells high, ending in a hyaline cell; flanks erect; ventral edge convex; dorsal tissue in 8-10 cell layers, thinner than ventral tissue, all cells thin-walled. Dioecious? Archegonial necks protruding, violet basally, hyaline above. Spores subspherical, 70-90 or 100-130 µm diameter, red-brown, wingless; distal face with 8-11 areoles across diameter, those of central part smaller (7-8 µm diameter) than those of periphery (9-14 µm diameter), with thin, high ridges and thin, 6 µm high tubercles; proximal face similarly ornamented, with somewhat lower ridges and tubercles, triradiate scar indistinct. Distribution and ecology. Riccia plano-biconvexa is a neotropical species recorded from Costa Rica (Jovet-Ast, 1981), the Galapagos Islands (Isabela (Gradstein & Weber, 1982)), Brazil (Alagoas, Bahía, Espíritu Santo, Goiás, Mato Grosso, Pernambuco, Rio Grande do Norte, Rio Grande do Sul), Paraguay, and Argentina (Misiones (Herzog, 1952)). It seems to be quite common in Brazil and has been collected on sandy, granitic or clayey soil, on soil over rocks, exposed on seasonal river banks, on borders of ponds, in pastures, between paving stones in towns, or sheltered by open vegetation, in caatinga or gardens, from sea level to 500 m. Specimens examined. BRAZIL. ALAGOAS: Maccio, Schäfer-Verwimp & Verwimp 8742 (Hb. Schäfer-Verwimp). BAHÍA: Ibotirama, Vital 8002 (SP); Bom Jesus da Lapa, Vital 7992 (SP). ESPÍRITU SANTO: Linhares, Vital 1870, 1871 (SP). GOIÁS: Tocantinopolis, Vital 2989 (SP). MATO GROSSO: Coxipo, Cuiaba, Lindman 358 (G). PERNAMBUCO: Fazenda Nova, Hürlimann H 524 (Hb. Hürlimann); Floresta, Vital 8181 (SP); Saltinho, Porto 1375 (PC). RIO GRANDE DO NORTE: Angicos, Vital 1974 (SP). RIO GRANDE DO SUL: Type; Tenente Portela, Barranco do Uruguai, Vianna 4257 (ICN); São Francisco de Assis, Picada do Padre, Vianna 3066 (ICN). PARAGUAY. Type of R. tenuilimbata. Riccia subplana is related to R. plano-biconvexa but the spores of the former species are pinkviolet or red-violet, with only 4-6 areoles across diameter of the distal face. Hässel de Menéndez (1963) included R. weinionis and R. corcovadensis as synonyms in Riccia plano-biconvexa. However, the former two taxa have two thickened strips in the cells of the dorsal tissue, a character not seen in R. plano-biconvexa.

26. Riccia ridleyi A.Gepp, J. Linn. Soc., Bot. 27: 74. 1890. Type. Brazil. Is. Fernando Noronha: Tangle Rock, 1887, Ridley s.n. (holotype, BM).

Figs. 38, 40, 79: H

Bischler et al. 61 Riccia bistriata A.Evans, Torreya 19: 85, 1919. Type. Peru. Cuzco: Santa Ana, 900 m, Cook & Gilbert 1481 (holotype, US). Thalli in gregarious patches; lobes 4.5 mm long, 1.3-2 mm wide, once forked, the dichotomy reaching half of lobe length, rounded apically; dorsal side greyish-green with narrow, acute, violet margin of large, hyaline or violet cells with thickened walls; flanks pink or pink-violaceous; ventral side pale green, sometimes pink-violaceous. Median groove deep. Epidermal cells disintegrating except in groove, pyriform, upper wall convex. Ventral scales imbricate, rounded apically, reaching or extending beyond lobe margins. Cross section of lobes 2-4 times as wide as high; dorsal edge rounded laterally; flanks oblique, with 1-2 hyaline or pink cells apically; ventral edge convex; dorsal tissue 1/3 of lobe height, in 5-7 cell layers, cell walls with two thickened, longitudinal strips. Dioecious? Spores tetrahedral, 98-120 µm diameter, red-brown or red-violet; wing incomplete, narrow, light brown; distal face with 10-15 incomplete areoles across diameter, distinguished by thick, irregular dark red or nearly black tubercles; proximal face with short ridges and more or less large tubercles of varying size, forming a network of incomplete areoles, triradiate scar weak or indistinct. Distribution and ecology. Riccia ridleyi is rare, known from Venezuela (Barinas (Infante-Sanchez & Heras Pérez, 2002), Falcón), Peru (Cuzco), and Brazil (Fernando Noronha), from sea level to 900 m. Its ecology remains to be investigated. Specimens examined. VENEZUELA. FALCÓN: 60 km S of Coro, 2 km S of Valle de Maria, Wingfield 13909 (Hb. Wingfield); Peninsula of Paraguana, Cerro Santa Ana above Santa Ana, Wingfield 13367 (Hb. Wingfield). PERU. CUZCO: Type of R. bistriata. BRAZIL. FERNANDO NORONHA IS.: Type; Topo do Pico, Vital 8314 (SP); without locality, 1877-1878, Jobert s.n. (PC). For differences with the allied R. weinionis and R. vitalii, see under R. vitalii.

27. Riccia sanguineisporis Jovet-Ast, Cryptog. Bryol. Lichénol. 12: 253. 1991. Type. Brazil. Mato Grosso do Sul: Mun. de Miranda, Vital 2335 (holotype, PC; isotype, SP).

Figs. 39, 40, 79: I

Thalli in gregarious patches; lobes 2-3 times deeply forked, 6-11 mm long, 0.8-1.2 mm wide, obtuse apically; dorsal side greyish green with narrow, pale margins, flanks brownish. Median groove deep from apex to base. Epidermis and the two layers below epidermis nearly hyaline, basal walls of epidermal cells and walls of the 4-5 hypodermal cell layers strongly thickened. Ventral scales hyaline, hardly extending beyond lobe margins. Cross section of lobes 0.8 mm high, as wide as high apically to 1.5(-3) times as wide as high near base; dorsal edge acute laterally; flanks suberect to oblique, often with the external layer of cells with thickened walls; ventral edge convex; cell walls of ventral tissue

Bischler et al. 62 thin. Monoecious? Spores subspherical, 130-145 µm diameter, dark red, wingless, crenulate; distal face without areoles, with large, red-black tubercles at the poles becoming smaller towards equator; proximal face with similar ornamentation, triradiate scar of large, dark red tubercles, not reaching the equator. Distribution and ecology. The species is known only from the type specimen. Specimen examined. BRAZIL. MATO GROSSO DO SUL: Type. The thickened walls of the epidermal and hypodermal cells and the marginal hyaline cell row of the narrow margin of the lobes, indicate an affinity with Riccia sorocarpa. However, in all other features of thallus and spores the two species are quite distinct.

28. Riccia sorocarpa Bisch., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 17(2): 1053. 1835. Type. Germany. Schriesheim near Heidelberg, Bischoff s.n. (syntype, HEID). Figs. 41, 44, 79: K Riccia antarctica Steph., Sp. hepat. 1: 32. 1898. Type. Chile. Without locality, Bertero 1279 (holotype, PC; isotypes, G, STR). Riccia chilensis Steph., Sp. hepat. 1: 33. 1898. Type. Chile. Valparaiso: El Salto, Dusen 141 (holotype, G) Riccia rauhii Buchloh, Nova Hedwigia 3: 510. 1961. Type. Peru. Cuzco: Ausangate, 4600 m, Rauh P 2504 (n.v.). Thalli in rosettes 2 cm in diameter; lobes 3-10 mm long, 0.5-2 mm wide, rounded apically, 2-3 times forked, the basal dichotomy deep, of 2/3-3/4 of lobe length; dorsal side pale green, rarely bluish green, with narrow, hyaline margins; flanks, parts of dorsal side and margins sometimes tinged with violet. Median groove deep at lobe apex, becoming shallower below but reaching lobe base. Epidermis of rounded or pyriform cells, disintegrating except in groove, basal walls and walls of the hypodermal cells strongly thickened. Ventral scales pale, not reaching lobe margins. Cross section of lobes slightly wider than high near apex, 2-3 times as wide as high below; dorsal edge obtuse and ascending laterally; flanks obliquely ascending, with independent row of long and narrow cells ending in a long, rounded cell apically. Monoecious. Antheridial necks hardly protruding, pale. Sporangia large, black, bulging on ventral side. Spores tetrahedral, 70-98 µm diameter, dark red with black-red ornamentation; wing 5-7 µm wide, wider near the pores, irregular, granulose and crenulate on margin; distal face with 8-10(-14) areoles across diameter, with rather thin ridges and large tubercles, the 4 areoles near pole often larger; proximal face covered with granules, or vermiculate, or with short, sinuose ridges, ridges of triradiate scar slender. Gametophytic chromosome number n = 8. Distribution and ecology. Riccia sorocarpa is widely distributed in Europe to 70° N, in the

Bischler et al. 63 Mediterranean area, Macaronesia, the Sahara, South Africa, Asia (SW Asia, Siberia, China), Australia, New Zealand, and Hawaii. It is known from north-western Greenland, from most of the states of the United States, but seems to be infrequent in the Neotropics. It has been recorded from Mexico (Baja California (Schuster, 1992b), D. F., Guadalupe Is. (Howe, 1934), Mexico, Oaxaca), Costa Rica, Peru (Ancash, Cuzco, Junín, La Libertad), Brazil (Pernambuco (Yano & Andrade Lima, 1987)), Chile (Valparaiso), Argentina (Catamarca (Hässel, 1958), Mendoza (Hässel, 1958), Neuquen (Hässel, 1963), Rio Negro (Hässel, 1963)), and Uruguay. The species has been collected in humid to semi-arid climatic areas, on various types of soils (acidic or basic, sandy, clayey or silty), over acidic or calcareous rocks, usually in exposed situations, also on river borders and in gardens, from sea level to 4530 m. Neotropical specimens examined. D. F. : Botanical Garden, Düll A 74 (Hb. Düll), Jovet-Ast s.n. (PC); MEXICO: 8 km E of Rio Frio Pass, Düll 98 (Hb. Düll); Popocatepetl, 15 km above Amecameca, Düll 2/395, 2/396 (Hb. Düll). OAXACA: Pan American Hwy, km 817, 22 km S Juchitlan, near Tehuantepec, Düll 2/610 (Hb. Düll). COSTA RICA. Taras de Cartago, road from Cartago to Turrialba, 1979, Jovet-Ast s.n. (PC). PERU. ANCASH: Quebrada Pachacolo, 1982, Geissler s.n. (G). CUZCO: Type of R. rauhii. JUNÍN: Jauja, La Oroya, Quebrada Shaco near Canchayllo, P. & E. Hegewald 5343 (Hb. Hegewald); Jauja, La Oroya, Laguna Mesapata near Canchayllo, P. & E. Hegewald 5368 (Hb. Hegewald); Jauja, Laguna Huaylacancha, P. & E. Hegewald 5865 (Hb. Hegewald). LA LIBERTAD: Otuzco, Salpo, P. & E. Hegewald 5109 (Hb. Hegewald); Trujillo, Cerro Campana, P. & E. Hegewald 7258 (Hb. Hegewald). CHILE. VALPARAISO: Type of R. chilensis. URUGUAY. Tacuarembo, Herter 78509 (PC). Riccia sorocarpa has a smooth thallus surface with a very narrow, hyaline margin, and epidermal and hypodermal cells with thickened walls. These characteristics distinguish the species from all other neotropical Riccia species. Riccia sorocarpa has a wide geographical range and is able to cope with a variety of habitats, from tropical areas to arctic zones. Its morphological characteristics are stable with exception of pigmentation of the flanks, degree of thickenings of the cell walls of the epidermal and hypodermal cells, spore diameter, and width of the spore wing. Geographical correlation of morphological variation has not been observed. Schuster (1992a) described the xeromorphic subsp. erythrophora Schust. (J. Hattori Bot. Lab. 71: 274. 1992. Type. U. S. A. California: San Diego C°, 1.5 km W of Potrero, Schuster 82-146) from the south-western U. S. A. and Mexico (Baja California: 20 km S of Tecate, Schuster 82-153). It differs from subsp. sorocarpa by the only 1-2 times forked lobes, the less conspicuously thickened walls of the epidermal and hypodermal cells, and the vinaceous or purplish scales. These characteristics hardly deserve taxonomic status.

Bischler et al. 64

29. Riccia squamata Nees, in Martius, Fl. Brasil. 1: 302. 1833. Type. Brazil. Minas Gerais: Serra de Piedade, Martius s.n. (holotype, STR; isotype, PC).

Figs. 42, 44, 79: L

Riccia subnigella Herzog, Feddes Rep. Nov. Spec. Regn. Veg. 55: 6. 1952. Type. Argentina. Misiones: Dep. Iguazu, Iguazu Preserve, Hosseus 362 (holotype, JE). Thalli in gregarious patches; lobes 2-3 mm long, 1.1-1.8(-2.3) mm wide, rounded apically; dorsal side light brown greenish, bordered by the black scales connate with the flanks, margins completely inflexed when dry; flanks dark red-violet; ventral side pale pink. Median groove shallow. Epidermal cells rounded or pyriform. Ventral scales black or red-violet, glossy, imbricate, with smaller marginal cells. Rhizoids all smooth. Cross section of lobes as wide as high apically to twice as wide as high below, and 2-2.5 times as wide as high basally; dorsal edge with a weak median groove; flanks ascending vertically; ventral edge convex; cells of dorsal tissue with thin, longitudinal strips. Monoecious. Sporangia hardly bulging. Spores tetrahedral, 96-125 µm diameter, brick red with darker ornamentation, wingless; distal face with 7-9(-11) areoles across diameter, with thin ridges and large, anastomosed tubercles, the areoles sometimes replaced by folds near equator; proximal face nearly smooth, with few, low, reddish-black tubercles and granules, triradiate scar weak, with narrow ridges. Distribution and ecology. Riccia squamata is an infrequent neotropical species and has been recorded from Mexico (Zacatecas), Southern Dutch Antilles (Curaçao), Brazil (Bahía, Minas Gerais, Paraíba, Pernambuco, Piauí), Chile (Valparaiso (Müller, 1955)), and Argentina (Misiones). The species has been collected on sandy or rocky soil, exposed or partially shaded, on river banks, in caatinga vegetation, or in deciduous forests, at low elevation. Specimens examined. MEXICO. ZACATECAS: Rio Grande, Las Masas, Parry et al. 445 a (CINC). SOUTHERN DUTCH ANTILLES. CURAÇAO: Seroe di Palomba, Christoffelsberg-area, van Slageren & Stoffers 8171 (U). BRAZIL. Without locality, Ule 504 (FI). BAHÍA: Jaguarari, Vital 8098 (SP). MINAS GERAIS: Type. PARAÍBA: São Mamede, Vital 2905 (SP); Pombal, Vital 2910 (SP). PERNAMBUCO: Salgueiras, Vital 1981 (SP). PIAUÍ: Picos, Vital 2930 (SP); Oeiras, Vital 2936 (SP); Elizeu Martins, Vital 8211 (SP). ARGENTINA. MISIONES: Type of Riccia subnigella. Riccia squamata is well characterized by spore shape, red colour and ornamentation. By its spore ornamentation R. squamata cannot be confused with any other neotropical Riccia species.

Bischler et al. 65 30. Riccia subdepilata Jovet-Ast, Cryptog. Bryol. Lichénol. 12: 228. 1991. Type. Brazil. Bahía: Mun. de Jeremoabo, Vital 8146 (holotype, PC; isotype, SP).

Figs. 43: A-K, 44, 79: M

Thalli forming complete or partial rosettes, of 13 mm diameter. Lobes with 3-4 deep dichotomies, 3-9 mm long, 0.7-1.1 mm wide, thin, rounded and slightly emarginate and inflexed apically; dorsal side light brown to pale green. Median groove nearly absent. Epidermal cells disintegrating. Ventral scales hyaline, not reaching lobe margins. Cilia few, scattered on lobe margins, hyaline, stiff, erect or slightly arched, 120-380 µm long, with thickened walls on one side, granulose from apex to 2/3 of length. Cross section of lobes 0.15-0.2 mm high, of 8-10 cell layers; dorsal edge rounded; flanks oblique; ventral edge hardly convex. Monoecious. Spores spherical to tetrahedral, 84-96 µm diameter, red-brown with darker ornamentation; wing granulose, 4-8 µm wide, to 12 µm wide near pores and delimiting a smooth, triangular area, margin slightly thickened and granulose; distal face with 8-11 complete areoles across diameter, each (7-)9-12 µm diameter, ridges with large tubercles, sometimes with strip of small tubercles only near equator; proximal face with less complete areoles but with similar ridges and tubercles, triradiate scar distinct. Distribution and ecology. Riccia subdepilata is known from Brazil only (Bahía, Pernambuco (Porto et al., 1999)). It has been collected on moist soil, in caatinga vegetation at low elevation. Specimens examined. BRAZIL. BAHÍA: Type, and from the same locality, Vital 8147 (SP). Riccia subdepilata is close to R. horrida, with which it shares a similar spore wall ornamentation. The distinctive characters of the two species are outlined under R. horrida.

31. Riccia subplana Steph., in Urban, Symb. antill. 3: 275. 1902. Type. Martinique. Morne Rouge, Pointe Fine, Duss 579 p.p. (holotype, G; isotype, PC).

Figs. 43: L-R, 47, 80: A

Thalli in complete or partial rosettes, 15-25 mm in diameter, or in gregarious patches; lobes 2-3 times forked, 5-15 mm long, 2.5-3.8 mm wide, rounded apically, thin and flattened when dry; dorsal side light green or glaucous, with acute, often slightly ascending margins; flanks and ventral side brown-violaceous. Median groove shallow. Epidermal cells rounded, hyaline, disintegrating except in groove. Ventral scales brown-violeaceous, usually not extending beyond lobe margins. Cross section of lobes 2-6 times as wide as high; dorsal edge with thin, ascending wing laterally; flanks oblique; ventral edge nearly flat; dorsal tissue occupying 1/3 of lobe height, of 4 cell layers, cells 40(-55) µm high, walls with narrow, pitted, longitudinal strip. Monoecious. Spores subspherical, 85-130 µm diameter, violaceous-brown; distal face with 4-6(-8) complete areoles across diameter, with thin, high ridges and thin, high tubercles, some connected by a thin, pink-violet membrane; proximal face with similar ornamentation, triradiate scar indistinct. Gametophytic chromosome number n = 8. Distribution and ecology. Riccia subplanais a neotropical species known from Guatemala, Costa Rica, Virgin Islands (St. John), Windward Islands (Guadeloupe (Duss, 1904), La Désirade, Les

Bischler et al. 66 Saintes, Martinique), Venezuela (Infante-Sanchez & Heras Pérez, 2002), Guyana, French Guiana, Peru (Loreto), and Brazil (Amazonas). It seems infrequent and has been collected on bare soil or rocky, volcanic soil, along streets among paving stones, in gardens, in cultivation, on borders of paths, or shaded under open vegetation, from sea level to 1000 m. Specimens examined. GUATEMALA. Suchitepequez, Naranjo, Hürlimann H 520 (Hb. Hürlimann). COSTA RICA. W of San José, 1979, Jovet-Ast s.n. (PC); near Alajuela, 1979, Jovet-Ast s.n. (PC); Turrialba, 1979, Jovet-Ast s.n. (PC); Atenas, 1979, Melendez s.n. (PC); Liberia, 1979, Jovet-Ast s.n. (PC); near Las Canas, Guanacaste, 1956, Hürlimann s.n. (Hb. Hürlimann). VIRGIN ISLANDS. ST. JOHN: Breutel s.n. (BM). WINDWARD ISLANDS. LA DESIRADE: Pointe des Colibris, 1989, Breuil s.n. (PC). LES SAINTES: Terre de Haut, Pointe Rodrigue, Le Gallo 5 (PC); Terre de Haut, Trace de Morne Morel, 1989, Breuil s.n. (PC). MARTINIQUE: Type; Fort de France, rue Ernest Renan, 1938, Stehlé s.n. (PC); Fort de France, Le Gallo s.n. (PC); Saint Pierre, Le Gallo 864, 886 (PC); Balata, Le Gallo 887 (PC). GUYANA. East Demerara, Timehri, Dakara Creek, Thomsons Farm, Gradstein 5746 (U). FRENCH GUIANA. Cayenne, Place des Palmistes, Aptroot 15087 A (CAY); Cayenne, Garden Châtons, Cremers 5208 (PC); Roura, Le Gallo 2 (PC); Saül, Gradstein 6111, 6191 (U); Saramaca, near Acarouany, Le Gallo 4 (PC); Salut Islands, Island Royal, Cremers 8642 (PC). PERU. LORETO: Maynas, Iquitos, Casas de Orquideas, Hegewald 6424 (Hb. Hegewald); Maynas, quebrada Yanomono, Timme 14287 (KSP). BRAZIL. AMAZONAS: Manaus, Rua Recife, Schäfer-Verwimp & Verwimp 9783 (Hb. SchäferVerwimp). Riccia subplana is allied to R. plano-biconvexa. The two species are best distinguished by the spores, those of R. subplana being red-violet, with fewer areoles across distal face, and protruding ridges and tubercles on the margins as seen from distal face. A thin, pink-violet membrane connects some of the ridges and gives the spores their particular appearance.

32. Riccia taeniaeformis Jovet-Ast, Cryptog. Bryol. Lichénol. 12: 270. 1991. Type. Brazil. Rio Grande do Sul: Tenente Portela, R. F. do Turvo, Bueno 4563 (holotype, PC; isotype, ICN). Figs. 45, 47, 80: B Thalli in crowded mats; lobes bifurcate, 10 mm long, 1-2.2 mm wide, thin, rounded or slightly emarginate apically; dorsal side pale green, sometimes with violet strip along margins. Median groove distinct apically only. Epidermal cells thin-walled, persistent, rectangular. Ventral scales colourless, small, with cells 48-75 x 36-48 µm, or absent. Cross section of lobes 0.7 mm high apically, 0.4-0.5

Bischler et al. 67 mm below; dorsal edge with thin wings laterally; flanks obliquely ascending, ending apically in the thin wings, with acute apical cell; ventral edge convex; dorsal tissue in 4(-5) layers. Dioecious? Sporangia bulging ventrally. Spores tetrahedral, (84-)110-125 µm diameter, red-brown with dark brown ornamentation; wing crenulate, irregular, yellow, 4-7 µm wide; distal face with 5-10 areoles across diameter, with thin ridges and conspicuous tubercles; proximal face with conspicuous tubercles or with tubercles mixed with short, sinuose or straight ridges, triradiate scar conspicuous. Distribution and ecology. Only one specimen of Riccia taeniaeformis has been collected. Ecological data are missing. Specimens examined. BRAZIL. RIO GRANDE DO SUL: Type. Riccia taeniaeformis is apparently related to R. albopunctata with which is shares the same ornamentation on the proximal spore face. However, idioblasts are lacking in R. taeniaeformis and the flanks of the lobes are not tinged with violet. Additionally, the scales are very small or absent in R. taeniaeformis.

33. Riccia trichocarpa M.Howe, Bull. Torrey Bot. Club 25: 184. 1898. Type. U. S. A. California: San Mateo C°, Campbell 138 (holotype, US; isotype, NY).

Figs. 46, 47, 80: C

Riccia canescens Steph., Sp. hepat. 1: 12. 1898. Type. Algeria. Oran, 1852, Balansa s.n. (PC). Riccia cana Durieu ex Trab., Rev. Bryol. Lichénol. 12: 15. 1942, nom. illeg. Type. Algeria. Oran, 1853, without collector (PC). Thalli forming rosettes or gregarious patches; lobes 2-4 times forked, 2-5.5 mm long, 0.7-1.5 mm wide; dorsal side light green, sometimes glaucous; margins, flanks, ventral side and scales dark violet to nearly black; flanks inflexed when dry. Median groove deep apically, wider and shallower below, vanishing towards base. Epidermal cells disintegrating except in groove, spherical. Cilia on flanks and margins, from top to bottom of lobes, numerous especially apically, intercrossing along midline, white, stiff, erect or slightly arched, acute, smooth, 400-1000 µm long, 30-40 µm wide basally, walls with helical thickenings, or flexuose, smooth, thin-walled setae, 500-900 µm long, 40-60 µm wide basally. Ventral scales small. Cross section of lobes as high as wide apically, to 1.5-2.5 times as wide as high below; dorsal edge rounded laterally; flanks arched; ventral edge convex. Monoecious. Antheridial necks protruding (50-100 µm), white or violet. Archegonial necks hardly protruding, usually violet. Sporangia often indicated by a red or dark violaceous spot on dorsal side of lobes, with 1-12 cilia shorter than those of flanks, and 1-2 setae similar to those of flanks. Spores 60-120 µm diameter, dark brown; wing narrow (2 µm), irregular, vanishing partly, granulose; distal face with 814 areoles across diameter, with thick, granulose ridges and conspicuous tubercles; proximal face with similar ornamentation, but with lower ridges, triradiate scar distinct. Gametophytic chromosome number n = 8 or 16.

Bischler et al. 68 Distribution and ecology. Riccia trichocarpa is widely distributed in Europe, the Mediterranean area, Macaronesia, tropical and subtropical Africa, Madagascar, SW Asia and, probably, Australia. It is widespread in central and south-western United States (California, Texas), and in the Neotropics, from Mexico (Baja California (Howe, 1934; Schuster, 1992b) , D., Guadalupe Is. (Sutliffe, 1932; Howe, 1934), S. Luis Potosí (Schuster, 1992b)) and was recorded from Chile (O'Higgins and Valparaiso (Mahu, 1994)). It is rare in the Neotropics where it has been collected on sandy or granitic soil and lava, in exposed sites in semi-arid areas, and in gardens, from sea level to 2300 m. Neotropical specimen examined. D. F.: Mexico City, Botanical Garden, Düll A 73 (Hb. Düll). Riccia trichocarpa is related to R. horrida but lobe morphology, length of cilia, spore diameter and number of areoles across distal face differ among the two species. Schuster (1992b) treated R. trichocarpa as a subspecies of R. ciliata.

34. Riccia viannae Jovet-Ast, Cryptog. Bryol. Lichénol. 12: 261. 1991. Type. Brazil. Rio Grande de Sul: Porto Alegre, Morro de Policia, Vianna 1199 (holotype, PC; isotype, ICN). Figs. 48, 51, 80: D Thalli in partial rosettes, or in gregarious patches; lobes 2-3 times forked, 5-15 mm long, 2-4 mm wide, rounded or obtuse apically; dorsal side light green greyish with hyaline, inflexed margins when dry, dark green when moist; flanks often with light pink spots. Median groove deep apically, vanishing below. Epidermal cells disintegrating except in groove, spherical; below epidermis two layers of hyaline cells, the upper one with thickened walls. Ventral scales hyaline, reaching or extending beyond lobe margins, bordered by cells with thickened walls. Cross section of lobes twice as wide as high apically, 3-4 times as wide as high below; dorsal edge with wings of 1-4 cells laterally; flanks obliquely ascending, then concave; ventral edge convex; cells of dorsal tissue with thin, longitudinal strips. Monoecious. Spores 76-120 µm diameter, light brown or red-brown, wingless; distal face with 4-6 areoles across diameter, granulose inside, the central larger, of 24-30 x 20-25 µm, those of the periphery 15-19 µm diameter, vanishing often towards equator and replaced by tubercles or sinuose, thin and low ridges with weak or without tubercles; proximal face with weak ornamentation of low tubercles or granules, or with numerous, nearly indistinct, small areoles, triradiate scar weak or indistinct. Distribution and ecology. Riccia viannae is known only from south-eastern Brazil (Rio Grande do Sul, Santa Catarina), where is seems not to be uncommon. It has been collected on soil, rocky boulders, borders of rivulets and drainages, in gardens and parks, at low elevation (0-200 m). Specimens examined. BRAZIL. RIO GRANDE DO SUL: Type; Porto Alegre, Morro de Policia, Homrich 1668 (ICN); Porto Alegre, Jardim do Fac. de Direito, Vianna 3999 (ICN); Tristesa, Vianna 9b (ICN); Vila Conceição, Vianna 2797 (ICN); Petropolis, Vianna 10, 339, 340, 2708 (ICN); Parque

Bischler et al. 69 Arroupilha, Vianna 1665 (ICN); Ponta Grossa, Vianna 2649, 3991 (ICN); Agronomia, Vianna 3072 (ICN); Gravatoi, Itacolomi, Vianna 3073 (ICN); Guajiba, Vianna 3071 (ICN); Lajeado, Parque de Souza, Vianna 337 (ICN); Pedro Osorio, Vital 9123 (SP); Sepé, Vital 9209 (SP); Ivoti, Cascata São Miguel, Vianna 630, 1196, 6390 (ICN); Viamão, Parque Saint-Hilaire, Vianna 8, 341 (ICN); Santa Maria, Boca de Monte, Vianna 6391 (ICN); Lavras do Sul, Vianna 3143 (ICN); Torres, Vianna 629 (ICN); São Francisco de Assis, Picada do Padre, Vianna 3065 (ICN); Cachoeira do Sul, Vital 9241 (SP); Uruguaiana, 40 km E of Porto Alegre, 1978, Volk s.n. (Hb.Volk). SANTA CATARINA: São Joaquim, Vital 2018 (SP). Riccia viannae is related to R. lamellosa. However, the spores in the two species are quite distinct, those in R. lamellosa being red-brown with black ridges, in R. viannae brown (when mature) and translucent, with granulose areoles.

35. Riccia violacea M.Howe, in E.Britton, Ann. Missouri Bot. Gard. 2: 51. 1915. Type. Puerto Rico. Mona Is.: between Sardinero and Ubero, Britton et al. 1749b (holotype, NY). Thalli in gregarious patches; lobes 1.5-5 mm long, 0.7-1 mm wide, 1-3 times forked, obtuse or slightly emarginate apically, margins obtuse; dorsal side blue-green, with dark violet margins when moist; flanks inflexed when dry, dark red-violet, with wide, purplish strip inside; ventral edge colourless. Median groove shallow apically, wider and flattened below. Epidermis disintegrating except in groove, cells rounded, mamilliform. Ventral scales short, irregular in shape, 2-3 cell layers in height, bordered by hyaline or violet projecting cells, or by hyaline, papilliform cells. Papillae numerous, on lobe margins and on upper part of flanks, hyaline, sometimes violaceous, smooth or granulose, rounded or obtuse apically, 25-130 µm long, 25-45 µm wide basally, erect or slightly arched, together with nearly acute, hyaline, strongly granulose papillae. Cross section of lobes 1-2 times as wide as high, without wings; dorsal side rounded laterally; flanks ascending, rounded apically; ventral edge convex or nearly flat. Dioecious. Antheridial necks protruding, violet. Spores subspherical, 85-115 µm diameter, black or dark brown, wingless, crenulate on margin; distal face with 12-17 areoles across diameter, with thick, black ridges and large tubercles; proximal face with similar ornamentation but usually without tubercles, triradiate scar weak. Riccia violacea is related to R. atromarginata var. atromarginata but in R. violacea the papillae are shorter, absent on the dorsal side of lobes. The species has been included by Schuster (1992b) as a synonym of R. atromarginata subsp. atromarginata. This synonymy is not accepted here (see under R. atromarginata).

KEY TO THE VARIETIES OF RICCIA VIOLACEA 1. Papillae granulose ...................................................................................... .....35a. var. violacea

Bischler et al. 70 1. Papillae smooth................................................................................................ ...35b. var. laevis

35a. Riccia violacea var. violacea

Figs. 49, 51

Papillae on lobe margins and upper part of flanks, hyaline or violet, granulose, obtuse or rounded apically. Distribution and ecology. Riccia violacea var. violacea is probably rare, known from Mexico (Baja California (Crum & Steere, 1959), Ceralvo Is. (Crum & Steere, 1959)), Cuba and Puerto Rico. The variety has been collected on exposed, calcareous soil, silt, or limestone cliffs, on borders of periodically dried-up rivers, at low elevations. Specimens examined. CUBA. Santiago de Cuba, Aguadores, Ekman 8701 (S); Habana, near Morro, Ekman 1155 (S); Oriente, near Imias, Rio Tacre, Dietrich Je-H 1554, 1556 (JE); Oriente, Baracoa, Imias, La Chivera, Meyer 2900 (JE). PUERTO RICO. MONA ISLAND: Type.

35b. Riccia violacea var. laevis Jovet-Ast, Cryptog. Bryol. Lichénol. 10: 100, 1989. Type. Leeward Is. Saint-Barthélémy: Between Anse Gouverneur and Pointe des Nègres, Le Gallo 626 (holotype, PC).

Figs. 50, 51, 80: F

Papillae on upper part of flanks and on lobe margins, hyaline, smooth, rounded or obtuse apically. Distribution and ecology. This variety is known from Mexico (Coahuila, Hidalgo, Veracruz), Bahamas (Pagán, 1939), Leeward Islands (Saint Barthélémy), Virgin Islands (Anguilla), Southern Dutch Antilles (Curaçao, Bonaire), and Peru (Amazonas). It seems to be less infrequent than var. violacea and has been collected on calcareous rocks and soil, on limestone, in steppes, under scrub vegetation and in evergreen or deciduous, dry forests, from sea level to 2200 m. Specimens examined. MEXICO. COAHUILA: 30 km NW of El Saltillo, Whittemore 2335 (Hb. Whittemore); 59.5 mi. N of Sabinas Hidalgo, Whittemore 2508 (Hb. Whittemore). HIDALGO: Andalgala, Barranco de los Venados, near Pachuca, Düll A 122 (PC). VERACRUZ: 15 km NW Perote, near El Limon, Düll A 77 (PC). LEEWARD ISLANDS. SAINT BARTHELEMY: Type; Between Anse Gouverneur and Pointe des Nègres, Le Gallo 510 (PC); Morne de l'Anse de Toiny, Le Gallo 1282, 1283 (PC). VIRGIN ISLANDS. ANGUILLA: Eastern, Le Gallo 1278, 1298 (PC). SOUHERN DUTCH ANTILLAS. BONAIRE: between Slagbaai and Labra, Stoffers & van Slageren 8346 (U). CURAÇAO: San Hironimo, Stoffers & van Slageren 8022 (U); Tafelberg of San Hironimo, Stoffers & van Slageren 8121, 8122 (U); Savonette side of Christoffelberg, Stoffers & van

Bischler et al. 71 Slageren 8171 (PC). PERU. AMAZONAS: Bagua Chica, near bridge to Bagua Grande, P. & E. Hegewald 7148 (Hb. Hegewald).

36. Riccia vitalii Jovet-Ast, Mem. New York Bot. Gard. 46: 283. 1987. Type. Brazil. Bahía: Mun. de Milagres, Vital 5941 (holotype, PC; isotype, SP).

Figs. 52, 53, 80: E

Thalli in gregarious patches, with fish smell; lobes deeply divided, 7 mm long, 1-2.3 mm wide, rounded apically; dorsal side pale or dark green; flanks and ventral side violet or pink-violet. Median groove shallow apically, vanishing below. Epidermis hyaline, disintegrating except in groove, cells convex. Ventral scales imbricate, violet or pink with hyaline marginal cells, extending beyond lobe margins. Cross section of lobes acute laterally with short, thin wings, of 1-3 hyaline or violaceous cells; flanks ascending, erect or oblique; ventral edge convex; dorsal tissue 1/3-1/2 of lobe height, cell layer below epidermis with two thickened longitudinal strips on cell walls. Dioecious (sometimes monoecious?). Spores subspherical or subtetrahedral, 92-152 µm diameter, dark red, or dark redbrown, or blackish, wingless; distal face with 6-9 more or less complete areoles across diameter, larger near pole than near equator, sometimes complete only near pole, with thick, irregular, rugulose ridges and irregular tubercles, at periphery with irregular granules and tubercles oriented towards equator; proximal face with more or less anastomosed granules, triradiate scar weak or indistinct. Distribution and ecology. Riccia vitalii is known from Costa Rica, Brazil (Alagoas, Amazonas, Bahía, Ceará, Espíritu Santo, Goiás, Maranhão, Mato Grosso do Sul, Paraíba, Pernambuco, Piauí, Rio Grande do Norte, Sergipe), and Paraguay. The species seems quite common in Brazil and has been collected on acidic soil over granitic rocks or sandstone, exposed in dry grasslands, on water edges, also along roadsides and in towns, or partially shaded in open caatinga vegetation, from sea level to 400 m. Specimens examined. COSTA RICA. Murciliago Bay, Howell 240 (NY). BRAZIL. ALAGÓAS: Arapiraca, Vital 2881, 2882 (SP). AMAZONAS: Manaos, Praça, near the Opera, Schäfer-Verwimp 7218 (Hb. Schäfer-Verwimp). BAHÍA: Type; Santa Ines, Vital 6520 (SP); Uaua, Vital 8117 (SP); Gaviao, Vital 1985 (SP); Riachao de Jaquine, Vital 1986 (SP). CEARÀ: Chorozinho, Vital 1976 (SP); Alto Santo, Vital 1978 (SP); Icos, Vital 1979 (SP); Campos Salas, Vital 2922 (SP). ESPÍRITU SANTO: Imbiracu, Vital 1929 (SP). GOIÁS: Santa Teresa de Goiás, Vital 3030 (SP); Tacantinopolis, Vital 2989 p.p .(SP). MARANHÃO: Montes Altos, Vital 2974 (SP). MATO GROSSO DO SUL: Aquidauana, Serra de Santa Barbara, Vital 8611 (SP); 2 km SE of Ciudade de Corumba, Vital 8603 (SP); Corumba, Pantanal, Fazenda Santa Clara, Schäfer-Verwimp 8456 (Hb. Schäfer-Verwimp). PARAÍBA: São Mamede, Vital 2904 (SP); Pombal, Vital 2909 (SP); Cajageiras, Vital 2912 (SP); Campina Grande, Vital 2895 (SP); Soledade, Vital 2899 (SP). PERNAMBUCO: Bom

Bischler et al. 72 Conselho, Vital 2883 (SP); Serra do Timoteo, Panelas, Vital 2890 (SP); Agrestina, Vital 2891 (SP); Salguairos, Vital 1981 (SP); Belém do São Francisco, Vital 8185 (SP); Paramirim, Vital 8193 (SP); Fazenda Nova, Hürlimann H 522, 524 , 525 (Hb. Hürlimann); between Recife and Caruaru, Hürlimann H 526 (Hb. Hürlimann). PIAUÍ: Fronteira, Vital 2925 (SP); São Juliao, Vital 2928 (SP); Oeiras, Vital 2935 (SP). RIO GRANDE DO NORTE: Lelmo Marinho, Vital 1973 (SP); Equador, Vital 2903 (SP). SERGIPE: São Miguel do Aleixo, Vital 2879 (SP). PARAGUAY. Cerro de Yagouaroux, Balansa 3724 (G). Riccia vitalii resembles R. weinionis and R. ridleyi by the cells of the dorsal thallus tissue having two longitudinal wall thickenings. The three species are distinguished by spore shape and ornamentation: in R. weinionis and R. ridleyi the spores are tetrahedral, less than 125 µm in diameter, and the proximal face is areolate; in R. vitalii they are subspherical or subtetrahedral, 92-150 µm in diameter, and the proximal face is granulose. Riccia weinionis differs from R. ridleyi by the fewer areoles on the spores, both on the distal face (7-9 across diameter; in R. ridleyi 10-12) and on the proximal face (15 areoles per facet; 30-50 in R. ridleyi).

37. Riccia weinionis Steph., Sp. hepat.1: 18. 1898. Type. Brazil. Rio de Janeiro: Weinio 111 (holotype, G).

Figs. 54, 55, 80: G

Riccia corcovadensis Steph., Sp. hepat. 1: 29. 1898. Type. Brazil. Rio de Janeiro: Corcovado, Riedel & Guillemin 289 (syntype, PC). Thalli in gregarious patches; lobes 5-6 times forked, 10-25 mm long, 2-3 mm wide, rounded apically; dorsal side greyish green or whitish when dry, green or blue-green when moist, margins narrow, light green or whitish, acute; flanks tinged with pink, or violet, or black, inflexed when dry; ventral side light green. Median groove deep apically, vanishing progressively towards base. Epidermal cells disintegrating except in groove, rounded. Ventral scales distant, not reaching lobe margins, pink-violet or violet, cells 60-85 x 25-36 µm, margins with two rows of long narrow cells. Cross section of lobes 3-5 times as wide as high; dorsal edge acute laterally with short wings, ending in 2-3 hyaline cells and in a terminal, hyaline, long, acute cell with thickened walls; flanks obliquely ascending and slightly arched; ventral edge convex; dorsal tissue 1/3-1/2 of lobe height, in 4-5 layers, cell walls with two longitudinal, partly anastomosed, thickened strips; cells of ventral tissue thinwalled, sometimes pitted. Monoecious. Antheridial necks hyaline, protruding, 80-170 µm long. Sporangia numerous, large, bulging dorsally. Spores tetrahedral, 90-145 µm diameter, red-brown to black-red, wingless but with lighter, granulose border; distal face with 7-9 areoles across diameter, with blackish ridges and large, often coalescent tubercles; proximal face with few complete areoles with thin ridges and tubercles, or with incomplete, irregular areoles with interrupted ridges, triradiate scar weak or nearly indistinct. Gametophytic chromosome number n = 8.

Bischler et al. 73 Distribution and ecology. Riccia weinionis is known from Mexico (D. F., Mexico, Oaxaca), Costa Rica, Panama, Leeward Islands (Saint-Barthélémy), Windward Islands (La Désirade, Les Saintes, Martinique, Guadeloupe (Jovet-Ast, 1959)), Southern Dutch Antilles (Bonaire, Curaçao), Colombia (Magdalena), and Brazil (Bahía, Cearà, Espíritu Santo, Goiás, Mato Grosso, Minas Gerais, Paraíba, Paraná, Rio de Janeiro, São Paulo, Sergipe). The species seems to be quite common; it has been collected on soil or rocky soil, or on thin layer of soil over rocks (lava, granite, limestone, sandstone, stone walls), exposed on boulders, along streams, and in gardens, or shaded beneath shrubs or around tree bases in deciduous or evergreen woodlands, from sea level to 2300 m. Specimens examined. MEXICO. D. F.: Mexio, Botanical Garden, Düll A 73 (Hb. Düll). MEXICO: Río La Puerta, above Quimixto, Whittemore 1683 (Hb. Whittemore); Hwy. 131 at km 35, 1.5 mi. S of Calipan, Whittemore 2268 (Hb. Whittemore); Hwy. 190, 3.2 mi. E of junction Hwy. 160, Whittemore 2278 (Hb. Whittemore). OAXACA: road Teotitlan-Cuitlapan, 14 km before Cuitlapan, Düll 2/229, 2/230 (Hb. Düll). COSTA RICA. Guanacaste, La Pacifica, near Las Canas, Hürlimann H 521 (Hb. Hürlimann); Santa Rosa, 1979, Jovet-Ast s.n. (PC). PANAMA. Isla de Barro Colorado, Chung & Aranda Vergara 205 (PMA). LEEWARD ISLANDS. ST. BARTHELEMY: Morne de Gris-Gris, Petite Saline, Le Gallo 895, 896, 897, 899, 901 (PC); Gustavia, near Le Morne, Le Gallo 1274, 1279, 1280, 1281 (PC). WINDWARD ISLANDS. LA DESIRADE: Pointe des Colibris, Le Gallo 1285, 1287, 1288 (PC). LES SAINTES: Ilet de Cabrit, De Foucault 265 (PC); Pointe Rodrigue, Terre de Haut, Le Gallo 5 (PC); Terre de Haut, Morne Morel, Sastre 8447, 8448 (PC); Terre de Haut, trace Morne Morel, 1989, Breuil s.n. (PC). MARTINIQUE: Rivière Salée, Stehlé 4195 (PC). SOUTHERN DUTCH ANTILLES. BONAIRE: road Slagbaai to Labra, van Slageren & Stoffers 8338, 8339, 8340, 8341, 8342, 8343, 8345, 8346 (U); slope of the Brandaris, van Slageren & Stoffers 8365 (U). CURAÇAO: Christoffelsberg area, van Slageren & Stoffers 8121, 8124, 8171 (U). COLOMBIA. MAGDALENA: without locality, Winkler 823 (TUB); Santa Marta, Cerro San Fernando, Schnetter C 435a (TUB). BRAZIL. BAHÍA: without locality, Salzmann 1830 (PC); Guaratinga, Vital 2858 (SP); Serra Preta, along road BA-052, Vital 5999 (SP); 3 km S of Irecé, Vital 6071 (SP); 9 km SW of Maracas, Vital 8728 (SP). CEARÀ: Alto Santo, Vital 1977 (SP). ESPÍRITU SANTO: Linhares, Vital 1872 (SP); Espiritu Santo, Schäfer-Verwimp 8834 (Hb. Schäfer-Verwimp). GOIÁS: Santa Teresa de Goiás, Vital 3031 (SP). MATO GROSSO: Miranda, Vital 2311, 2329, 2357, 2375 (SP). MINAS GERAIS: Nanuque, Vital 1948 (SP); Medina, Vital 1994 (SP); Lagoa Santa, Grutas da Lapinha, Vital 5502 (SP); near Medina on BR-116, km 898, Vital 5910 (SP). PARAÍBA: 2 km W of Esperança, Vital 5447 (SP). PARANÁ: along the road BR-060, km 154, Vital 6339 (SP); without locality, Schiffner s.n. (FH). RIO

Bischler et al. 74 DE JANEIRO: Type; type

of Riccia corcovadensis; Sapucaia, Vital 1995 (SP). SÃO PAULO: Vista

Alegre do Alto, 5 km SE de Vista Alegre, Vital 10391(SP). SERGIPE: Ribeiropolis, Vital 2875 (SP); São Miguel do Aleixo, Vital 2878 (SP). Riccia weinionis belongs to a group of species characterized by cells of the dorsal tissue with longitudinal thickenings (R. ridleyi, R. vitalii, R. weinionis). The species of this group are readily distinguished by spore wall ornamentation (see under R. vitalii). Moreover, in R. vitalii the thallus lobes are narrower (1-2.3 mm) and the pink or violet scales extend beyond lobe margins; in R. weinionis the thallus lobes are wider (2-3 mm), with pale margins, and the scales are shorter and do not pass beyond the lobe margins.

2b. RICCIA subgen. LEPTORICCIA R.M.Schust., Phytologia 56: 72. 1984. Type. Riccia membranacea Gottsche & Lindenb. The subgenus is monospecific. 38. Riccia membranacea Gottsche & Lindenb., in Gottsche, Lindenb. & Nees, Syn. hepat. 608. 1846. Ricciella membranacea (Gottsche & Lindenb.) A.Evans, Rhodora 12: 196. 1910. Type. Mexico. Without locality, Liebmann s.n. (n.v.).

Figs. 56, 57, 80: K

Riccia lanigera Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 570. 1885. Type. Brazil. Pará: river Ramos, tributary of river Amazonas, Spruce s.n. (syntype, BM). Riccia amazonica Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 571. 1885. Type. Brazil. Pará: near Obidos, Paraná-mirí dos Ramos, Spruce s.n. (syntype, BM). Riccia echinatispora Schiffn., Oesterr. Bot. Z. 58: 463. 1908. Type. Brazil. São Paulo: Santos, garden of Julio Conceição, Schiffner 854 (holotype, W). Thalli in complete or partial rosettes, 5-12 mm diameter, deeply divided into 4-6 lobes; lobes 3-5 mm long, 2-4 mm wide, rounded or truncate, often emarginate apically, widened to the tips; dorsal and ventral sides light or yellowish green, thin, membranaceous, shiny. Median groove distinct apically only. Pores bounded by 4 cells, 50-80 µm diameter, or by 6 cells towards base of lobes, 100250 µm diameter. Ventral scales absent. Cross section of lobes 100-150 µm high, with thin wings; ventral tissue in 2-3 layers. Monoecious. Archegonia scattered near median groove. Sporangia bulging ventrally. Spores light brown to red-brown, spherical, 40-65 µm diameter, wingless, spiny, with rounded or truncate spines. Gametophytic chromosome number n=8. Distribution and ecology. Riccia membranacea has been recorded from tropical Africa (Sierra Leone, Ghana, Nigeria, Cameroon, Congo, Tanzania, Angola), and U. S. A. (Arkansas, Connecticut, Delaware, Florida, Illinois, Indiana, Kentucky, Louisiana, New Jersey, Missouri, New Mexico, North Carolina, Ohio, Oklahoma, Tennessee, Texas, West Virginia). In the Neotropics, the species seems to

Bischler et al. 75 be quite common and has been recorded from Mexico (Veracruz), Costa Rica, Puerto Rico (Howe, 1923), Venezuela (Infante-Sanchez & Heras Perez 2002), Ecuador (Pichincha), Peru (Huancabamba, La Libertad, Loreto, San Martin), Brazil (Acre (Vital & Visnadi, 1994), Amazonas (Hell, 1969), Espíritu Santo, Mato Grosso, Pará, Pernambuco (Yano et al., 1987), Rio Grande do Sul, São Paulo), Argentina (Formosa (Herzog, 1952)). It colonizes humid, acidic soil, on river margins, borders of lakes and marshes, or cultivated areas (gardens), from lowland to 2700 m. Neotropical specimens examined. MEXICO. VERACRUZ: near Pepe’s Bar, Esquilon, above Naolinca, Long et al. 29654 (E); river gorge below Teocelo, Long 29694 (E). COSTA RICA. San José, Barrio Mexico, 1979, Jovet-Ast s.n. (PC); Paso Ancho, near San José, 1979, Melendez-Howell s.n. (PC); San Pedro de Montes de Oca, 1979, Melendez-Howell s.n. (PC). ECUADOR. PICHINCHA: Road Quito to Sto. Domingo de los Colorados, Gradstein et al. 6987 (U). PERU. HUANCABAMBA: Piura, W Porculla pass, P. & E. Hegewald 7160 (Hb. Hegewald). LA LIBERTAD: Otuzco, Huancamarca, Quebrada Hornillo, P. & E. Hegewald 5148 (Hb. Hegewald). LORETO: Prov. Maynas, Yanayaco near Iquitos, P. & E. Hegewald 6410, 6411 (Hb. Hegewald). SAN MARTIN: Tarapoto, Ahashiyacu falls, 15 km from Tarapoto, road to Yurimaguas, Gradstein 1828 (U); Tarapoto, Rio Huallaga, Spruce s.n. (G). BRAZIL. ESPIRITU SANTO: Rio Doce, close to Linhares Bridge, Vital 1865 (SP). MATO GROSSO: Mun. Miranda, 5 km E from Fazenda Bodoquena, sect. Guaicurus, Vital 2327 (SP); Corumba, Fazenda Sta Clara, Schäfer-Verwimp 8454 (Hb. Schäfer-Verwimp); near Rio Ramos, Ule 251 (G). PARÁ: syntype of Riccia amazonica; type of Riccia lanigera. RIO GRANDE DO SUL: Cambara do Sul, Itaimbrezinho, Bueno 6388 (ICN); São Luiz Gonçaga, Rio Piratini, 1982, Bueno s.n. (ICN). SÃO PAULO: Type of Riccia echinatispora; Barra do Turvo, Vital 1864 (SP); Mun. Conchas, Rio Conchas, Vital 2471 (SP); Rio Pardo, Sanharao Farm, Barra do Turvo, Vital 1844 (SP); Salto Grande de Paranapanema, Schiffner 2253 (FH). The very thin thallus of Riccia membranacea (2-3 cell layers thick) and the spiny spores distinguish the species from all other neotropical Riccia species.

2c. RICCIA subgen. THALLOCARPUS (Lindb.) Jovet-Ast, Rev. Bryol. Lichénol. 41: 449 . 1976. Thallocarpus Lindb., Not. Sällsk. Fauna Fl. Fenn. Förh. 13: 377. 1874. Cryptocarpus Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 231. 1870 (nom. illeg.). Angiocarpus Trevis., Mem. Reale Ist. Lombardo Sci. Ser. 3, Cl. Sci. Mat. 4: 444. 1877 (nom. illeg.). Type. Riccia curtisii (James ex Austin) Austin. Often heterothallic, male thalli smaller than female. Epidermal pores bounded by 4-6 cells. Dorsal

Bischler et al. 76 tissue with wide air chambers. Ventral scales absent. Spores remaining in tetrads, with large tubercles on distal face. The subgenus is monospecific in the Neotropics and occurs disjunct in Costa Rica and southern South America.

39. Riccia curtisii (James ex Austin) Austin, Bull. Torrey Bot. Club 6: 305. 1879. Cryptocarpus curtisii James ex Austin, Proc. Acad. Nat. Sci. Philadelphia 21: 231. 1870. Thallocarpus curtisii (James ex Austin) Austin, Bull. Torrey Bot. Club 6: 21. 1875. Angiocarpus curtisii (James ex Austin) Trevis., Mem. Reale Ist. Lombardo Sci. Ser. 3, Cl. Sci. Mat. 4: 444. 1877. Type (Schuster, 1992b). U. S. A. Carolina: without locality, 1853, Curtis s.n. (lectotype, BM). Figs 58, 59, 80 H-I Ricciella subsimilis Steph., Bih. Kongl. Svensk. Ventensk.-Akad. Handl. 23 (III, 2): 31. 1897. Riccia subsimilis (Steph.) Steph., Sp. hepat. 1: 42. 1898. Type. Paraguay. Colonia Presidente Gonzales, Lindman 307 (holotype, G; isotype, M). Riccia spegazziniana C.Massal., Atti Accad. Sci. Med. Nat. Ferrara 80: 15. 1906. Type. Argentina. Buenos Aires: La Plata, in the garden of Spegazzini, 1905, Spegazzini s.n. (LPS). Riccia synspora Schiffn., Oesterr. Bot. Z. 58: 462. 1908. Thallocarpus synsporus (Schiffn.) Schiffn., Hedwigia 76: 149. 1936. Syntypes. Brazil. Rio Grande do Sul: Navegantes, Reineck & Czermak 19 (PC); Porto Alegre, Rua dos Voluntarios de Patria, Reineck & Czermak 87 (PC). Female thalli in complete or partial rosettes, to 20 mm diameter, male thalli in rosettes, to 6 mm diameter; lobes 1-3 times forked, the basal dichotomy deep, rounded and hardly emarginate apically; dorsal side yellow-green. Median groove hardly distinct. Pores bounded by 4-6 cells, disintegrating in older parts. Ventral scales absent. Thallus cross section with dorsal tissue 2/3 of lobe height, with 1-2 cell layers bordering 1-2 layers of air chambers, ventral tissue in 2-5 layers; dorsal edge convex, with horizontally spreading, thin wings; flanks obliquely ascending; ventral edge strongly convex. Dioecious. Antheridia in 2-4 rows along median groove, necks red-brown or violaceous. Archegonia scattered along median groove. Sporangia mainly ventrally bulging. Spore tetrads 120-200 per sporangium, 86-112(-148) µm diameter, red-brown to dark brown, wingless, with rounded or truncate tubercles. Gametophytic chromosome number n = 8. Distribution and ecology. Riccia curtisii is known from central and southern Africa, and northern India. It is widely distributed in southern United States (Arkansas, Connecticut, Florida, Louisiana, Mississippi, North Carolina, South Carolina, Texas). In the Neotropics it seems infrequent. It has been recorded from Costa Rica, Brazil (Espíritu Santo, Rio Grande do Sul, São Paulo), Paraguay, Argentina (Buenos Aires, Formosa, Misiones) and Uruguay. It grows on humid, exposed or shaded, sandy or clayey, acidic soil along paths and roads, on river borders, and in cultivated areas (gardens,

Bischler et al. 77 parks, cultivation), from sea level to 1400 m. Selected specimens examined. COSTA RICA. Granadilla, N of Mts. Oca, 1979, MelendezHowell s.n. (PC). BRAZIL. ESPÍRITU SANTO: Linhares, Vital 1874, 1875 (SP). RIO GRANDE DO SUL: Syntypes of Riccia synspora; Porto Alegre, Ponta Grossa, Vianna 1495, 4578 (ICN); Independencia, Vianna 3997 (ICN); Viamão, Parque Saint-Hilaire, Vianna 4178 (ICN); Iguazú, Vianna 256 (ICN); Cachoeira do Sul, Vital 9235 (SP). SÃO PAULO: Salto Grande of Paranaparema, Schiffner 2253 (FH). PARAGUAY. Type of Riccia subsimilis. ARGENTINA. BUENOS AIRES: Type of Riccia spegazziniana. FORMOSA: Pirané, border of Rio Pilaga, Hosseus 52, 53 (CORD). MISIONES: Foz de Iguazú, Schäfer-Verwimp & Verwimp 14872 (Hb. Schäfer-Verwimp). URUGUAY. Dep. Colonia, Estanzuela, Herter 8, 3592 (PC); Montevideo, 25 m, Miguelete, Herter 1471 (PC). The spores, remaining in tetrads, separate Riccia curtisii from all other neotropial Riccia species.

2d. RICCIA subgen. RICCIELLA (A.Braun) Reichenb., Deutsch. Bot. Herbarienb. 23, 213. 1841. Ricciella A.Braun, Flora 4: 756. 1821. Type. Riccia fluitans L. (Ricciella fluitans (L.) A.Braun). Thallus more or less deeply divided into wide or narrow lobes, rosette-forming or not, spongy or not spongy, with wide air chambers. Pores bounded by 4-6 cells. Ventral scales often absent, sometimes large. Sporangia immersed or ventrally bulging. Spores not remaining in tetrads. Riccia subgen. Ricciella is distributed over all continents and over the entire Neotropics, where it is represented by 14 species, in two sections.

KEY TO THE NEOTROPICAL SECTIONS OF SUBGEN. RICCIELLA 1. Thalli spongy or not spongy, deeply divided into long, narrow or wide lobes. Sporangia bulging ventrally. .............................................................................................................................sect. Ricciella 1. Thalli spongy, shallowly divided into wide lobes. Sporangia immersed, hardly bulging ..................... ......................................................................................................................................... sect. Spongodes

4a. Subgen. RICCIELLA sect. RICCIELLA Thallus not spongy, deeply divided into long, narrow or wide lobes, with wide air chambers.

Bischler et al. 78 Sporangia bulging ventrally. Sect. Ricciella has a wide distribution over all continents and has been recorded from most of the neotropical regions, but is rare in the tropical Andes.

KEY TO THE NEOTROPICAL SPECIES OF SECT. RICCIELLA 1. Distal face of spores with worm-like, thin ridges radiating from the distal pole, not areolate ........... ............................................................................................................................................. 44. R. frostii 1. Distal face of spores with more or less complete areoles .................................... .............................2 2. Lobes 1.5-4 mm wide. Spores with 8-20 areoles across diameter of distal face .........................3 3. Ventral scales large, reaching lobe margin. Epidermal pores s very mall, without distinct bounding cells, . Spores wingless .................................................................... 45. R. geissleriana 3. Ventral scales not reaching lobe margin, or absent. Epidermal pores large or with distinct bounding cells, . Spores winged ..................................................................................................4 4. Ventral scales absent. Spore wing 2-3 µm wide. Dioecious, male thalli smaller than female ..................................................................................................... 49. R. paraguayensis 4. Ventral scales short. Spore wing complete, 5-12 µm wide, or irregular and often discontinuous, 2-5 µm wide. Monoecious..............................................................................5 5. Distal face of spores with complete areoles...................................................................6 6. Spores 84-96 µm in diam., areoles of proximal face complete. Lobe margin thick, not wavy........................................................................................... 41. R. chiapasensis 6. Spores 72-86 µm diam., areoles of proximal face often incomplete. Lobe margin thin, wavy............................................................................................... 43. R. dussiana 5. Distal face of spores with ill-delimited areoles .............................. 46. R. hegewaldiana 2. Lobes 0.3-1.5 mm wide. Spores with 3-8 areoles across diameter of distal face .........................7 7. Ventral scales present. Spore wing complete, 3.5-7 µm wide; at least some areoles with a central branched ridge..................................................................................................................8 8. Ventral scales not reaching lobe margin. Lobes 0.7-1(-1.5) mm wide. ............................... .....................................................................................................................47. R. jovet-astiae 8. Ventral scales conspicuous, the apical sometimes extending beyond lobe margin. Lobes 0.3-0.5 mm wide .......................................................................................... 51. R. stenophylla 7. Ventral scales absent, or present apically only, inconspicuous. Spore wing complete or

Bischler et al. 79 discontinuous or absent; areoles of distal face without or with unbranched central ornamentation ..............................................................................................................................9 9. Distal face of spores with 3-5 partly complete, partly incomplete areoles across diam., areoles of proximal face irregular in shape ..................................................... 40. R. bahiensis 9. Distal face of spores with 5-8 complete areoles across diam.; areoles of proximal face regular in shape .....................................................................................................................10 10. Spores 80-92 µm diam.; wing 5-7 µm wide, complete.......................... 48. R. limicola 10. Spores 55-76(-84) µm diam., wing discontinuous, vestigial, or absent .....................11 11. Dorsal edge of lobes projecting above the flanks as wing with unistratose margin, as seen in cross section. Proximal face of spores with areoles similar in size to those of distal face ............................................................................................42. R. crassifrons 11. Dorsal edge of lobes rounded, as seen in cross section. Proximal face of spores with areoles smaller than those of distal face .................................. 50. R. paranaensis

40. Riccia bahiensis Steph., Sp. hepat. 1: 50. 1898. Type. Brazil. Bahía: S of Joazeiro, Martius s.n. (holotype, G).

Figs. 60: A-F, 62, 80: L

Thalli in gregarious patches; lobes bifid, linear, 4-6 mm long, 1-1.5 mm wide, rounded and slightly emarginate apically; dorsal side light brown. Median groove wide (fide Stephani, 1898). Pores 28-30 µm diameter, bounded by 5 cells. Ventral scales absent. Cross section of lobes 0.7-0.8 mm high, usually twice as wide as high; dorsal edge rounded laterally; flanks ascending; ventral side slightly convex; air chambers in 2-3 layers; ventral tissue in 5 layers. Monoecious (fide Stephani, 1898). Archegonia deeply embedded in ventral tissue, separated from ventral epidermis by a single layer of cells. Sporangia ventrally bulging. Spores tetrahedral, 70-84 µm diameter, red-brown with darker ornamentation; wing 4.8-5.5 µm wide, complete or incomplete, granulose; distal face with 3-5 complete or incomplete areoles, with thick and more or less sinuose ridges, in some with a central ornamentation of 1-3 thickenings; proximal face with irregular areoles, more numerous and smaller than those of distal face, with thinner ridges, triradiate scar well marked. Distribution and ecology. Known only from the type specimen. Collected on sandy, acidic soil. Specimen examined. BRAZIL. BAHÍA: Type. Riccia bahiensis has often been considered a synonym of R. cavernosa (e.g., Schuster, 1992b). However, in R. bahiensis the flanks of the lobes are nearly erect and the spores are smaller, with thicker ridges delimiting complete areoles on distal face and with more numerous areoles on proximal face.

Bischler et al. 80

41. Riccia chiapasensis Jovet-Ast, Cryptog. Bryol. Lichénol. 14: 235. 1993. Type. Mexico. Chiapas: Tapachuela, 1960, Riba s.n. (holotype, MEXU; isotype, PC).

Figs. 60: G-N, 62, 80: M

Thalli in gregarious patches; lobes 2-3 times forked, 7-8 mm long, 1.5-2.5 mm wide, truncate and slightly emarginate apically; dorsal side light green when dry. Median groove shallow and wide, hardly distinct. Pores bounded by 4-5 thin-walled cells. Ventral scales hyaline, small, not reaching lobe margins. Cross section of lobes 0.8-1 mm high, usually 2.5-3 times as wide as high; dorsal side obtuse laterally, without wings; flanks obliquely ascending; ventral side slightly convex; air chambers in one layer. Monoecious. Sporangia hardly bulging dorsally. Spores 84-96 µm diameter, brown; wing irregular, 2-5 µm wide, undulate, crenulate, slightly thickened on margin, granulose; distal face with 9-12(-14) areoles of 7 µm diameter across diameter, with thin, granulose, red-brown ridges and tubercles; proximal face with facets of 60-70 areoles, smaller than those of distal face, 1.5-2 µm diameter, with a tubercle in each corner, triradiate scar weak. Distribution and ecology. Riccia chiapasensis has been collected only once, on acidic, siliceous soil. Specimen examined. MEXICO CHIAPAS: Type. Riccia chiapasensis is distinguished from the other neotropical Riccia species by its spores having numerous areoles (60-70) on the proximal face.

42. Riccia crassifrons Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 571. 1885. Type. Brazil. Amazonas: Rio Negro, Spruce s.n. (holotype, BM).

Figs. 61, 62, 81: A

Riccia spruceana Steph., Sp. hepat. 1: 37. 1898. Type. Brazil. Amazonas: Rio Ramos, 1850, Spruce s.n. (holotype, G). Thalli in gregarious patches; lobes 7-10 mm long, 0.7-1.2 mm wide, rounded or slightly emarginate apically, 3-4 times forked, the basal dichotomy deep, 4-5 mm long; dorsal side light brown or light greenish brown when dry. Median groove shallow, short. Pores 20-40 µm diameter, bounded by 4 cells. Ventral scales absent. Cross section of lobes 0.5-0.7 mm high, as wide as high; dorsal edge projecting above the flank as wing with unistratose margin; flanks obliquely ascending, abruptly extending into the wings; ventral edge convex; air chambers in two layers; ventral tissue in 2-3 layers. Monoecious (fide Stephani, 1898). Sporangia bulging ventrally. Spores tetrahedral, light brown or red-brown, 55-72 µm diameter; wing vestigial, 1-5 µm wide, or absent; distal face with 6-8 areoles across diameter, with thin or slightly thickened ridges and weak tubercles, sometimes with a brown or pale tubercle in the centre; proximal face with areoles similar to those of distal face, with tubercles and sometimes a central tubercle, triradiate scar weak.

Bischler et al. 81 Distribution and ecology. Riccia crassifrons is rare and has been recorded only from Venezuela (Barinas, Infante-Sanchez & Heras Pérez, 2002), Peru (Loreto) and Brazil (Amazonas). It was collected on acidic silt on river borders. Specimens examined. PERU. LORETO: Prov. Maynas, Yanayaco near Iquitos, P. & E. Hegewald 6411 (Hb. Hegewald). BRAZIL. AMAZONAS: Type of R. crassifrons; type of R. spruceana. Riccia crassifrons can be distinguished from the other species of subgen. Ricciella by its thallus lobes consisting of dorsal and ventral epidermis bounding two layers of air chambers, by the thin lateral thallus wings, and by its spores with a narrow, irregular and partly vanishing wing.

43. Riccia dussiana Steph., in Urban, Symb. antill. 3: 275. 1902. Type. Martinique. Fonds St. Denis, Duss 577 (holotype, G).

Figs. 63, 66, 81: B

Thalli spongy, with gregarious lobes; lobes 5-7 mm long, 1.5-2 mm wide, rounded and slightly emarginate apically, 3-4 times forked, the basal dichotomies deep, the apical short; dorsal side pale yellowish when dry, or pale green with pale yellow, narrow, wavy and translucent margins. Median groove deep apically, vanishing below. Pores 55-85 µm diameter, bounded by 2 rings of 4 cells. Ventral scales short, not reaching lobe margins, hyaline, or violet, or spotted with violet. Cross section of lobes 0.4-0.5 mm high, 2-4 times as wide as high; dorsal edge with narrow, thin lateral wings ending in a single cell, often reflexed towards ventral side; flanks more or less concave, violaceous, or hyaline, or spotted with violet; ventral edge convex; dorsal tissue spongy, 1/3-1/2 of lobe height, cells with longitudinal, pitted strips; air chambers in 1(-2) layers; ventral tissue with thinwalled, pitted cells. Monoecious. Antheridial necks white, protruding. Archegonial necks colourless, hardly protruding. Sporangia bulging ventrally. Spores light brown, 72-86 µm diameter; wing yellowbrown, 5-7 µm wide, margin thickened and granulose; distal face with 8-11 areoles across diameter, with dark red-brown, thick ridges and weak tubercles; proximal face with areoles smaller than those of distal face, often incomplete, with irregularly thickened ridges, triradiate scar weak. Distribution and ecology. Riccia dussiana has been recorded from Martinique, French Guiana, and Peru (Loreto). It is rare and has been collected on siliceous soil, along paths and trails, at low elevation. Specimens examined. WINDWARD ISLANDS. MARTINIQUE: Type; Fonds St. Denis, near St. Pierre trail, M. & H. Stehlé 4022 (PC); Balata, Le Gallo 887 (PC). FRENCH GUIANA. Carbet Saramaca, Acarouany, Le Gallo 3, 4 & s.n. (PC); Salut Islands, Island Royale, Cremers 8642 (PC). PERU. LORETO: Maynas, 1.3 km E of Rio Sucasari, Timme 12269 (KSP).

Bischler et al. 82 Riccia dussiana is distinguished from other species of sect. Ricciella by its yellowish thalli and its thin, wavy lobe margins.

44. Riccia frostii Austin, Bull. Torrey Bot. Club 6: 17. 1875. Type. U. S. A. Nevada: without locality, Watson s.n. (holotype, NY).

Figs. 2 D, 64, 66, 81: C

Female thalli larger than male, in complete or incomplete rosettes, spongy, to 10 mm diameter, with crowed lobes partially creeping over each other; lobes 1-1.4 mm long, 1-1.3(-2) mm wide, rounded or truncate apically, 2-4 times forked, segments to 2 mm long; dorsal side green, or bluish green, or with pink spots on margins. Male thalli 0.3-1.5 mm diameter, spongy, rounded or with 2-4 lobes, often cross-shaped, pink on borders or over the whole dorsal side. Median groove short or absent. Pores small in younger parts, larger in older. Ventral scales absent. Cross section of lobes 0.30.5 mm high, 3-6 times as wide as high; dorsal edge obtuse laterally; flanks obliquely ascending; ventral edge hardly convex; air chambers 2/3-3/4 of lobe height, in 1-2 layers; ventral tissue of large cells. Dioecious. Archegonia deeply immersed, necks white or violet, protruding. Antheridial necks protruding, often purplish. Sporangia usually numerous, black, bulging mainly ventrally. Spores redbrown or yellow-brown, 40-65 µm diameter; wing narrow, 2-3 µm wide, punctate-vermiculate; distal face with irregular, branched, vermiculate ridges radiating from distal pole; proximal face with vermiculate ridges, often absent near proximal pole, triradiate scar of thin ridges. Gametophytic chromosome number n = 8. Distribution and ecology. Riccia frostii is widely distributed in central and southern Europe, the Mediterranean area, Saharan Africa, SW Asia, India, and China. It also has a wide range in the United States, where it has been recorded from 25 states (Schuster, 1992b). The species is rare, however, in Central and South America, where it has been collected in Mexico (Baja California (Howe, 1923)), Argentina (Neuquen, Rio Negro), and Uruguay, on sandy or clayey, calcareous or siliceous soil. The species is not known from the Neotropics. Specimens examined. ARGENTINA. NEUQUEN: Paso Flores, Hässel de Menéndez 305 (BA); Arroyito, Hässel de Menedez 314 (BA). RIO NEGRO: Río Pichileufu, Hässel de Menéndez 319 (BA); La Balsa, Pte General Roca, Ribeira Río Negro, Bachmann 10008 (BA) URUGUAY. Montevideo, Miguelete, Herter 78804 (PC). Riccia frostii can be distinguished from the other species of subgen. Ricciella by its spore ornamentation, with worm-like, thin ridges radiating from the distal pole.

45. Riccia geissleriana Jovet-Ast, Cryptog. Bryol. Lichénol. 14: 236. 1993. Type. Peru. Ancash: NE Huaraz, beyond laguna Llaca, 4250 m, Geissler 8885 (holotype, G).

Figs. 65, 66, 81: D

Bischler et al. 83 Thalli in gregarious patches; lobes 7-10 mm long, 3-3.5 mm wide, rounded-truncate and crisped apically; twice forked, the basal dichotomy 5-6 mm deep, the following 1-3.5 mm; dorsal side light brown when dry, margins wavy, hyaline or violaceous; flanks tinged with violet; ventral side violaceous. Median groove deep, wide and convex. Subepidermal cells with slightly thickened walls. Pores bounded by 4 cells. Ventral scales large, reaching lobe margins, hyaline apically, violaceous basally. Cross section of lobes 2.5-4 times as wide as high; dorsal edge acute with short wings, tinged with violet, ending in a large basal cell and a narrow, acute, terminal cell; ventral edge weakly convex; air chambers in one layer, partitions 3-5 cells high; ventral tissue of small, thin-walled cells. Monoecious. Archegonial necks violaceous. Sporangia numerous. Spores tetrahedral, light brown, 7296 µm diameter, wingless; distal face with 18-20 areoles across diameter, each of 3-7 µm diameter, convex inside, ridges without tubercles; proximal face with three well distinct facets, with 50-60 areoles per facet, areoles less regular in shape but of the same size than those of distal face, without tubercles, triradiate scar indistinct. Distribution and ecology. Riccia geissleriana has been collected once only, at high elevation in the Peruvian Andes (4250 m), on acidic soil in a Polylepis woodland. Specimen examined. PERU. ANCASH: Type. Riccia geissleriana resembles R. chiapasensis, a.o. by the similar lobe width, and also shows characteristics of subgen. Riccia (general habit, scale length, air chambers in single layer, simple pores). However, the air chambers are wide as in subgen. Ricciella. The simple pores, without distinct bounding cells, separate R. geissleriana from all other neotropical species of the subgen. Ricciella.

46. Riccia hegewaldiana Jovet-Ast, Cryptog. Bryol. Lichénol. 14: 238. 1993. Type. Peru. La Libertad: prov. Otuzco, Casmiche, 2000 m, P. & E. Hegewald 5050 (holotype, Hb. Hegewald). Figs. 67, 70, 81: E Thalli in gregarious patches; lobes 5-8 mm long, 2-4 mm wide, slightly emarginate apically, 2-3 times forked, the basal dichotomies deep; dorsal side pale light brownish green when dry, margins wavy and crisped. Median groove very short. Epidermal cells granulose. Pores 40-55 µm diameter, bounded by 5-6 hyaline, thin-walled cells. Ventral scales hyaline, large but not extending beyond lobe margins. Cross section of lobes 0.4-0.7 mm high, to seven times as wide as high; dorsal side convex and acute laterally; flanks obliquely ascending, slightly concave and recurved apically; ventral edge nearly flat; air chambers in two layers, or in one layer and then high and narrow, partitions 1, rarely 2 cells wide; ventral tissue 1/3-1/4 of lobe height. Monoecious? Archegonial necks violet. Sporangia hardly bulging dorsally. Spores tetrahedral, red-brown with darker ridges, 65-79 µm diameter; wing 5-6(-12) µm wide, regular, thickened on margin, pores large; distal face with approx. 8 ill-delimited areoles across diameter, with thick and more or less sinuose ridges radiating from pole to equator; proximal face with numerous, small, often incomplete areoles, with slender ridges and tubercles,

Bischler et al. 84 triradiate scar weak. Distribution and ecology. Riccia hegewaldiana has been collected once only, on clayey, acidic soil, on the borders of a trail, at 2000 m elevation. Specimen examined. PERU. LA LIBERTAD: Type. Riccia hegewaldiana is related to R. frostii and R.. paraguayensis by its spore ornamentation pattern but the thallus lobes are thicker than those of R. paraguayensis and the thalli do not form rosettes. From R. frostii the species differs, moreover, by its wider, dorsally pale green lobes with scales reaching lobe margins, and the larger spores.

47. Riccia jovet-astiae Vianna, Bol. Inst. Bioci. Univ. Fed. Rio Grande do Sul 38: 165. 1985. Type. Brazil. Rio Grande do Sul: Canoas, Kay Saafeld 2706 (holotype, ICN).

Figs. 68, 70, 81: F

Thalli in gregarious patches; lobes 10-13 mm long, 0.7-1(-1.5) mm wide, rounded and slightly emarginate apically, sometimes tripartite, 4-5 times forked, the basal dichotomies 2.5 mm deep, the apical 1-2 mm; dorsal side light yellow-green to dark yellow-green when dry. Median groove short, deep apically, shallow and evanescent towards base. Pores 36 µm diameter, bounded by 4 cells and by an additional ring of 4-5 larger cells. Ventral scales rounded apically, 0.3 mm wide. Cross section of lobes 0.3-0.6 mm high, much wider than high; dorsal edge rounded or obtuse laterally; flanks slightly ascending; ventral edge convex; air chambers in 2-3 layers; ventral tissue in 2-4 layers, of smaller cells than those of dorsal tissue. Monoecious. Archegonial necks red-violet. Sporangia strongly bulging ventrally. Spores tetrahedral, light brown or brown, 59-84 µm diameter; wing 3.5-7 µm wide, pale, nearly smooth, with granulose margin; distal face with 4-5(-8) areoles across diameter, with dark brown ridges without tubercles and, in each areole, a central thickening or a branched ridge forming an X or Y; proximal face with 10-15 irregular, complete or incomplete areoles per facet, with thin, brown, somewhat sinuose ridges, triradiate scar well marked. Distribution and ecology. Riccia jovet-astiae is known only from southern Brazil where is seems to be infrequent. It has been collected on sandy, acidic soil, in gardens and on river borders. Specimens examined. BRAZIL. RIO GRANDE DO SUL: Type; Cascata S. Miguel, Vianna 626 (ICN); Porto Alegre, Fac. de Direito, Vianna 665 (ICN); Porto Alegre, Ponta Grossa, Vianna 3870 (ICN); R. F. do Turvo, Tenente Portela, Vianna 4261 (ICN). Riccia jovet-astiae is related to members of the R. fluitans species complex. The lobes are wider than those of R. stenophylla, the neotropical representative of this species complex, and the spores have a different ornamentation pattern.

48. Riccia limicola Jovet-Ast, Rev. Bryol. Lichénol. 44: 422. 1978. Type. Ecuador. Galapagos Is.:

Bischler et al. 85 Santa Cruz, beside Alf Karstdalens house above Bella Vista, Weber B-41244 (holotype, PC; isotype, U).

Figs. 69: A-N, 70, 81: G

Thallus lobes 0.3-0.5(-0.7) mm wide, rounded or emarginate apically; 2-3 times forked, basal dichotomy deep, 6-10 mm long, apical shorter, 1-2.5(-4) mm long; dorsal side greyish green or light green and shiny. Median groove deep on the terminal branch, evanescent elsewhere. Pores 25 µm diameter, bounded by 4 thin-walled cells. Ventral scales not seen. Cross section of lobes 2-3 times as wide as high near apex of lobes to 5 times wider at base; dorsal edge rounded laterally; flanks obliquely ascending; ventral edge convex; air chambers in 1-2 layers; ventral tissue in approx. 5 layers. Monoecious. Sporangia bulging ventrally. Spores tetrahedral to spherical, brown or redbrown, 80-92 µm diameter; wing light brown, 5-7 µm wide, granulose and wavy; distal face with (6-) 7-8 areoles across diameter, with thick ridges and triangular tubercles in the corners, and small tubercles on the ridges; proximal face with 12-17 areoles per facet, smaller than those of distal face, triradiate scar weak. Distribution and ecology. Riccia limicola is rare, known only from the Galapagos Islands. It has been collected on acidic, clayey soil, on borders of trails, around pools, and in summit grassland, from sea level to about 650 m. Specimens examined. ECUADOR. GALAPAGOS ISLANDS: Type; Santa Cruz, base of El Puntudo, Weber B-412438 (U); vicinity of Academy Bay, La Copa, Weber B-13, 191 (U). Riccia limicola is related to R. stenophylla but its lobes are less subdivided, with basal dichotomies deeper, more than 6 mm long. The areoles on the distal face of the spores are complete, with numerous tubercles on the ridges, but without central ornamentation inside. On the proximal face, the triradiate scar is less marked, and the ridges are more conspicuous.

49. Riccia paraguayensis Spruce, Bull. Soc. Bot. France 36 (suppl. Congr. Bot. 1889): CXCVI [196]. 1890. Type. Paraguay. Near Asunción, Balansa 1280 (holotype, PC; isotype, G). Figs. 69: O-R, 71, 81: H Ricciella macropora Steph., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 23 (III, 2): 30. 1897. Type. Paraguay. Asunción, Lindman 220b (isotypes, G, S). Female thalli in rosettes of 2-4 cm diameter, male thalli smaller, 1 cm diameter; lobes 10-15 mm long, 2-3 mm wide, thin, translucent, obtuse and slightly emarginate apically; 3-4 times forked, the basal dichotomy deep and diverging; dorsal side bright green and shiny when dry. Median groove distinct apically, evanescent below. Pores rounded or ovate. Ventral scales absent. Cross section of lobes very thin, of few layers, often restricted on the lateral sides to dorsal and ventral epidermis and one layer of air chambers. Dioecicous. Antheridial necks white, protruding. Sporangia bulging ventrally. Spores brown, 60-70(-80) µm diameter; wing thin, yellowish, 2-3 µm wide, granulose;

Bischler et al. 86 distal face with 9-10(-13?) areoles across diameter, with thin ridges and small tubercles; proximal face with 30-40 areoles per facet, rarely with a central tubercle, smaller than those of distal face, triradiate scar thin. Distribution and ecology. Riccia paraguayensis is known from Brazil (Mato Grosso do Sul, Rio Grande do Sul, São Paulo), Paraguay, Argentina (Buenos Aires (Hässel, 1963), Corrientes, Jujuy (Hässel, 1963), Misiones (Hässel, 1963), Tucumán (Müller 1955)), and Uruguay. This rare species has been collected on sandy, granitic, shaded soil, at water border and in cultivated areas, from sea level to 700 m. Specimens examined. BRAZIL. MATO GROSSO DO SUL: without locality, Schäfer-Verwimp 8453 (Hb. Schäfer-Verwimp). RIO GRANDE DO SUL: Porto Alegre, Ponta Grossa, Vianna 4580 (ICN); Porto Alegre, Fac. de Direito, Vianna 13 (ICN); Guaiba, Gueitteito 6395 (ICN); Cachoeira do Sul, Vital 9235, 9242, 9229 (SP). SÃO PAULO: Pirajú, Vital 2176 (SP). PARAGUAY. Type; type of R. macropora. ARGENTINA. CORRIENTES: border of river Paraná, Bonpland 323 (PC). URUGUAY. Dep. Colonia, Estanzuela, Herter 8, 3592 (PC). The shiny lobes of Riccia paraguayensis recall R. membranacea, but the two species have completely different spore ornamentation patterns. Hässel de Menéndez (1963) treated R. subsimilis as a synonym of R. paraguayensis. However, by its spores remaining in tetrads R. subsimilis is a synonym of R. curtisii.

50. Riccia paranaensis Hässel, Opera Lilloana 7: 228. 1963. Type. Argentina. Buenos Aires: delta of Río Paraná, Arroyo Abra Vieja, Nov 1956, Hässel de Menéndez 3 (holotype, BA). Figs. 72, 73, 81: I Thalli in complete or partial rosettes, 0.7-3 cm diameter, often with crowded lobes in 2-3 layers; lobes 1-3 mm long, 0.5-1.2 mm wide, deeply 3-4 times forked, rounded or emarginate apically; dorsal side light green or yellowish green when dry, basally violet, or tinged with violet or pink only on margins and ventral side. Median groove deep, distinct apically on terminal segments only. Pores bounded by 3-4 cells. Ventral scales absent, or apically present, colourless. Cross section of lobes 22.5 times (towards base of lobes 3-4 times) as wide as high; dorsal edge rounded laterally; flanks oblique; ventral edge hardly convex; air chambers in 1-3 layers; ventral tissue in 3-5 layers. Monoecious. Archegonial necks protruding, 25 µm long, violet. Sporangia bulging ventrally. Spores light brown with darker ornamentation, or red-brown, 62-76(-84) µm diameter; wing irregular and narrow, and weakly undulate, sometimes discontinuous in part, more or less granulose, 2-2.5 µm wide; distal face with 5-8 areoles across diameter, with thin ridges and triangular tubercles; proximal face with approx. 20 areoles with thin ridges per facet, smaller than those of distal face, triradiate scar

Bischler et al. 87 weak. Distribution and ecology. Riccia paranaensis is known from southeastern Brazil (Espíritu Santo, Mato Grosso, Rio Grande do Sul, São Paulo) and Argentina (Buenos Aires). The species is infrequent and has been collected on humid sand and acidic soil, exposed or shaded, on borders of trails and at water edges, in lowland areas. Specimens examined. BRAZIL. ESPÍRITU SANTO: Linhares, Rio Doce, Vital 1874, 1875 (SP). MATO GROSSO: Miranda, Vital 2323, 2324 (SP); Corumba, Fazenda Santa Clara, Schäfer-Verwimp 8453 (Hb. Schäfer-Verwimp). RIO GRANDE DO SUL: São Lourenço, Vital 8957 (SP); Cachoeira do Sul, Vital 9227, 9251 (SP); Torres, Lagoa Itapova, Vianna 72 (ICN); T. F. do Turvo, Tenente Portela, rio Miuguai, Oliveira 1312, Vianna 6389 (ICN); Ivoti, Cascata São Miguel, Homrich 1660 (ICN); Sta Maria, Boca do Monte, Vianna 4558 (ICN). SÃO PAULO: Serra do Mar, Schäfer-Verwimp & Verwimp 14992 (Hb. Schäfer-Verwimp). ARGENTINA. BUENOS AIRES: Type. Riccia paranaensis is related to R. stenophylla with which it shares narrow lobes. However, the thalli of R. paranaensis are crowded and arranged in several layers in the patches, and the spore wall of the distal face lacks ridges or tubercles inside the areoles but instead has tubercles on the ridges. Riccia paranaensis may also be compared to R. huebeneriana Lindenb. In the latter species the lobe dichotomies are distinctly shorter, the dorsal side of the lobes is bright green, often tinged with purple or violet near apex and above sporangia, and the scales are often violet. In R. paranaensis, the scales are hyaline and the spores are smaller.

51. Riccia stenophylla Spruce, Bull. Soc. Bot. France 36 (suppl. Congrès Bot. 1889): CXCV [195]. 1890. Type. Paraguay. Asunción, Balansa 3706 (holotype, PC).

Figs. 74, 75, 81: K

Riccia oerstediana Lindenb. & Hampe, in Hampe, Linnaea 24: 304. 1851. Type. Costa Rica. Without locality, Oersted s.n. (holotype, G). Riccia hosseusii Herzog, Feddes Repert. Nov. Spec. Regni Veg. 55: 5. 1952. Type. Argentina. Formosa: Dept. Pirané, Río Pilaga, Hosseus 531/A (holotype, JE) Thallus in crowded mats or forming partial rosettes; lobes 3-5(-20) mm long, 0.3-0.5 mm wide, slightly wider at level of sporangia, 2-5 times forked, obtuse or slightly emarginate apically; dorsal side light green. Median groove distinct apically, vanishing below. Pores bounded by 4(-6) cells, irregular in size. Ventral scales semi-lunate, sometimes the apical conspicuous, extending beyond lobe apex. Cross section of lobes 2-3 times as wide as high; dorsal edge rounded or obtuse laterally; flanks oblique; ventral edge convex; air chambers in 2-3 layers; ventral tissue in 2-4 layers. Monoecious. Sporangia large, 0.5 mm diameter, bulging ventrally. Spores brown or light red-brown, 65-74 µm diameter; wing lighter, complete, 5-7 µm wide, tuberculate-papillose; distal face with 4-5

Bischler et al. 88 more or less complete areoles across diameter, each with a tubercle or a simple or branched ridge in the centre; proximal face with areoles smaller than those of distal face near equator, delimited by low ridges, becoming incomplete towards pole and replaced by more or less sinuose and branched ridges, triradiate scar well marked. Gametophytic chromosome number n = 8. Distribution and ecology. Riccia stenophylla is known from the United States (Florida, North Carolina, Texas) and the Neotropics: Mexico (Aguascalientes, Jalisco), Guatemala, Costa Rica, Panama, Peru (Cajamarca, Piura), Brazil (Bahía, Espíritu Santo, Goiás, Mato Grosso, Paraná, Pernambuco, Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo), Paraguay, Argentina (Buenos Aires (Hässel, 1963), Cordoba (Hässel, 1963), Formosa), and Uruguay. The species seems to be quite common and grows on sandy or clayey, sometimes granitic soil, or on rocks, exposed or sheltered, in gardens, plantations, and along forest borders, from low elevations to 2000 m. Neotropical specimens examined. MEXICO. Without locality, Arsène 7833, 7836 (G). AGUASCALIENTES: 0.9 mi. S of Hwy. 70 on turnoff to Tapas Viejas, 12 mi. W of Aguascalientes, Whittemore 1784 (Hb. Whittemore). JALISCO: 16.8 mi. NW of Ayutla, road to Los Volcanes and Talpa de Allende, Whittemore 2164 (Hb. Whittemore). VERACRUZ: Xalapa town, Hotel garden, Long 29611 (E); Rio Jamapa between Huatusco and Coscomatepec, Long & Garcia 29685 (E). GUATEMALA. Naranjo, N of Chicacao, Hürlimann H 518 (PC). COSTA RICA. Type of R. oerstediana; San José, 1979, Jovet-Ast s.n. (PC); El Rodeo, 1979, Jovet-Ast s.n. (PC); Turrialba, 1979, Jovet-Ast s.n. (PC); near Alajuela, 1979, Jovet-Ast s.n. (PC); Santa Ana, 1979, Melendez-Howell s.n. (PC); Tres Rios, Melendez-Howell s.n. (PC); Paso Ancho near San José, 1979, Melendez-Howell s.n. (PC). PANAMA. Prov. Cocle, Las Mozas, Valle de Anton, Salazar et al. 13498 (PMA). PERU. CAJAMARCA: Contumaza, near Tambo de Lima, between Contumaza and Cascas, Hegewald 7394 (Hb. Hegewald). PIURA: Huancabamba, Porculla-Pass, Hegewald 7161, 7162 (Hb. Hegewald). BRAZIL. BAHÍA: Itabuna, Schäfer-Verwimp & Verwimp 12930 (Hb. Schäfer-Verwimp); Mun. Urandi, Vital 7932 (SP). ESPÍRITU SANTO: 8 km E from Linhares, Vital 1928 (SP); Linhares, Vital 1868 (SP); Vila Velha, Morro da Penha, Schäfer-Verwimp 8837 (Hb. Schäfer-Verwimp). GOIÁS: Ceres, Vital 3040 (SP). MATO GROSSO: Miranda, Vital 2358 (SP). PARANÁ: Cachoeiras de Sete Quedas, Prainha, Vital 8362 (SP). PERNAMBUCO: Reserva de Saltinho, Reserva de Brejo, Porto (PC); Garanhuns, Schäfer-Verwimp & Verwimp 12987 (Hb. Schäfer-Verwimp). RIO DE JANEIRO: Glaziou 3822, 5399 (PC). RIO GRANDE DO SUL: Mun. Bom Jesus, Vital 2021, 2124 (SP); Bagef, Bueno 2678 (ICN); San Leopoldo, Vianna 2712 (ICN); R. F. do Turvo, Tenente Portela, Bueno 4599 (ICN). SANTA CATARINA: Mun. Lajes, Vital 9388 (SP). SÃO PAULO: Mun. Matão, Vital 2182 (SP); Mun. Buritizal, Vital 2405 (SP); Ibira, Termas, Vital 2294 (SP); Rio Santo Anastacio, Vital 2546 (SP); 1.5

Bischler et al. 89 km SW Pilar do Sul, Vital 2601 (SP); Mun. Porto Feliz, Vital 2461 (SP); Mun. Teodoro Sampaio, Porto Angeli, Vital 2173 (SP); Bauru, Vital 2514 (SP); Mun. Guararapes, Vital 2480 (SP); Litoral, Itanhaem, Schäfer-Verwimp & Verwimp 11132 (Hb. Schäfer-Verwimp); Litoral, Restinga near Bertioga, Schäfer-Verwimp & Verwimp 7745 (Hb. Schäfer-Verwimp); Serra do Mar, Ubatuba, Schäfer-Verwimp 9270 (Hb. Schäfer-Verwimp); Aguas de Santa Barbara, Schäfer-Verwimp & Verwimp 9466 (Hb. Schäfer-Verwimp). PARAGUAY. Type; Pto Hayes, Geissler 14720, 14740, 14954, 14955, 14982 (G); Boquerón, Petito Alonsito, Geissler 14986 (G). ARGENTINA. FORMOSA: Type of R. hosseusii. URUGUAY. San José, Herter 86056 (PC). Riccia stenophylla resembles Riccia fluitans and Riccia limicola. However, the spores of the latter species have 5-7 areoles with granulose ridges across distal face, and the areoles of the proximal face are complete. Riccia fluitans has granulose spores with 5-7 areoles across diameter of the distal face, without tubercles or ridges in the centre, whereas in R. stenophylla the spores are not granulose and have 4-5 areoles across diameter of distal face, with tubercles or ridges in the centre. The spores of Riccia canaliculata Hoffm., a species known from Europe, the Mediterranean and North America, are larger (80-97 µm).

2d. Subgen. RICCIELLA sect. SPONGODES Nees, Naturgesch. eur. Leberm. 4: 391. 1838. Type. R. crystallina L. emend. Raddi. Sect. Cavernosae R.M.Schust., J. Hattori Bot. Lab. 71: 270. 1992 (Hep. N America 6: 486. 1992). Type. R. cavernosa Hoffm. Thallus spongy, shallowly divided into wide lobes. Sporangia immersed, hardly bulging. Sect. Spongodes has a wide distribution over all continents.

KEY TO THE NEOTROPICAL SPECIES OF SECT. SPONGODES 1. Lobes green or yellow-green dorsally. Pores large. Spores red-brown, with papillose wing and incomplete areoles on distal face, with ridges drawing at pole four squares, or an X, or an Y ............... ...................................................................................................................................... 53. R. cavernosa 1. Lobes blue-green dorsally. Pores very small. Spores yellow-brown, with crenulate wing, and 9-10(11) well delimited areoles across diameter of distal face ........................................... .52. R. crystallina

Bischler et al. 90 52. Riccia cavernosa Hoffm., Deutschl. Fl. 2: 95. 1795. Type. Germany. Harz Mts. near Göttingen, Hoffmann s.n. (n.v.).

Figs. 76: A-G, 77, 81: L

Riccia brandegei Underw., Bot. Gaz. 19: 275. 1894. Type. Mexico. Baja California: Brandegee s.n. (n.v.) Riccia catalinae Underw., Bot. Gaz. 19: 275. 1894. Type. U. S. A. California: Santa Catalina I., McClatchie 44 (n.v.). Riccia montagnei Steph., Sp. hepat. 1: 44. 1898. Type. Chile. Without locality and collector (holotype, PC; isotype, G). Thalli in rosettes of 35 mm diameter, spongy, deeply divided to 3-8 mm, or in gregarious patches; lobes narrow, 1-2(-3) mm wide, rounded or emarginate apically; dorsal side green or yellow-green when dry, sometimes tinged with pink on margins and above sporangia. Median groove indistinct. Epidermis often disintegrating. Pores small at apex, larger and numerous in older parts of lobes, 2040 µm diameter, bounded by 4(-5) cells. Ventral scales absent. Cross section of lobes 3-5 times as wide as high, not narrowed into lateral wing; air chambers in 2-4 layers, 2/3-3/4 of lobe height. Monoecious. Antheridial necks colourless, hardly protruding. Archegonial necks pink violaceous, hardly protruding. Sporangia black, mainly bulging ventrally. Spores tetrahedral, 150-300 per sporangium, red-brown or black-brown, 60-120 µm diameter; wing 3.5-6 µm wide, light red-brown, papillose, complete or locally narrowed and becoming evanescent, sometimes weakly thickened at margin; distal face with dark red-brown ridges, irregularly crenulate, with complete areoles at periphery or with a branched net, delimiting 4 squares at the pole, with central short ridge or tubercle, or drawing an Y or an X; proximal face with incomplete areoles or with short and distant ridges, or with tubercles mixed with short ridges, triradiate scar well marked. Gametophytic chromosome number n = 8. Distribution and ecology. Riccia cavernosa is widely distributed in central Europe to Scandinavia, in the Mediterranean area, Macaronesia (Canary Is.), Africa (Ethiopia, Cameroon, Zaire, eastern Africa, South Africa), Asia (SW Asia, Siberia, Mongolia), and Australia. It has been recorded from the United States (25 states) and the Neotropics: Mexico (Baja California (Underwood, 1894), Querétaro), Bahamas (Evans, 1911), Ecuador (Cotopaxi), Peru (Cajamarca, La Libertad), Brazil (Rio Grande do Sul), Chile, Paraguay (Hässel, 1958), and Argentina (Buenos Aires (Hässel, 1963), Chubut (Hässel, 1963), Neuquén (Hässel, 1963), Rio Negro (Hässel, 1963)), perhaps also Cuba and Trinidad (Schuster, 1992b). The species seems to be quite common and has been collected on calcareous or siliceous, sandy or clayey soil, near water edges, from sea level to 2800 m. Neotropical specimens examined. MEXICO. QUERÉTARO: N bank of Rio Santa Maria near Ayutla, Long & Delgadillo 29723 (E). ECUADOR. COTOPAXI: Latacunga, Volk 6/0702 (JE).

Bischler et al. 91 PERU. CAJAMARCA: Contumaza, near Tambo de Lima between Contumaza and Cascas, P. & E. Hegewald 7397 (Hb. Hegewald). LA LIBERTAD: Otuzco, Samne, between Trujillo and Otuzco, P. & E. Hegewald 9522 (Hb. Hegewald). BRAZIL. RIO GRANDE DO SUL: Torres, Parque da Guarita, 1974, Vianna s.n. (ICN). CHILE. Type of R. montagnei. The spore wall ornamentation in R. cavernosa is quite variable. The polar region of the distal face shows thickenings forming an X or an Y; sometimes, the branches of the X and the Y extend towards the ridges of the marginal areoles and draw 3-4 central areoles with some tubercles and ridge fragments in the centre, especially in Argentinean specimens. The ornamentation of the proximal face also varies, from complete areoles with thick ridges to individual, incurved, sinuose and branched ridges. Riccia cavernosa might comprise allopatric variants. Schuster (1992b) treated Riccia bahiensis as a synonym of R. cavernosa.

53. Riccia crystallina L., Sp. pl. ed. 1, 1138. 1753. Type. Europe (OXF-Dill., H-SOL). Figs. 76: H-P, 77, 81: M Riccia ochrospora Mont. & Nees ex Lindenb., Nova Acta Phys.-Med. Akad. Caes. Leop.-Carol. Nat. Cur. 18(1): 504b. 1837. Type. Chile. Valparaiso: Quillota, Bertero 1279 (holotype, PC). Riccia plana Taylor, London J. Bot. 5: 414, 1846. Type (Perold, 1991). Australia, Swan River, 1843, Drummond s.n. (holotype, K; isotypes, MEL). Thallus in dense, spongy rosettes, 10-15 mm diameter, with few, deep partitions, sometimes in gregarious patches; lobes approx. 2 mm long, to 14 mm wide, subdivided into short segments 2 mm wide, truncate, or rounded, or emarginate apically; dorsal side light blue-green and crystalline when moist, bluish green when dry. Median groove deep or absent. Pores small, larger towards central part of the rosettes. Ventral scales absent. Cross section of lobes 6 layers high; air chambers numerous in upper 3/4; ventral tissue thinner than dorsal, of large cells with pitted walls. Monoecious. Antheridial necks white, protruding. Archegonial necks pink, hardly protruding. Sporangia more numerous towards base of lobes, bulging ventrally. Spores yellow-brown, with darker ridges, approx. 400 per capsule, 60-87 µm diameter; wing complete, 4-6 µm wide, margin crenulate; distal face with 9-10 (11) complete areoles across diameter, with rounded or bifid tubercles at corners of ridges; proximal face with complete areoles with similar tubercles, triradiate scar well marked, dark brown. Gametophytic chromosome number n = 8. Distribution and ecology. Riccia crystallina has a wide range in Atlantic Europe, the Mediterranean area, Macaronesia, tropical and southern Africa (Ethiopia, South Africa), Australia and New Zealand. It has been recorded also from Arizona. In the Neotropics the species is rare and is

Bischler et al. 92 known from Mexico (Mexico), Chile (O'Higgins (Gottsche et al. 1847), Valparaiso), Argentina (Rio Negro), and Uruguay. Many former records may belong to R. cavernosa, or to other species; those of the Bahamas (Evans 1911), Peru (Montagne, 1845b), Brazil (Bahía, Nees, 1833), Paraguay (Spruce, 1888b) need confirmation. Riccia crystallina has been collected in the Neotropics on humid, weakly acidic or calcareous silt, mud, and sandy or clayey soil, from sea level to 3300 m. Neotropical specimens examined. MEXICO. MEXICO: Popocatepetl, 15 km above Amecameca, Düll 2/395 (Hb. Düll). CHILE. VALPARAISO: Type of Riccia ochrospora. ARGENTINA. RIO NEGRO: Rio Pichileufu, Hässel de Menéndez 316 (BA). URUGUAY. Montevideo, Miguelete, Herter 77804 (PC); Dep. Colonia, Estanzuela, Herter 3592 (PC). Many authors have considered Riccia crystallina and R. cavernosa as conspecific (others used the name R. plana; e.g., Hässel, 1963). However, the different thallus colour (bluish or glaucous in R. crystallina, yellow-green in R. cavernosa), spore colour (yellow-brown in R. crystallina, red-brown to dark-brown in C. cavernosa), and spore ornamentation (compete areoles in R. crystallina, incomplete ones in R. cavernosa) readily distinguish the two species.

3. RICCIOCARPOS Corda, in Opiz, Beitr. Naturg. 651. 1829. Type. Ricciocarpos natans (L.) Corda. Named for the sporangia resembling those of Riccia. The genus is monospecific.

1. Ricciocarpos natans (L.) Corda, in Opiz, Beiträge z. Naturgesch. 651. 1829. Riccia natans L., Syst. Nat., ed. 10, 1339. 1759. Type (Grolle, 1976). Europe. Great Britain: Suffolk, near Hadley, Buddle s.n. (holotype, OXF, n.v.; isotype, H-SOL, n.v.).

Figs. 2 : F, 82, 83, 104: C

Riccia natans var. brasiliensis Mont., Ann. Sci. Nat., Bot. sér. 2, 12: 49. 1839. Brazil. Rio Grande do Sul: Aquilegio near Porto Allegre, St. Hilaire s.n. (PC). Ricciocarpos natans fo. terricola Steph., Sp. hepat. 1: 54. 1898. Types from various continents. Neotropical syntypes: Brazil, Ule, Lindman; Mexico, Schaffner (n.v.). Ricciocarpos velutinus (Wilson ex Hook.) Steph., Sp. hepat. 1: 55. 1898. Riccia velutina Wilson ex Hook., Ic. Pl. pl. 249. 1849. Type. U. S. A. Texas: Drummond s.n. (syntype, BM). Thallus yellow green to green, often tinged with purple, forming hemi-rosettes or rosettes,

Bischler et al. 93 segments 2-5 mm wide, with groove apically. Epidermis well developed, often with oil-cells. Epidermal pores with single ring of 4-8 cells, radial walls thin. Air chambers in 2 to several layers. Ventral tissue in 3-4 layers, with oil-cells. Ventral scales usually purplish, in several ill-defined, transverse rows, very long in floating forms, shorter in terrestrial, with oil-cells, without marginal papillae, margins serrulate. Asexual reproduction by specialized propagules lacking. Monoecious. Antheridia in dorsal groove along thallus midline. Archegonia in 2-3 rows in dorsal groove. Spores approx. 450 per capsule, 42-72 µm diameter, proximal and distal faces with more or less complete, tuberculate areoles and tuberculate ridges, trilete scar distinct. Gametophytic chromosome number n = 9. Distribution and ecology. The distribution of the species is worldwide in the Northern and Southern Hemispheres, from 62°N to 45°S (Europe, the Mediterranean area, tropical and temperate Africa, SW Asia, northern Asia, India, China, Japan, Australia, and New Zealand), and includes Canada, U. S. A. (recorded from 35 states) and Central and South America. In the Neotropics it is known from Mexico (Jalisco, Mexico, Morelos, San Luis Potosí, Tabasco), Panama, Venezuela (Infante-Sanchez & Heras Pérez, 2002), Brazil (Amazonas, Bahía (Yano, 1981b), Espíritu Santo (Yano, 1981b), Pará, Paraná (Yano, 1989), Pernambuco (Porto & Yano, 1985), Rio Grande do Sul, Santa Catarina (Yano, 1981b; Vianna, 1976), São Paulo), Bolivia, Chile (Valparaiso), Argentina (Buenos Aires, Formosa, Entre Ríos (all recorded by Hässel, 1963)), and Uruguay; perhaps also from Cuba (Howe, 1923). Ricciocarpos natans grows floating on stagnant water, at the margins of temporary ponds, streams and in swamps, or on very wet soil. It is rare everywhere and has rarely been seen with sporophytes. The species is not drought-tolerant but is probably able to shorten its life cycle to a few weeks. It seems to be absent from areas with long, cold winters. In the Neotropics, R. natans has been collected from sea level to 2800 m. Specimens examined. MEXICO. JALISCO: Laguna Escondida, ca 30 km N of Ciudad Colima, Novelo & Mijangos 938 (MEXU). MEXICO: Laguna de San Pedro Tultapec de Quiroga, 2 km S of Lerma, Novelo 527 (MEXU). MORELOS: Lago Hueyapan, Parque Nacional Lagunas de Zempoala, 8 km N of Hutzilac, Bonilla 596 (MO). SAN LUIS POTOSÍ: Schaffner 4 (NY). TABASCO: Ejido el Capricho, 8-10 km N of Plan Chontalpa, Cowan 2824 (MEXU, MO, NY). PANAMA. Vicinity of Juan Mina, Chagres River, Bartlett & Lasser 16828 (NY); La Jagua, McDaniel 8223 (NY). PERU. LORETO: Yurimaguas, lower Huallaga, Killip & Smith 27708 (NY). BRAZIL. Tropical Brazil, Burchell 7426 (NY). AMAZONAS: Iguapó forest, near Manaus, Griffin III, Vital & Yano 819 (FLAS). MATO GROSSO: Rio Tapuari, Fazenda Taruma, Prance & Schaller 26263 (NY). PARÁ: Santarem, Spruce 843 (NY). RIO GRANDE DO SUL: type of Riccia natans var. brasiliensis. (PC). SÃO PAULO: Campinas near Jaguary, Mosén GY (NY); Ilha Grande do

Bischler et al. 94 Paranapanema, Fazenda Bella Vista, Rio Turvo, Schiffner 2261 (TENN); near S. Anna, rio Tieté, near São Paulo, Schiffner 331 (TENN). BOLIVIA. SANTA CRUZ: Velasco Prov., Arroyo Toledo, Ritter 2449 (MO), Reserva El Refugio, Guillen 3309 (MO, U). CHILE. VALPARAISO: Quintero, Bosque Las Petras, Mahu 12928 (PC). URUGUAY. S. José, Barra, Herter 322 A (NY). The main diagnostic feature distinguishing Ricciocarpos natans from other Marchantiales are the scales with serrulate margins. In terrestrial populations the scales are short and wide whereas in floating populations they are very long and often slender. Aquatic populations of R. natans have sometimes been distinguished taxonomically from terrestrial ones, but other diagnostic characters have not been found. In the Neotropics, Ricciocarpos velutinus, R. natans var. brasiliensis and R. natans fo. terricola belong to the terrestrial form.

3. CORSINIACEAE Engl., Syll. Pflanzenfam., Grosse Ausgabe 44. 1892. Type. Corsinia Raddi. Thallus green or whitish, tinged or not tinged with purple or violet; branching dichotomous. Epidermis sometimes partly disintegrating in older parts of thalli. Epidermal pores simple, bounded by a single ring of 5-7 cells, without hyaline inner ring. Air chambers in 1(-2) layers, wide, with at least rudimentary chlorophyllose filaments. Basal tissue thick. Oil cells present. Ventral scales in several ill-defined rows, or in 2 rows, with filiform appendages. Asexual reproduction by specialized propagules lacking. Monoecious or dioecious. Male and female gametangia dorsal. Antheridia in groove on median line of thallus. Archegonia in cavities. Calyptrae thick, warty, or 1-4-layered, smooth. Involucre a posterior scale. Sporophytes remaining included in thallus tissue, with few-celled foot and short seta. Capsules cleistocarpous, wall with or without annular thickenings. Spores few, large, mixed with sterile cells or with short elaters, not more than twice as long as wide, with single helical band. Gametophytic chromosome number x = 8, or 9. The family includes two genera, which are both represented in the Neotropics. Schuster (1992b) considered them to be sufficiently distinct to represent two subfamilies, Corsinioideae and Cronisioideae.

KEY TO THE GENERA OF CORSINIACEAE 1. Thallus without purplish pigmentation. Ventral scales in several ill-defined rows. Calyptra thick, warty. Involucre a thick, pluristratose posterior scale, not involute around the archegonia. Capsule wall without annular thickenings. n=8 or 16.......................................................................4. Corsinia

Bischler et al. 95 1. Thallus often with purplish pigmentation. Ventral scales in two rows. Calyptra thin (1-4 cell layers), not warty. Involucre a thin, unistratose scale, involute around the archegonia. Capsule wall with or without annular thickenings. n=9........ ...................................................................5. Cronisia

4. CORSINIA Raddi, Opusc. Sci. 2: 354. 1818. Type. C. coriandrina (Spreng.) Lindb. Genus dedicated to the Italian botanist Tomas Corsini. The genus is monospecific.

1. Corsinia coriandrina (Spreng.) Lindb., Hepaticologiens Utveckling 30. 1877. Riccia coriandrina Spreng., Anleitung zur Kenntnis der Gewächse 3: 320. 1804. Corsinia marchantioides Raddi, Opusc. Sci. 2: 354. 1818, nom. illeg. Lectoype (Schuster, 1992b). Italy, near Florence, Micheli s.n. (n.v.).

Figs. 1: B, 2: A-C, 3: F, 84, 85, 104: D-E

Corsinia coriandrina fo. gymnocarpa Massal., Bull. Soc. Bot. Ital. 1917, nom. nud. Material. Argentina. Buenos Aires: La Plata, Spegazzini s.n. (n.v.) Thallus 3-6 mm wide, forked, pale or whitish green, not tinged with purple; when fresh, the plants emanate an odour of coriander. Epidermal cells thin-walled; epidermal pores with ring of 5-7 cells, radial walls thin. Air chambers in 1(-2) layers with chlorophyllose filaments. Basal tissue with oilcells, without mucilage cavities. Rhizoids smooth and pegged. Ventral scales hyaline, in several illdefined rows, with oil-cells and marginal papillae and with filiform appendage, not constricted basally. Monoecious or dioecious. Antheridia in dorsal groove along thallus midline bordered by cilia. Archegonia 1-10 in dorsal cavities with slightly proliferating edges, forming a toothed-ciliate crest. Calyptrae several-layered after fertilization, warty or spinose. Involucre an irregular, pluristratose, posterior scale. Sporophytes 1-3 per involucre. Capsule wall without annular thickenings. Spores 300350 per capsule, 90-150 µm diameter, light to dark red-brown, distal face with plates or flat protuberances, proximal face faintly tuberculate, trilete scar distinct. Elaters lacking, but sterile cells mixed with spores. Gametophytic chromosome number n = 8 or 16. Distribution and ecology. Corsinia coriandrina is distributed in warm-temperate areas with Mediterranean-type climates, in the Mediterranean-Atlantic area, Macaronesia, SW Asia, and the southern U. S. A. (Texas). Records from Afghanistan, China and Japan belong to other taxa. In the Neotropics, the species is rare and known only from the northern and southern borders of the area. It has been recorded from Mexico (Hicks, 1993; Düll, 1999; Mexican specimens examined belong to Exormotheca pustulosa!), Brazil (Paraná (Vianna, 1976), Rio Grande do Sul (Vianna, 1976)), Chile (Valparaiso (Mahu, 1994)), and Argentina (Buenos Aires, Corrientes, Misiones (Hässel, 1963)). The species is drought-tolerant and grows on compact, periodically moist sand or clay of low pH,

Bischler et al. 96 in rocky habitats or on bare soil, exposed or shaded by rather open vegetation, at low elevations. Specimens examined. ARGENTINA. BUENOS AIRES: Adrogaré, Pujals 10838 (BA); 25 km W Tandil, Volk 6/0631 (BA). CORRIENTES: Mercedes, 25 km de Paso de los Libres, ruta 23, Matteri 26966 (BA). The thallus of Corsinia coriandrina resembles that of Riccia but differs by its scales (in illdefined rows) and the structure of its epidermal pores, and by the characteristics of the male and female receptacles. Spore ornamentation also separates Corsinia from all Riccia species. The species comprises morphologically indistinct haploid dioecious and polyploid monoecious populations with allopatric distributions in the Mediterranean-Atlantic area. The single New World population analysed was polyploid and differed genetically from Old World polyploids (BoisselierDubayle & Bischler, 1998). Neotropical populations of the species have not yet been investigated genetically.

5. CRONISIA Berk., Introd. Crypt. Bot. 434. 1857. Type. Cronisia paradoxa Berk. (= Cronisia weddellii (Mont.) Grolle). Anagram of Corsinia. Boschia Mont., Ann. Sci. Nat., Bot. sér. 4, 5: 351. 1856, nom. illeg. Funicularia Trevis., Mem. Reale Ist. Lombardo Sci. Ser. 3, Cl. Sci. Mat. 4: 443. 1877, nom. illeg. Type. Boschia weddellii Mont. Myriorrhynchus Lindb., Acta Soc. Fauna Fl. Fenn. 2, 5: 7. 1884. Type. M. fimbriatus (Nees) Lindb. (= Riccia fimbriata Nees). Thallus light, whitish or yellowish green, often tinged with purple or violet. Epidermal cells thinwalled; epidermal pores with ring of 5-10 cells, without hyaline ring, radial walls thin, sometimes strongly elevated. Air chambers in one layer, with chlorophyllose filaments. Basal tissue often with mucilage cavities. Ventral scales in 2 rows, with oil-cells, with or without papillae; appendages filiform or subdivided into numerous uniseriate filaments, not constricted basally. Monoecious. Antheridia in dorsal groove along thallus midline. Archegonia in dorsal cavity. Calyptrae 1-4-layered, not warty. Involucre unistratose, involute around archegonia. Sporophytes 1-2 per involucre. Capsule wall with or without annular thickenings. Spores approx. 100-200 per capsule, 60-108 µm diameter, distal face with flat protuberances, proximal face faintly tuberculate, trilete scar distinct. Elaters short, not more than twice as long as wide, with single helical band, or sterile cells intermixed with spores. Gametophytic chromosome number n = 9. A neotropical genus of two species occurring in tropical, winter-dry savannas of Central and South America.

KEY TO THE SPECIES OF CRONISIA

Bischler et al. 97

1. Epidermal pores usually strongly elevated above epidermis. Mucilage cavities in ventral thallus tissue numerous. Ventral scales projecting beyond thallus margins, appendage subdivided into uniseriate and furcate filaments, inserted beyond apex of scale body. Involucres without pigmented cells. Capsule walls thin, annular thickenings lacking. Sterile cells mixed with spores short-elongate, without helical thickenings. ............................................................................... 1. Cronisia fimbriata 1. Epidermal pores not strongly elevated above epidermis. Mucilage cavities in ventral thallus tissue few. Ventral scales not extending beyond thallus margins, with 1-6 filiform appendages, inserted at apex of scale body. Involucres dotted with dark purplish cells. Capsule wall with annular thickenings. Elaters with helical band..................................................................................... 2. Cronisia weddellii

1. Cronisia fimbriata (Nees) Whittem. & Bischl., Cryptogamie, Bryol. 22: 4. 2001. Riccia fimbriata Nees, in Martius, Fl. Brasil. 1: 301. 1833. Myriorrhynchus fimbriatus (Nees) Lindb., Acta Soc. Fauna Fl. Fenn. 2, 5: 7. 1884. Type. Brazil. Minas Gerais: Sierra de Piedade, Martius s.n. (isotypes, G, BM).

Figs. 86, 87, 104: F-G

Cronisia mexicana Hicks, Tropical Bryology 7: 1. 1993. Type. Mexico. Tlaxcala: next to waterfall on Rio Zahuapan near Amaxoc, 2300 m, Sharp 419 (type, TENN; isotype, MEXU). Thallus 1.5-2.5 mm wide, whitish or yellow-green, ventral side tinged with dark red, margins hyaline. Epidermal pores with ring of 5-10 cells, radial walls thin, strongly or moderately elevated above epidermis. Air chambers with well developed chlorophyllose filaments. Basal tissue with numerous mucilage cavities. Ventral scales extending beyond thallus margins, hyaline apically, bordered by small cells, with oil-cells, red-orange to dark red basally; appendage inserted beyond apex, subdivided into uniseriate, furcate filaments. Monoecious. Antheridia in several rows, in dorsal groove. Involucre unistratose, plicate apically, without idioblasts, with numerous yellowish oil cells. Capsule wall not thickened. Spores 100-200 per capsule, 92-108 µm diameter, distal face with rounded protuberances, proximal face faintly tuberculate, trilete scar distinct. Elaters lacking, but sterile cells mixed with spores. Gametophytic chromosome number n = 9. Distribution and ecology. The species is rare, known from Mexico (Chihuahua, Tlaxcala), Brazil (Bahía, Cearà (Schäfer-Verwimp & Gincotto, 1993), Mato Grosso do Sul (Schäfer-Verwimp & Gincotto, 1993), Minas Gerais, Paraíba (Schäfer-Verwimp & Gincotto, 1993), Pernambuco, Piauí), and Argentina (Salta (Massalongo, 1906)). The species grows on rocky soil, sandstone, lava or granite, or on calcareous soil, exposed, in grassland, caatinga, in semi-arid zones, from 300 m in Brazil to 2400 m in Mexico. Selected specimens examined. MEXICO. CHIHUAHUA: on dirt road, 3.8 mi. E of S. J.

Bischler et al. 98 Bachiniva, on hwy. to Cuauhtemoc, 2000 m, Whittemore et al. 3459, 3460 (Hb. Whittemore). TLAXCALA: type of C. mexicana. BRAZIL. BAHÍA: Jeremoaba, Vital 8153 (PC); Bom Jesus de Lapa, Vital 1651 (FLAS). MINAS GERAIS: type. PERNAMBUCO: Serra Talhaa, Porto 8095 (PC); Fazenda Nova near Caruaru, Hürlimann H 503 (JE). PIAUÍ: Mun. Fronteira, Vital 2924 (PC); Mun. Picos, Vital 2929 (PC). The type material of Cronisia fimbriata is sterile. Because of the epidermal pores, which are strongly elevated above the epidermis, the species has formerly been included in the genus Exormotheca. However, the recently discovered gynoecia clearly show that the species belongs to Cronisia (Bischler & Whittemore, 2001). The absence of capsule wall thickenings, the elaters without helical bands, and the peculiar structure and insertion of the scale appendage separate the species from C. weddellii.

2. Cronisia weddellii (Mont.) Grolle, J. Bryol. 9: 532. 1977. Boschia weddellii Mont., Ann. Sci. Nat., Bot. sér. 4, 5: 352. 1856; Bull. Soc. Bot. France 3: 576. 1856. Funicularia weddellii (Mont.) Trevis., Mem. Reale Ist. Lombardo Sci. Ser. 3, Cl. Sci. Mat. 4: 443. 1877. Type. Brazil. Goiás: Weddell s.n. (BM, PC).

Figs. 88, 89, 104: H-I

Riccia paradoxa Hook. & Wilson, London J. Bot. 3: 166. 1844, nom. illeg. Cronisia paradoxa Berkeley, Introd. Crypt. Bot. 434. 1857. Type. Brazil. Piauí: near Oeiras, Gardner s.n. (BM, NY). Funicularia bischleriana Jovet-Ast, Rev. Bryol. Lichénol. 32: 201. 1963. Type. Colombia. Meta: Río Guayabero, rapids of the Macarena, 350 m, Bischler 1671 (PC). Thallus 2-3.5 mm wide, often tinged with purple or violet. Epidermal pores with ring of 5-7 cells, radial walls thin. Air chambers with well developed or rudimentary chlorophyllose filaments. Basal tissue with some mucilage cavities. Ventral scales with oil-cells and marginal papillae, and with 1-6 filiform appendages, one usually longer than the others. Antheridia in 2-3 rows. Archegonia 2-4 per cavity. Involucre unistratose, with dark purplish idioblasts. Capsule walls with annular thickenings. Spores approx. 120 per capsule, 60-97 µm diameter, distal face with flat protuberances, proximal face faintly tuberculate, trilete scar distinct. Elaters very short, with single helical band. Gametophytic chromosome number n = 9. Distribution and ecology. This rare neotropical species has been recorded from Mexico (Veracruz (Schuster, 1992b)), Southern Dutch Antilles (Curaçao (van Slageren, 1979)), Colombia (Meta), Brazil (Alagoas (Vital, 1974a), Bahía, Cearà (Vital 1974a), Espíritu Santo, Goiás, Mato Grosso (Vital, 1974a), Piauí), from areas with a pronounced seasonal climate. Cronisia weddellii is drought-tolerant and grows on periodically moist, exposed or shaded, sandy soil, on moist banks, along river borders, in caatinga vegetation and in deciduous, seasonal forest, from sea level to 800 m. Specimens examined. SOUTHERN DUTCH ANTILLES. CURACAO: Christoffelsberg area,

Bischler et al. 99 rooi Beru on NNE side, van Slageren & Stoffers 8319 (U). COLOMBIA. META: type of Funicularia bischleriana. BRAZIL. “Brasilia tropica, ” Burchell8 8903 (BM). BAHÍA: Jaguarari, Vital 8099 (JE). ESPÍRITU SANTO: Linhares, Rio Doce, Vital 1877 (FLAS, JE). GOIÁS: type. PIAUÍ: Oeiras, Gardner 6, 130 (BM, NY).

CYATHODIACEAE

by Noris Salazar Allen

4. CYATHODIACEAE (Grolle) Crandall-Stotler & Stotler, in J. Shaw & B. Goffinet, Bryophyte Biology 55, 63. 2000. Targioniaceae subfam. Cyathodioideae Grolle, J. Bryol. 7: 208. 1972. Cyathodiaceae Müll. Frib. ex Rabenhorst, Krypt.-Fl. Deutschl. ed. 2, 6 (Ergänzungsb.): 182. 1940, nom. inval. Type. Cyathodium Kunze. The family is monogeneric.

6. CYATHODIUM Kunze, in Lehmann, Nov. Stirp. Pugillus 6: 17. 1834. Type. C. cavernarum Kunze. Name from cyathus = cup. Thallus 1-4(-16) mm wide, not tinged with purple. Branching dichotomous, ventral and apical; thalli forming often hemi-rosettes. Epidermis persistent. Epidermal pores simple, with 1 ring of 5-21 cells, without hyaline inner ring, wide open, radial walls thin. Air chambers in one layer, empty. Basal tissue reduced, in 0-2 layers, with or without oil-cells. Rhizoids mostly smooth, some tuberculate, sinuous and with a spiral twisting. Ventral scales in 2 rows, often restricted to apical part of thallus, of 2-10 cells, with or without oil cells, with marginal papillae, sometimes with small, filiform appendage not constricted basally. Asexual reproduction in some species by “gemmae” (fragmenting thallus segments) from thallus margins or by ventral tubers. Monoecious or dioecious. Antheridia embedded in small ventral or apical branches with large scales. Archegonia 1-30, in archegonial cavity located ventrally beyond thallus apex, . Archegonia terminal in bivalved involucre beyond thallus apex, sometimes bounded by thallus scales larger than those of sterile parts of thallus Sporophytes with foot and hardly elongating seta. Calyptrae 2-4-layered after fertilization. Sporophytes 1-4 per involucre. Capsules opening by a lid and 6-8 irregular valves to 1/3-1/2 of capsule length, wall with annular thickenings in upper part only. Spore : elater ratio more than 4 to 1.

Bischler et al. 100 Spores 25->100 per capsule, 25-85 µm diameter, proximal and distal faces tuberculate or short- to long-spinose, or with more or less complete, tuberculate areoles and ridges, trilete scar hardly distinct. Elaters 4-100 per capsule, with 2-4 helical bands. Gametophytic chromosome number n = 9. A genus of about 12 mesophytic-hygrophytic species able to grow under weak light intensities, found on rock, soil or bark in forests, caves, along river banks, on rocks around waterfalls, on cement floors, stairs and flower pots in humid areas, sometimes weedy in arable fields, distributed in the tropical belt in southern Japan, southern Africa, South America, Central America and the Caribbean at low elevations. The species are not drought-tolerant but are able to perennate with ventral tubers. Srivastava & Dixit (1996) subdivided Cyathodium into two subgenera, subgen. Metacyathodium with ventral tissue several layers rhick along de midline of the thallus, and subgen. Cyathodium with only a unistratose ventral epidermis below the air chambers.

KEY TO THE NEOTROPICAL SPECIES OF CYATHODIUM

1. Thallus with midrib, or multistratose area at least under the antheridial receptacles ............................... 2 2. Dorsal pores with innermost ring of cells projecting into air chambers. Tuberculate rhizoids with few tubercles. Dioecious. Involucre sac-like, opening by two lips apically, not convoluted. Spores irregularly lamellate................................................................................................ 5. Cyathodium steerei 2. Dorsal pores with innermost ring of cells not projecting into air chambers. Tuberculate rhizoids with numerous tubercles. Monoecious. Involucre not bilabiate, convoluted. Spores verrucosetuberculate....................................................................................................3. Cyathodium foetidissimum 1. Thallus without midrib.............................................................................................................................. 3 3. Thallus elongate, dichotomously branched, each thallus cell with chloroplasts and 4-8 oil bodies. Dioecicous. Spores lamellate-reticulate......................................................... 4. Cyathodium spruceanum 3. Thallus in rosettes, oil bodies 1(-2) in specialized cells devoid of chloroplasts. Monoecious. Spores baculate-spinose or echinate ........................................................................................................ 4 4. Dorsal pores prominent, in two rows along thallus length. Sporophytes with operculum of 4(8) outer cells. Spores baculate-spinose, dark brown................................. 2. Cyathodium cavernarum 4. Dorsal pores not prominent, 1-2 on mid-thallus, at base of apical lobes. Sporophytes without operculum. Spores echinate, green to light brown................................1. Cyathodium bischlerianum

Bischler et al. 101

KEY TO STERILE PLANTS

1. Thallus in compact rosettes.............................................................................................................2 2. Thallus border with marginal cells with outer walls slightly to strongly concave. Dorsal pores numerous and large.............................................................................. 2. Cyathodium cavernarum 2. Thallus border with marginal cells with outer walls strongly convex. Dorsal pores 1-2............. ..........................................................................................................1. Cyathodium bischlerianum 1. Thallus elongate, dichotomously branched.....................................................................................3 3. Midrib (multistratose area) absent. Thallus cells with oil bodies and chloroplasts ..................... .............................................................................................................4. Cyathodium spruceanum 3. Midrib (multistratose area) present, oil bodies in specialized cells (idioblasts) ........................4 4. Tuberculate rhizoids slightly tuberculate. Midrib present at least under antheridial receptacles and in young thalli. Thalli thin, delicate ...............................5. Cyathodium steerei 4. Tuberculate rhizoids strongly tuberculate. Midrib prominent in young and adult thalli. Thalli fleshy ........................................................................................... 3. Cyathodium foetidissimum

1. Cyathodium bischlerianum Salazar Allen, Bryologist 104: 141. 2001. Type. Panama. Parque Nacional Soberanía, Río Pedro Miguel, Camino de Cruces, Salazar Allen, Chung & Rivera 14390. (PMA). Paratypes. Parque Nacional Soberanía, Sendero El Charco, Rodríguez 192, 216 (PMA); Río Pedro Miguel, Camino de Cruces, Salazar Allen et al. 14386, 14393, 16612 p.p. (PMA). Figs. 90, 91: A-D, 97: A-B Thallus metallic green to yellowish-green, delicate, small, 3.5-6.5 mm long, 1.8-2.7 mm wide, dichotomously branched, in compact or discrete rosettes, fan-shaped, thallus broadly rounded at apex, forming two ear-shaped lobes; marginal cells with outer walls strongly convex, oil cells triangular, smaller than adjacent border cells. Dorsal epidermis delicate, chlorophyllose, chloroplasts larger than those of ventral cells, cells 22-54(-80) µm long, (19-)22-52(-60) µm wide, walls thin, lacking trigones, oil bodies light brown, 1(-2), in cells lacking chloroplasts. Epidermal pores 1-2, simple, wide open, surrounded by 1-3(-4) concentric rows of 5-12 quadrate to botuliform cells. Ventral epidermis chlorophyllose, cells polygonal, larger than those of dorsal or adaxial epidermal layer, (37-) 40-123(-144) µm long, 22-55(-67) µm wide. Midrib absent. Rhizoids long and colourless, of two

Bischler et al. 102 types, large and straight-walled and narrower and sinuous. Ventral scales simple, without appendages, near thallus apex, on ventral walls of involucre and around antheridial receptacles, filamentous to ± elongate triangular, 3-8 cells long, with apical or lateral idioblasts containing 1(-3) oil bodies. Monoecious. Antheridial receptacles lateral, cushion-shaped to oblong triangular, 165 µm in longest diameter, with basal scales, ostioles raised above surface. Involucres divided into two lips, with 1-2 archegonia, marginal cells elongate in 2(-3) rows, the outermost row with thickened inner tangential walls, brown-red in colour, mid-cells polygonal, larger, thin-walled, hyaline towards base. Sporophytes with a seta of 9 uniseriate, narrow cells. Capsules ovoid, 1(-2) per involucre, 0.35-0.52 mm long, 0.32-0.46 mm wide, orange brown when mature, without operculum; capsule wall one cell thick, cells in upper 1/2-1/3 with transversal, bar-shaped thickenings, dehiscent by 4-6(-7) irregular valves, cells of lower 2/3-1/2 without thickening bands and with large oil bodies, non-dehiscent. Spores green to light brown, (37-)40-55(-60) µm in longest diameter, echinate, oval-triangular. Elaters attached to base of capsule, varying in number with size of capsule from 2-7, 410-585 µm long, 12-15 µm diameter, with 4 narrow helical bands, attenuated on distal end, widened and funnelshaped proximally. Distribution and ecology. The species has been collected in the area of the Canal (sea level to 50 m) and in the Province of Coclé (580 m) in Panama. It is probably more widely distributed in Central America and the Neotropics but due to its small size may have been overlooked. The species usually grows in mixed collections together with C. cavernarum and C. spruceanum in very moist, shaded areas, on rocks near or in creeks, along eroding river banks, subject to partial submergence during high water levels, and on tree bark, e.g., of Ficus sp. Selected specimens examined. PANAMA. PANAMA: Parque Nacional Soberanía, Río Pedro Miguel, Camino de Cruces, Salazar Allen et al. 14386 p.p., 14393, 14395 p.p, 16599, 16600, 16612 p.p., 16613 p.p., 16643 (PMA), Rodríguez 209, 211, 216, 220 (PMA). COCLÉ: El Valle de Antón, Río Las Mozas, Salazar Allen et al. 14447 (PMA). Cyathodium bischlerianum differs from C. cavernarum in having slender, more elongate and more delicate thalli with strongly crenulate borders due to pronounced conical projection of marginal cells. Pores on the dorsal epidermis are few (usually a single one at base of involucre) and divisions of air chambers are not readily observed. The apical lobes are large, ear-shaped and abruptly constricted towards base at insertion of antheridial receptacles. When mature, the lips of the involucre are below or at the same level of the thallus rim and the sporophytes project from the upper and ventral side of the involucre. The echinate spore ornamentation, consisting of spines with a bulbous base, is unique among neotropical species of Cyathodium and otherwise only known for the Asiatic C. aureonitens. The dioecious condition, hairy involucre, presence of an operculum, ornamentation of upper cells of capsule, and spore colour readily separate the latter species from C. bischlerianum.

Bischler et al. 103 2. Cyathodium cavernarum Kunze, in Lehmann, Nov. Stirp. Pugillus 6: 17. 1834. Type. Cuba. Without locality, 1824, Poeppig s.n. (holotype, W; isotype, S).

Figs. 93, 94, 97: C-D

Cyathodium africanum Mitt., J. Proc. Linn. Soc. London 22: 327. 1887. Type. Tanzania, Usagara Mountains, J. Hannington s.n. (holotype, K). Cyathodium mexicanum Steph., Rev. Bryol. 36: 139. 1909. Type. Mexico. Jalisco: barranca Guadalajara, 26 Sep.1908, Pringle 10677 (holotype, G, n.v.) Cyathodium barodae Chavan, Bryologist 11: 57. 1937. Type. India, Baroda State, Baroda, 1934-1935, Chavan s.n. (n.v.). Thallus green to yellowish-green, delicate, small, dichotomously branched, in compact or discrete rosettes, fan-shaped, (2.2-)3.8-9.0 mm long, 0.7-2.2 mm wide; marginal cells chlorophyllose, rectangular to quadrate with outer walls slightly to strongly concave, oil cells triangular, devoid of chloroplasts. Dorsal epidermis delicate, chlorophyllose, cells (33-)45-60(-67) µm long, (20-)30-52(60) µm wide, walls thin, lacking trigones; oil bodies light brown, 1(-2), in idioblasts lacking chloroplasts, 15-27 m long, 15-24 µm wide. Epidermal pores simple, smaller towards apex, wide open, slightly raised above epidermal cells, surrounded by 3-4(-5) concentric rows of 4-6(-10) elongate cells, the uppermost sometimes with sinuous internally abutting cell walls. Ventral epidermis chlorophyllose, chloroplasts smaller than those of dorsal epidermis, cells polygonal, larger than those of dorsal layer, 90-187 µm long, (30-)45-67 µm wide. Midrib absent. Rhizoids abundant, long and colourless, of two types, large and ± straight-walled and narrower and sinuous, not tuberculate. Ventral scales near thallus apex and on ventral walls of involucre, simple, without appendages, filamentous to ± triangular deltoid, 3-6 cells long, with lateral or terminal cells containing an oil body. Monoecious. Antheridial receptacles lateral, sometimes occurring in close proximity of female receptacles, cushion-shaped to oblong elliptical, 150-210 µm long, 142-230 µm wide. Involucres sacshaped, divided into two lips, marginal cells elongate, reddish-brown, in 1-2 rows with inner tangential walls thickened, mid-cells polygonal, thin-walled, brown on distal portion, hyaline below; archegonia 2-5 per involucre, maturation asynchronous. Sporophytes with a delicate seta of 6 uniseriate, narrow cells. Capsules ovoid, 1(-3) per involucre, (0.2-)0.35-0.6 mm long, dark brown to black when mature, one cell thick; operculum of 2 tiers of cells, outermost 4(-8) cells, the innermost 8-12 cells; cells in upper 1/2-1/3 of capsule, below the operculum with annular to spiral thickenings, splitting into 4-6 irregular valves when drying, cells of lower half without thickening bands, nondehiscent, with chloroplasts and large oil bodies. Spores dark brown, (32-)35-54(-60) µm diameter, densely to sparsely baculate-spinose, with rugose surface. Elaters (2-)4-16 per capsule, 390-585 µm long, 15-16 µm diameter, varying in number with size of capsule, trispiral at middle, bispiral on the attenuated ends. Illustrations. Stephani (1985, as C. mexicanum); Montagne (1842, Fig. 4); Lang (1905, Fig. 2548); Chavan (1937a, Figs. 1-50; 1937b, Figs. 1-83, as C. barodae); Schiffner (1938, p. 133, Fig. 1);

Bischler et al. 104 Vital (1974b, Fig. 2, as Cyathodium africanum); Yano & Costa (1992, Fig. 2, as C. africanum); Srivastava & Dixit (1996, Figs. 8: 1-21, 19: 1-4). Distribution and ecology. Cyathodium cavernarum has been collected in Mexico, Central America (Costa Rica, Panama), the Caribbean (Cuba), and South America (Colombia and Brazil). It is the most widely distributed species of the genus and probably occurs throughout the Neotropics. In the Old World tropics the species has been reported from Africa, India, Western and Eastern Himalayas, Burma, and Indonesia (Java, Srivastava & Dixit, 1996). It grows in soil, on rocks, on eroded banks of rivers or creeks, along roadsides or trails, on bark of trees (palms and other angiosperms), on shaded sites, caves, on cement gutters and walls along roadsides, on cement flower pots. It grows on calcareous substrata but it can also be found on slightly acidic substrates where underlying bedrock is calcareous. In exposed habitats the species usually grows associated with members of the Pottiaceae; in ravines and river banks with species of Philonotis, Fissidens, Notothylas, Racopilum tomentosum (Hedw.) Brid., Cyathodium spruceanum and Lejeuneaceae. Cyanobacteria are also closely associated with this and other species of Cyathodium, forming spherical colonies attached to various parts of the thallus. Selected specimens examined. MEXICO. MÉXICO: Barranca de Malinaltenango, near the colonial bridge at valley bottom, Pócs 9552C (MEXU). NAYARIT: 50 km east of Ruiz, 9.8 km east of San Pedro Ixcatán on road to Mesa de Nayar, Whittemore et al. 3244 (MEXU). VERACRUZ: Hwy. 180, 3.3 mi. north-west of turnout to San Andrés de Tuxtla, Whittemore et al. 2184 (MEXU); tropical montane forest below Fortin de las Flores, Düll 2/106 (MEXU). COSTA RICA. San José, Zoológico Nacional Simón Bolívar, Salazar Allen et al.16082-16087, 16184-16187 (PMA, SJ); Alajuela, Universidad Adventista de Centroamérica, Salazar Allen et al. 16090 (PMA, SJ); Alajuela, road to Carrara, Salazar Allen et al. 16099-16104 (PMA, SJ); San Miguel de Sarapiqui, Cerro Congo, Salazar Allen et al. 16162 (PMA, SJ); Atenas, Salazar Allen et al. 16106, 16108-160109 (PMA, SJ), 16111-16123 (PMA, SJ); San Mateo, Salazar Allen et al. 16124-16129, 16132-16136 (PMA, SJ); San Mateo, Quebrada Fierro, Salazar Allen et al. 16137-16141 (PMA, SJ). PANAMÁ. COCLÉ: El Valle de Antón, Río Las Mozas, Salazar Allen et al. 14457, 14520, 16555, 16556, 16558, 16566, 16571, 16572, 16579, 16581 p.p. (PMA), Rodríguez 169, 170 (PMA); Quebrada El Guayabo, Salazar Allen et al. 16592, 16595 (PMA). COLÓN: Aduana de Portobelo, Salazar Allen & Santamaría s.n. (PMA). LOS SANTOS: Parque Nacional Cerro Hoya, E. of Río Progreso, Rivera s.n. (PMA); PANAMÁ: Cuevas de Chilibre, Salazar Allen et al. 16660, 16661, 16665 (PMA); Parque National Soberanía, Sendero El Charco, Salazar Allen et al. 14311, 14825, 14832, 14856, 14859, 14865 (PMA), La Cascada, Rodríguez 226, 234, 235 (PMA), Río Pedro Miguel, Camino de Cruces, Rodríguez 179, 220, 221 (PMA), Salazar Allen et al. 14383, 14387, 14395, 15236, 16599, 16606, 16614, 16619, 16625, 16626 (PMA); Parque Nacional Altos de Campana, Salazar Allen & H. Korpelainen 15692, 15694 (PMA).

Bischler et al. 105 CUBA. Unknown locality, Wright 28501 (HAC); Santiago de Cuba, Alto Villalón, Caney, Pócs & Caluff 9164/V (HAC). BRAZIL. CEARÁ: Ubajara, Vital 1338 (PC, SP). GOIÁS: Serra de Lobeira, Itaberai, Vital 2 (PC, SP). MATO GROSSO: Barra do Garças, Vital 1350 (PC, SP). MINAS GERAIS: Lagoa Santa, Vital 1207 (PC, SP). Populations of Cyathodium cavernarum are quite variable in size of thallus, shape and size of pores, involucral and sporophyte position, size of sporophytes, number and size of spores and number of elaters per capsule (Vital, 1974b). Thalli may be short and tumid to elongate and delicate. Plants growing on cement gutters along roadsides in Costa Rica are rather large in size, with thalli in tumid rosettes, deep green and with large pores. Populations on river banks tend to have more elongate, delicate thalli and less pronounced pores. In both variants the two rows of air chambers are distinctive. Involucral lips may be slightly to widely divergent, projecting beyond the rim of lobes, or located slightly below them, or at the same level. Sporophytes can be found deeply within the involucre or protruding at apex. Very small sporophytes (0.2 mm long) have been observed in lateral branches of thalli carrying large sporophytes (0.6 mm long) at apex. There may only be two or three elaters in the small capsules. Cyathodium cavernarum closely resembles C. bischlerianum in its bright metallic-green colour. It grows intermixed and usually above the latter species. The more robust, fan-shaped thallus with larger pores and air chambers in two rows, and the presence of an operculum, clearly distinguish C. cavernarum from C. bischlerianum. The thickenings of the upper capsule cells and spore ornamentation are also very different between the two species (see under C. bischlerianum).

3. Cyathodium foetidissimum Schiffn., Expos. plant. Itineris Indici I: 2. 1898. Type. Indonesia. West Java: “In monte Megamendong; in silva primaeva apud locumTelaga Warna ad terram inter arb. radices. Regio pluvialis, “ alt. ca. 1400 m, 5 Jan 1894, V. Schiffner 8 (holotype, PC). Figs. 94-95, 97: E Thallus light green, delicate, elongate, dichotomously branched, 4-17 mm long, 1-6 mm wide; marginal cells chlorophyllose, quadrate to short-rectangular, oil cells triangular. Dorsal epidermis delicate, without trigones, chlorophyllose, cells (15-)20-37.5 µm long, (15-)18-30 µm wide, oil cells smaller, 15 µm long, 19-23 µm wide, with a single, hyaline to greyish oil body. Epidermal pores simple, smaller towards apex, surrounded by 3-4 irregular rows of elongate cells, these with minute chloroplasts. Ventral epidermis chlorophyllose, chloroplasts smaller than those of dorsal epidermis, cells polygonal, larger than those of upper epidermis 75-160 µm long, (24-)33-68(-75) µm wide. Midrib (= central multistratose area) present. Rhizoids abundant, long and colourless, of two types, narrow and strongly tuberculate, or wide and smooth, basal rhizoid area brownish. Ventral scales

Bischler et al. 106 large, multicellular, long rectangular to irregularly deltoid, 4-7 cells wide, 7-8(-12) cells long, with oil cells in idioblasts. Monoecious. Antheridial receptacles sessile in apical sinus, cushion-shaped, oval to oblong, flanked on both sides by archegonia. Archegonia on both sides of male receptacle, enclosed by sinuous laminar involucre. Sporophytes with a short seta, 0.6-0.8 mm in diameter, dark brown, operculum of 12 cells, projecting into the capsule cavity; cells in upper 1/2 to 1/3 of capsule, below operculum, with tangential bar thickenings, lowermost cells smooth. Spores dark brown, subtriangular in shape, verrucose-tuberculate 31-42 µm in longest diameter (not including verrucae). Elaters 390600 µm long, 12-15 µm wide, with three thickened bands, attenuated at apex. Illustrations. Lang (1905, Pl. XXII: 25-48); Goebel (1915, Fig. 645; 1930, Fig. 827); Schiffner (1939, Figs. 37-40); Jones (1952, Fig. 1: e, f); Srivastava & Dixit (1996, Figs. XI: 1-12, XXI: 1-4). Distribution and Habitat. Cyathodium foetidissimum is widespread in the Old World tropics and had not previously been reported from the Neotropics. This species has recently been collected in Costa Rica and Ecuador. In Costa Rica the species was found growing at 1150 m elevation in small patches in depressions on a sandy limestone rock wall hidden by vascular vegetation, in very shady conditions, next to Dumortiera hirsuta (Sw.) Nees and Marchantia sp., and mixed with Fissidens flaccidus Mitt., Taxiphyllum taxirameum (Mitt.) Fleisch., and a species of Lejeuneaceae. A collection from Ecuador (150 m elevation), identified by Dr. Srivastava as Cyathodium cf. foetidissimum, has strongly tuberculate rhizoids typical of C. foetidissimum but the uppermost thallus cells are larger than in the material from Costa Rica. In addition, the specimen from Ecuador has vegetative tubers, which are unknown in C. foetidissimum. Probably this Asiatic species is more widely distributed in the Neotropics; the scarcity of collections may be due to the small size and delicacy of the thallus. Specimens examined. COSTA RICA. CARTAGO: Cantón de Paraíso, Valle del Río Grande de Orosí, “en pared rocosa de arenisca caliza antes de entrar al puente colgante, ” Salazar Allen, Lepiz & De Gracia 17047 (PMA), 17048, 17049 (PMA, SJ). ECUADOR. LOS RÍOS: Río Palenque Forest Reserve, on bare soil under oil palm in plantation, Gradstein, Mues & Frahm 6958 (U). The two collections examined are the only records of this species from tropical America. In general aspect, sterile thalli of this species closely resemble small or young thalli of Cyathodium spruceanum. The presence of a multistratose central area (“midrib”), oil bodies in specialized idioblasts, strongly tuberculate rhizoids and the inner row of cells surrounding pores not abutting into the opening, separate C. foetidissimum from C. spruceanum. Fertile plants are distinctly different in the involucral morphology and sexual condition. Cyathodium steerei is dioecious, with a saccate involucre opening by two apical lips, whereas C. foetidissimum is monoecious and has involucres that

Bischler et al. 107 are laminar and occur on both sides of the apically located male receptacle. Other species having oil bodies in idioblasts are C. cavernarum and C. bischlerianum but the latter two species grow in rosettes or produce very minute thalli without midrib. Also, when fertile, these two species produce their male receptacles laterally, unlike C. foetidissimum which produces them apically.

4. Cyathodium spruceanum Prosk., Bryologist 54: 243. 1951. Type. Peru. Huanuco: Tingo María, Cueva de los pavos, on soil under boulder, ca. 750 m, 3 Sep 1949, Proskauer s.n. (holotype, UC 924947).

Figs. 91: E-J, 96, 97: F-H, 99

Thallus light green to greenish-yellow, elongate, dichotomously branched, in large to discrete patches, 13-20 mm long, 1.5-8.0 mm wide, variable in size; marginal cells hyaline or with few chloroplasts and oil bodies, in 1-2 rows, turning brown with age. Dorsal epidermis delicate, cells at midthallus (25-)34-60(-90) µm long, (18-)30-96 µm wide with chloroplasts and 4-6(-8) translucent to light brown oil bodies, walls thin, lacking trigones, markedly distended on their inner (air chamber) side. Epidermal pores simple, in two rows, slightly raised above epidermal cells, widely opened, surrounded by 4-5 rows of narrow, botuliform cells, the uppermost abutting into pore. Ventral epidermis chlorophyllose, chloroplasts smaller than those of dorsal epidermis, cells hexagonal to polygonal, at midthallus (60-)75-127(-180) µm long, (25-)37-60(-70) µm wide. Midrib absent. Rhizoids brown at insertion, abundant, long, of two types, smooth and tuberculate, lightly sinuous with spiral twisting. Ventral scales variable in appearance and number of cells, near thallus apex, on ventral walls and inside of involucre, larger on ventral walls of involucre, without appendages, hyaline to slightly brown, elongate triangular, 5-10 cells long, with lateral and terminal cells containing oil bodies. Dioecious. Antheridial receptacles apical, sessile, oval, elliptical, transversely elongate, lobate to horseshoe shaped, 1.3-3.0 mm long, 0.4-1.5 mm wide, surrounded by a lamina of elongate cells that projects from thallus lobes, scale wider distally; receptacles turning brown when mature with more than 100 antheridial chambers flanked by 4-5 rows of elevated cells surrounding pore. Female plants similar to male ones. Involucre large, with 5-30 archegonia and sometimes with 1(-2) inner scales, brown and truncate at apex; larger scales and rhizoids on its ventral surface, opening by two smooth to slightly dentate, ± recurved lips on its distal surface, the lips below thallus rim or the upper portion of involucre projecting outwardly beyond rim and funnel-shaped when containing 10-30 archegonia; outermost cells of involucral lips reddish brown, oval, square to rectangular with thickened inner tangential walls. Sporophytes 1(-2) per involucre, seta delicate 2(-3) seriate, 6 cells long. Capsules dark brown to black when mature, operculum of 6-12(-13) cells, slightly projecting apically, upper cells of capsule with irregular thickenings on their radial and tangential walls. Spores dark brown, (30-)37-42(-55) µm diameter, lamellate-reticulate, lamellae projecting from surface of spore and vertically striated, 2-5 µm high, 0.7-2.8 µm wide, distal face with reticulum forming closed areoles, opened on proximal face, with a transverse cingulum dividing the

Bischler et al. 108 spore in two asymmetrical halves; cingulum with a central, linear row of gemmae and minute perforations. Elaters 20-80(-100) per capsule, 600-780 µm long, 16-19(-24) µm wide, trispiral at middle, bispiral at attenuated ends. Vegetative propagation by ventral tubers and branches; tubers variable in size, with rounded, imbricate lobes, ventral scales and numerous rhizoids. Illustrations. Proskauer (1951, Figs. 1-22, 25); Srivastava & Dixit (1996, Fig. 10: 1-12). Distribution and ecology. The species has been collected in Central America (Costa Rica, Panama) and South America (Peru, Brazil). It grows in humid areas, on eroding banks of gorges, rivers and creeks, in shaded banks of roads, covered by herbs and Selaginella. Selected specimens examined. COSTA RICA. Alajuela, Cantón de Atenas, Alto del Monte, Salazar Allen et al. 16121(PMA); Cantón de San Mateo, Desamparados, Río Negro, Salazar Allen et al. 16124b, 16128, 16129 (PMA, SJ). PANAMÁ. COCLÉ: El Valle de Antón, Río Las Mozas, Salazar Allen et al. 14165, 14432, 15715, 15717, 15722, 15723, 16560, 16568, 16572, 16574, 16581(PMA); Rodríguez 171 (PMA). LOS SANTOS: Rivera 202, 213, s.n. (PMA). PANAMÁ: Condado de El Rey, Salazar Allen et al. 16080 (PMA); Cuevas de Chilibre, Salazar Allen et al. 16667 (PMA); Parque National Altos de Campana, Salazar Allen et al. 16549, 16674, 16675, 16679 (PMA); Parque National Soberanía, La Cascada, Salazar Allen et al. 16688, 16693, 16699 (PMA), Rodríguez 161, 162 (PMA), Sendero de El Charco, Salazar Allen et al. 16699, 16700 (PMA), Río Pedro Miguel, Camino de Cruces, Salazar Allen et al. 14383, 14389, 16549, 16602, 16603, 16609, 16610, 16614, 16625, 16627, 16631, 16633, 16640, 16648, 16650, 16654, 16663, 16741 (PMA). Cyathodium spruceanum grows in medium-sized to large patches. The thallus is elongate and its size varies depending on environmental conditions. The rather opaque appearance contrasts with the bright metallic-green, rosette-forming thalli of C. cavernarum and C. bischlerianum that grow in the same habitats. Spore ornamentation is also variable, some populations having high, well-developed lamellae arranged in regular areoles while others, collected at the end of the rainy season, have low, cord-shaped, thickened lamellae and a loose, open reticulum. Ventral tubers appear to germinate while favourable conditions prevail and are abundant during the peak of the rainy season. New thalli have been observed growing at the onset of the dry season among populations that have started to die; pH of soil analysed for 14 sites in Panama ranges from acidic to slightly alkaline (5.4-7.6).

5. Cyathodium steerei Hässel, Rev. Bryol. Lichénol. 30: 223-231. 1961. Type. Argentina. Tucumán: Valley of Río de las Sosas, El Nogalar, 4 Dec 1960, Hässel de Menéndez 390 (holotype, BA). Figs. 98, 99 Thallus bright green, thin, with a midrib, 2.5-8.0 mm long, 1.5-3.0 mm wide, dichotomously branched, marginal cells slightly crenulate, with specialized idioblasts between two or each

Bischler et al. 109 consecutive cell, decomposed in older portions of thallus. Dorsal epidermis delicate, cells lenticular, markedly bulging internally into air chambers, thin-walled, chlorophyllous, at midthallus 36-60 µm diameter, oil bodies brown, in specialized cells devoid of chloroplasts, 28-30 µm diameter. Epidermal pores simple, increasing in size in older portions of thallus, 120-180 µm diameter, surrounded by 1-3 (-4) rings of thin-walled cells, projecting slightly above surface, uppermost ring of 5-21 cells abutting like papillae into pore, second and third (-4) rings of cells incomplete, with 5-10 elongate cells. Ventral epidermal cells elongate, thin-walled, at midthallus 60-96 µm long, 24-26 µm wide, longer and wider towards base; multistratose area present in tubers and near antheridial receptacles, with few chloroplasts, without oil bodies. Rhizoids of two types, wide and slightly undulate, and narrow with spiral twisting and tuberculate. Ventral scales lanceolate, 5-7 cells long and 2-3 cells wide with 1-6 marginal cells with a brown oil body. Dioecious. Antheridial receptacles apical, hemispherical, without lobules, rugose, nearly sessile, 1-1.5 µm diameter, surrounded at base by scales. Involucres to 2 mm long, with 5-10 archegonia, sac-like, opening by two lips apically, marginal cells with sinuous walls, brownish, ventral surface with scales. Sporophytes one per involucre, operculum of 4 thickwalled central cells and inner tier of 8-9 cells with annular thickenings (Srivastava & Dixit, 1996). Spores brown, 33-40(-44) µm diameter, with irregular lamellae, 7-19 µm long, 2-6 µm high, upper border sinuous. Elaters reddish brown, 600-780 µm long, 16-19 µm wide, attenuate at both ends, with 3-4 spirally thickened bands, bispiral at extremes. Vegetative propagation by tubers with triangular or rounded imbricate, lunulate lobules, scales and rhizoids. Illustrations. Hässel de Menéndez (1961, Pl. I; 1963, Figs. 17: A-G, 18: A-F); Srivastava & Dixit (1996, Fig. 13: 1-9). Distribution and ecology. The species is only known from Argentina and grows on soil and on rock covered by thin soil. It is possible that with further collecting the range of the species will extend to other neotropical countries. Specimens examined. ARGENTINA. TUCUMÁN: Valley of Río de las Sosas, El Nogalar, M. Grassi s.n. (BA); Tafi, S. Javier, Horco Molle, Vervoorst 6877 bp (BA). Cyathodium steerei is similar in spore ornamentation to C. spruceanum, but thallus size, presence of oil bodies in idioblasts, number of opercular cells, and the presence of inner and outer involucral scales readily distinguish C. steerei from its sister species.

5. TARGIONIACEAE Dumort., Analyse Fam. Pl. 68, 70. 1829. Type. Targionia L. The family is monogeneric.

7. TARGIONIA L., Sp. pl. ed. 1, 1136, 1753. Type. Targionia hypophylla L. Genus dedicated to

Bischler et al. 110 Giovanni Targioni-Tozzetti (1712-1783), Professor in Florence. Thallus 1.5-4 mm wide, deep green to light green, often tinged with black-purple. Branching dichotomous and ventral. Epidermal pores simple, with 1-3 rings of 6-9 cells, radial walls thin or thickened. Air chambers in one layer, with chlorophyllose filaments. Oil-cells numerous. Ventral scales in 2 rows, dark purple, with oil-cells, marginal papillae and a single large appendage. Asexual reproduction by specialized propagules lacking. Monoecious. Antheridia in irregular groups dorsal or terminal on main thallus, or on short ventral branches; scales associated with androecia lacking. Archegonia 1-10 per archegonial cluster, terminal in mussel-shaped involucre on ventral side of thallus apex. Involucre dark purplish, 4-8-layered; scales associated with gynoecia lacking. Calyptrae 3-4-layered after fertilization. Sporophytes 1-3 per involucre, with bulbous foot and hardly elongating seta. Capsules opening by an irregularly disintegrating lid and 5-8 irregular valves to 1/3 of capsule length; walls with annular thickenings. Spore : elater ratio 4 : 1. Spores approx. 3000 per capsule, 50100 µm diameter, proximal and distal faces with reticulate areoles and reticulate ridges, trilete scar distinct. Elaters with 2-3 helical bands. Gametophytic chromosome number n = 9 or 27. The genus Targionia is widely distributed in warm-temperate regions, between 55°N to 42°S. It is particularly frequent in Mediterranean-type climates and is absent in humid tropical lowland and arctic and boreal regions. The genus comprises two drought-tolerant species in the Neotropics, colonizing rock crevices and rock outcrops in areas with dry seasons. The two species have been attributed to distinct subgenera by Schuster (1992b): subgen. Targionia (Type. T. hypophylla L.) and subgen. Prototargionia R.M.Schust. (Type. T. stellaris (Müll. Frib.) Hässel).

KEY TO THE NEOTROPICAL SPECIES OF TARGIONIA 1. Epidermal pores 80-170 µm diameter, bounded by 2-3 concentric rings of cells, radial walls thin. Appendage of ventral scales lanceolate-ovate, more or less constricted basally, often fimbriate. Antheridia in small ventral branches. ................................................. ........................1. T. hypophylla 1. Epidermal pores 35-50 µm diameter, bounded by a single ring of 6-7 cells, radial walls strongly thickened. Appendage of ventral scales broad-based, triangular, with basal, pluricellular teeth. Antheridia in irregular groups on dorsal side of thallus...................................................2. T. stellaris

1. Targionia hypophylla L., Sp. pl., ed. 1, 1136. 1753. Type. Italy, Spain, Turkey: Istanbul (syntypes, OXF, n.v.).

Figs. 3: H, 100, 102, 104: K

Targionia mexicana Lehm. & Lindenb., in Lehmann, Nov. stirp. pug. 4: 27. 1832. Type. Mexico. Oaxaca: Schiede s.n. (n.v.). Targionia bifurca Nees & Mont., Ann. Sci. Nat., Bot. sér. 2, 9: 40. 1838. Type. Chile. Quillota,

Bischler et al. 111 Bertero s.n. (PC, syntype). Targionia convoluta Lindenb. & Gottsche, in Gottsche, Lindenb. & Nees, Syn. hepat. 576. 1846. Type. Mexico. Puebla: Chinantla, Liebmann 520 (n.v.). Targionia lorbeeriana Müll. Frib., Hedwigia 79: 78. 1940. Type. Italy. Sicily: Taormina, Huber s.n. (S). Thallus 2-5 mm wide, dark or bright green. Epidermal cells smooth, with slightly thickened walls and nodulose trigones. Epidermal pores 80-170 µm diameter, with 2-3 concentric rings of cells, radial walls thin. Ventral scales purplish-black, with lanceolate appendage, entire, crenulate or irregularly toothed, often slightly constricted basally. Monoecious. Antheridia on small ventral branches. Archegonia in purplish-black involucres. Spores brown, 65-75(-100) µm diameter, areolate on proximal and distal faces, areoles and ridges reticulate. Gametophytic chromosome number n =9 or 27. Distribution and ecology. The species is distributed throughout warm-temperate and tropical parts of the world. It has been recorded from Europe, up to southern Scandinavia and Iceland in the north, the Mediterranean area, Macaronesia, Saharian, tropical and South Africa, Mascarenes, Ascension Is., Madagascar, SW and Central Asia, Afghanistan, northern India, Sri Lanka, Sikkim, Nepal, China, Tailand, Korea, Taiwan, Japan, Malaysia, Java, Oceania, New Zealand, Australia, and Tasmania. It is known from Canada, U. S. A. (Arizona, California, Colorado, Nevada, New Mexico, Oregon, Texas, Utah, Washington) in the New World, and in the Neotropics from Mexico (Guadalupe Is. (Sutliffe 1932), Baja California, Chiapas, D. F., Durango, Hidalgo, Mexico, Michoacán, Morelos, Oaxaca, Puebla, Querétaro, Tlaxcala, Veracruz), El Salvador (Haupt, 1942), Nicaragua, Costa Rica, Puerto Rico, Colombia (Meta, Magdalena (Winkler, 1976)), Ecuador (Chimborazo, Galápagos (Isabela)), Peru (Amazonas, Ancash, Apurimac, Arequipa, Cajamarca, Cuzco, Huancavelica (Buchloh, 1926), Huanuco, Junín (Stephani, 1916), La Libertad, Lima (Bryan, 1929)), Brazil (Goiás, Rio Grande do Sul (Vianna, 1976)), Bolivia (Chuquisaca, Cochabamba (Hässel, 1963), La Paz, Potosí, Tarija), Chile (Santiago, Valparaiso (Montagne, 1838)), Argentina (Buenos Aires, Catamarca, Cordoba (Hässel, 1963), Mendoza (Hässel, 1963), Neuquén, Rio Negro (Hässel, 1963), Salta (Hässel, 1963), Tucumán (Hässel, 1963)). The species is quite common in areas with dry seasons where it grows in dense colonies on periodically moist or wet clay, sandy clay, or sand, overlying limestone, dolomite, tuff or other calcareous rocks, basalt, lava, or volcanic rocks, or on soil in rock and wall crevices, often exposed, sometimes in partial shade, on banks along intermittent rivers, in grassland, open shrub vegetation and open forests, also in towns and ruins, from 600 to 4420 m, but mostly above 2000 m. Selected specimens examined. MEXICO. S. loc., Ehrenberg s.n. (G), Deppe 1100 (G). BAJA CALIFORNIA: Ceralvo Is., Steere 17566 (NY). CHIAPAS: Road San Cristóbal-Ocosingo, km 15, Gradstein 8319 (MEXU); 2 km N of San Cristóbal de las Casas, den Held & van Rhijn HH 42

Bischler et al. 112 (MEXU, U); Mun. Angel Albino Corzo, Cuxtepeque, Breedlove 67458, 71090 (MEXU); N of Mapastepec, Sharp 4443b (TENN). D. F.: Botanical Garden, 1979, Jovet-Ast s.n. (PC); Desierto de los Leones, 1944, Ruiz Oronoz s.n. (MEXU), Sharp 495b (TENN); Pedregal de San Angel, Universidad, Delgadillo 2088 (MEXU); Salazar, 1949, Herrera s.n. (MEXU). Parque Ejidal San Nicolas, Totolapan, Long 29550 (E). DURANGO: 1896, Palmer 709 (FH). HIDALGO: Road Pachuca to Real del Monte, 1948, Ruiz Oronoz & Herrera s.n. (MEXU); San Miguel de Regla, 1948, Ruiz Oronoz s.n. (MEXU). MEXICO: Amecameca, P. Cortez, Eggers & Frahm 792236 (PC); La Escondida, 8 mi. E of Lerma, Hermann & Crum 20862 (PC); Mun. Oculilan, Lagunas de Zempoala, E side of Laguna Quila, Pócs 9546 P (MEXU); Cerro Gordo, W side of Santiago Tolman, N of Teotihuacan, Whittemore 4085 (MEXU). MICHOACÁN: Cerro de Tecolote near Zacapu, Sharp 3735 (MEXU, Hb. Fulford); peak SE of Angangueo, Whittemore et al. 3142 (MEXU); El Salto, 23 km E Morelia, Iwatsuki & Sharp 2975 (TENN). MORELOS: Tepoztico, 1949, Orega s.n. (MEXU). OAXACA: Sierra Juarez, Road Oaxaca-Tuxtepec, Düll 5/510 (PC); 108 km of Juchitlan, Panamerican road km 903, Düll 610/5 (PC); 23 km above Oaxaca, below La Cumbre, on road to Ixtlan de Juarez, Sharp et al. 2437 (MEXU, TENN); Miahuautlan, Sharp et al. 2207 (MEXU, TENN). PUEBLA: Mun. Xochitlan, Arroyo Atena, 15 km SE de Zapotitlan, Grimes 2744 (MEXU); above Tezuitlan, Sharp 9804 (MEXU, TENN). QUERÉTARO: Peña de Bernal, Long & Delgadillo 29700 (E); Sierra Gorda, Puerto el Pino, near Pinal de Amoles, Long & Delgadillo, 29711 (E). TLAXCALA: Acuit Palpilco, Arsène s.n. (PC); Amazoc, Sharp 384 (TENN). VERACRUZ: Pico de Orizaba, Galeotti 6966 (PC); Orizaba, Mohr 30 (G); 5 km N of Naolinco, Sharp et al. 2938 (TENN); Pedregal de la Joya, NW of Xalapa, Long et al. 29616 (E); Rio Jamapa between Huatusco and Coscomatepec, Long & Garcia 29682 (E). NICARAGUA. Dept. Esteli, Stevens & Grijalva 15688 (MO). COSTA RICA. El Rodeo, 1979, Jovet-Ast s.n. (PC). PUERTO RICO. P. R. Rucio, Peñuleas, Pagán 1044 (MEXU). COLOMBIA. META: Paramo de Sumapaz, Cerro Nevado de Sumapaz, Cleef 1432 a, 1453 (U). ECUADOR. CHIMBORAZO: 15 km N of Alausi, Gradstein et al. 3367 (U). GALÁPAGOS ISLANDS: Isabela, Volcán Alcedo, SE side, Gradstein & Weber H 213 (JE, MEXU), Gradstein & Sipman H 446 (U). PERU. S. loc., d'Orbigny s.n. (PC). AMAZONAS: Prov. Chachapoyas, above Leimebamba, Frahm 2347 (U). ANCASH: NE of Huaraz, Laguna Llaca, Geissler 8855 (G). APURIMAC: Andahuaylas, near Chincheros, Hegewald 5764 (PC). AREQUIPA: Arequipa, Volk 6/0698 (JE, U). CAJAMARCA: Rio Chucsen (Mata-Namura), Hegewald 6144 (PC); Cerro Cunatan, S Contumaza, Hegewald 7344 (PC); Prov. Contumaza, Trujillo, Guzmango-Santiago, Sagastegui 10598 (U). CUZCO: Prov. Aula, Indahuacho (Cuzco-Abancay), Hegewald 5674 (PC). HUANUCO: La Unión, Huanuco Viejo, Geissler

Bischler et al. 113 8732 (G). LA LIBERTAD: Otuzco, Casmiche, Hegewald 5026 (PC); Huancamarca, Quebrada Hornillo, Otuzco, Hegewald 5167 (PC). BOLIVIA. CHUQUISACA: Luis Calvo, 4 km E of Muyapampa, Lewis 84-1019 (F); Oropeza, 10 km NE of Sucre, Lewis 83-1687 (F). LA PAZ: Inquisivi, 3 km NE of Quime, Lewis 83-3352 (F); Murillo, 33 km N of La Ceja de El Alto La Paz, Lewis 82-419, 453 (F, PC). POTOSÍ: Chayanta, 1 km NE of Macha, Lewis 84-1650 (F, PC). TARIJA: Gran Chaco, 15 km W of Palos Blancos, Lewis 842896 (F); O'Connor, 4 km NW of Canaletas, Lewis 84-2509 (F). BRAZIL. GOIÁS: Itebarai, Vital 1463 (PC). CHILE. S. loc., Bertero 97 (PC). SANTIAGO: Santiago, Costes 87 (PC). ARGENTINA. BUENOS AIRES: Sierra de la Ventana, Kühnemann 4145b (PC). CATAMARCA: Andalgala, Estancia Yunka Suma, Sleumer 1788 (LIL). NEUQUÉN: Lago Guillermo, Kühnemann 2272 (PC). Targionia hypophylla is defined here to include the morphologically similar and sympatric haploid and triploid cytotypes. The triploid was taxonomically separated from the haploid by Müller (1940), but as shown by Boisselier-Dubayle & Bischler (1999) the morphological characteristics of the two cytotypes (structure of epidermal pores, scales, spore size and ornamentation) overlap and their separation on morphological grounds therefore precluded. The chromosome number of the Linnean type of T. hypophylla is unknown. When the cytotypes of the species would be given specific rank, lectotypification of T. hypophylla is not possible. Both cytotypes are probably present in the Neotropics but chromosome counts are not yet available. Targionia hypophylla is easily distinguished in fertile condition by its bivalved, purplish-black involucres at apex on ventral side of thallus, and by its small, discoid male branches.

2. Targionia stellaris (Müll. Frib.) Hässel, Opera Lilloana 7: 74. 1963. Grimaldia stellaris Müll. Frib., Feddes Repert. 58: 61. 1955. Type (Hässel, 1963). Argentina. Jujuy: Dept. Tilarca, near San Gregorio, 4050 m, Sleumer 3596 (lectotype, LIL).

Figs. 101, 102

Thallus 1.5-2.5 mm wide, light green. Epidermal cells roughened, with strongly bulging trigones. Epidermal pores 35-50 µm diameter, with single ring of 6-7 cells, radial walls thickened. Ventral scales purplish-black, with triangular, acute appendage, not constricted basally, with one or several pluricellular teeth. Monoecious. Antheridia in several, irregular groups on dorsal side of thallus. Archegonia in purplish-black involucre. Spore morphology unknown. Gametophytic chromosome number unknown. Distribution and ecology. The species is rare and apparently endemic in southern South America. It was collected at high elevations (2800-4200 m) in northern Argentina (Jujuy, Tucumán).

Bischler et al. 114 Specimens examined. ARGENTINA. JUJUY: the type. TUCUMÁN: Valle de Tafi, Infiernillo, Lamb 5387 (LIL). The taxonomic status of Targionia stellaris needs further investigation based on living collections and sporophytes, which are unknown in this species. The material of T. stellaris reported from the Galápagos Islands (Gradstein & Weber, 1982) belongs to T. hypophylla. The epidermal pores in this sample are quite large and bounded by two rings of cells.

6. EXORMOTHECACEAE Müll. Frib. ex Grolle, J. Bryol. 7: 208. 1972. Type. Exormotheca Mitt. Thallus often perennating with tuberous apices, branching dichotomous, ventral and apical. Epidermal pores simple, with single ring of 5-10 cells, without hyaline inner ring. Air chambers in one layer, with chlorophyllose filaments, or empty. Ventral scales in 2 rows. Asexual reproduction by specialized propagules lacking. Monoecious or dioecious. Antheridia in dorsal groove, or in irregular groups behind gynoecia. Archegonia in terminal cushions. Archegoniophore developing after fertilization; receptacle with simple pores on dorsal side. Calyptrae 2-4-layered. Involucres tubular. Sporophytes with hardly or slightly elongating seta. Capsule wall with annular thickenings. Spore : elater ratio 4 : 1 or almost 1 : 1. Elaters with helical bands. Gametophytic chromosome number x = 8, or 9. The family includes three genera. Two are endemic in the western Himalayas, one is represented in the Neotropics.

8. EXORMOTHECA Mitt., in Godman, Nat. Hist. Azores West. Isl. 325. 1870. Type. Exormotheca pustulosa Mitt. Name from exormos = extruded, theca = capsule, referring to the short-exserted capsules. Thallus 1-4 mm wide, not tinged with purple. Epidermal pores very strongly elevated above epidermis, with ring of 5-6 (-8-10) hardly differentiated cells, radial walls thin. Air chambers with chlorophyllose filaments. Basal tissue with mucilage cavities. Ventral scales hyaline, with oil-cells and marginal papillae, without or with 1-2 filiform, often forked appendages, not constricted basally. Monoecious or dioecious. Archegonia 1-4 per archegonial cavity. Archegoniophore stalk without assimilatory strip, with 1 rhizoid furrow and scales at top; receptacle usually 2-lobed, with pores bounded by a ring of approx. 7 cells. Calyptrae 2-3-layered after fertilization. Sporophytes 1-2 per involucre. Capsules opening by an irregular lid and 4-5 irregular valves to 1/2 of capsule length. Spore : elater ratio almost 1 : 1. Spores 500-1500 per capsule, 50-150 µm diameter, distal face with plates or flat protuberances, proximal face granulose-tuberculate, trilete scar distinct. Elaters with 1-4 helical bands. Gametophytic chromosome number n = 8 or 16.

Bischler et al. 115 A genus of about seven drought-tolerant species of exposed rock outcrops and rocky soil, distributed in Mediterranean-type climates from 43°N to 33°S. The genus is everywhere rare. A single species occurs in the Neotropics.

1. Exormotheca pustulosa Mitt., in Godman, Nat. Hist. Azores West. Isl. 326. 1870. Type. Madeira, Pico de Barcellos, Johnson s.n. (NY, n.v.).

Figs. 1: D, 2: K, 103, 104: L-M, 109

Thallus 1-3 mm wide, blue-green, whitish when dry. Epidermal pores with ring of 8-10 cells. Basal tissue with numerous oil cells. Ventral scales extending beyond thallus margins, hyaline, appendage short, filiform. Monoecious. Spores brown, 52-67 µm diameter, distal face with plates. Elaters with 3 helical bands. Gametophytic chromosome number n = 16. Distribution and ecology. A rare but widespread species, recorded from the Mediterranean area, Macaronesia, tropical and southern Africa, the Mascarenes, and St. Helena. Two localities are known from Mexico (D. F. and Puebla). The species grows on sandy soil, on acidic cliffs and rock outcrops in rocky grassland at 2150-2300 m elevation. Neotropical specimens examined. MEXICO. D. F.: Botanical Garden, 1979, Jovet-Ast s.n. (PC), Düll A 78 (JE, PC), Düll s.n. (PC); Pedregal de San Angel, University, Delagadillo 1645, 2088, 2090 (MEXU). PUEBLA: 10 km E Puebla, Düll A 76 (PC). The species was thought to have been introduced in the Botanical Garden of Mexico (Gradstein et al., 1983) but has since been collected also in Puebla, in a natural environment.`

7. CLEVEACEAE Cavers, New Phytol. 10: 42. 1911. Type. Clevea Lindb. (Athalamia W. Falc.). Name honouring P. T. Cleve, Swedish phycologist and friend of the bryologist S. O. Lindberg. Thallus often perennating with tuberous apices, branching dichotomous, sometimes ventral or apical. Epidermal pores simple, with 1 or 2-3 concentric rings of 4-8 cells, with or without hyaline inner ring. Air chambers in 1-4 layers, empty. Ventral scales in several ill-defined rows. Asexual reproduction by specialized propagules lacking. Monoecious or dioecious. Antheridia dorsal, in irregular groups, often behind gynoecia, or on small ventral branches. Archegonia in dorsal or terminal cushions. Archegoniophore stalk developing after fertilization; receptacle with simple or without epidermal pores on dorsal side. Calyptrae 2-3(-5)-layered after fertilization. Involucres bilobed. Sporophytes with hardly short seta. Capsule wall with annular thickenings. Spore : elater ratio 4 : 1. Elaters with helical bands. Gametophytic chromosome number x = 9. A family comprising two or three genera, distributed mostly in arctic and alpine areas. Two genera are represented in the Neotropics.

Bischler et al. 116

KEY TO THE NEOTROPICAL GENERA OF CLEVEACEAE 1. Thallus often tinged with purple. Ventral scales without oil-cells, with or without marginal papillae. Archegoniophores dorsal (not terminal), stalk without rhizoid furrow. Involucre and sporophyte standing out obliquely from the side of the receptacle. ................................................... 9. Athalamia 1. Thallus without purplish pigmentation. Ventral scales with oil-cells and marginal papillae. Archegoniophores terminal, stalk with a single rhizoid furrow. Involucre and sporophyte hanging down from beneath the receptacle. ................................................................................... 10. Sauteria

9. ATHALAMIA W. Falc., Trans. Linn. Soc. London 20: 397. 1851. Type. Athalamia pinguis W. Falc. Name from a = lacking, thalamus = receptacle, referring to the lack of terminal receptacles. Clevea Lindb., Notiser Sällsk. Fauna Fl. Fenn. Förhandl. 9: 289. 1868, nom. illeg. Type. Clevea hyalina (Sommerf.) Lindb. (Marchantia hyalina Sommerf.). Thallus 1.5-6 mm wide, bright green, often tinged with purple. Branching dichotomous and ventral. Epidermal pores with single ring of 4-7 cells, radial walls thin or thickened. Air chambers in 1-4 layers. Basal tissue without mucilage cavities. Ventral scales without oil-cells and usually without marginal papillae, with 1(-3) filiform appendages, not constricted basally. Monoecious or dioecious. Antheridial groups often behind archegoniophore; scales associated with androecia lacking. Archegonia single per archegonial cavity, in dorsal cushions bounded by scales. Archegoniophore stalk dorsal (not terminal), without assimilatory strip or rhizoid furrows, with scales; receptacle 2-8lobed, with simple or without epidermal pores on dorsal side. Involucres 2-8, outwardly projecting, opening obliquely. Sporophytes one per involucre, with hardly elongating seta. Capsules splitting into 4-7 irregular valves to 1/2-3/4 of capsule length. Spores 2000-3000 per capsule, 40-70 µm diameter, distal and proximal faces with rounded protuberances, ornamented at top, trilete scar distinct. Elaters with 2-3 helical bands. Gametophytic chromosome number n = 9 or 18. A genus of 12-15 species of rocky habitats, distributed worldwide in cold and temperate areas. Two species are recorded from the Neotropics, one from the Andes (600-2800 m), the other from southern U. S. A. and Mexico.

KEY TO THE NEOTROPICAL SPECIES OF ATHALAMIA 1. Thallus 3.5-5 mm wide. Air chambers low and elongate, in 3-4 layers. Ventral scales not extending beyond thallus margins, usually with a basal tooth. Monoecious. Antheridia in narrow groups along thallus midline. Spores with 7-8 protuberances across diameter .............. 1. Athalamia andina

Bischler et al. 117 1. Thallus 1.5-3.5 mm wide. Air chambers high and narrow, in 1-2 layers. Ventral scales extending beyond thallus margins, without basal tooth. Dioecious. Antheridia scattered in large groups. Spores with 5-6 protuberances across diameter................................................ 2. Athalamia pygmaea

1. Athalamia andina (Spruce) S.Hatt., J. Hattori Bot. Lab. 12: 54. 1954. Clevea andina Spruce, Trans. & Proc. Bot Soc. Edinburgh 15: 569. 1885. Type. Ecuador. Tunguragua: river Pastaza near Baños; Ambato, 2000-2800 m, Spruce s.n. (syntypes, G).

Figs. 105, 107, 136: A

Targionia robusta Steph., Sp. hepat. 6: 3. 1917. Athalamia robusta (Steph.) S. Hatt., J. Hattori Bot. Lab. 12: 54. 1954. Type. Peru. Junín: Youli, turbera de Distichia, 4400-4500 m, Weberbauer 377 (n.v.). Thallus 3.5-5 mm wide, light green, margins purplish. Epidermal cells thin-walled. Epidermal pores 60-84 µm diameter, with ring of 6-7 cells, without hyaline inner ring, radial walls thin. Air chambers in 3-4 layers, low and elongate. Ventral scales purplish, lanceolate, acute apically, often with a tooth basally. Monoecious. Receptacle of archegoniophore 2-3-lobed, with hyaline scales. Spores 44-50 µm diameter, with 7-8 protuberances across diameter. Elaters with 3 helical bands. Distribution and ecology. A rare species, known only from Ecuador (Tunguragua), Peru (Huanuco (Bryan, 1929), Junín (Weberbauer, 1945), Lima (Carillo & Chanco, 1971), Bolivia (Cochabamba, Stephani, 1916, as Targionia robusta), and Argentina (Catamarca). It has been collected on wet clay along river borders, on shaded damp rocks, or on soil at the foot of steep, dripping rocks in river valleys and canyons, from 600 to 4500 m. Specimens examined. ECUADOR. TUNGURAGUA: Baños, Spruce s.n. (G); Ambato, Spruce s.n. (G). ARGENTINA. CATAMARCA: S. Ambato, Volk 6/0670 (JE). The spores of Athalamia hyalina have 7-8 protuberances across diameter, those of A. spathysii 6-7 and those of A. pygmaea 5-6. Spore wall ornamentation is very similar in the three species. The specimen attributed to Athalamia andina, cited by Carillo & Chanco (1971), belongs to Plagiochasma rupestre.

2. Athalamia pygmaea R.M.Schust., Phytologia 57: 411. 1985. Type. U. S. A. Texas: near Boot Spring, Chisos Mts., Big Bend Natl. Park, Dec 1981, Schuster 81-1251a (Hb. Schuster) (n.v.). Figs. 106, 107, 136: B Thallus 1.5-3.5 mm wide, light green, margins crisped, purplish. Epidermal cells thin-walled, with distinct trigones. Epidermal pores 40-80 µm diameter, with ring of 4-6 cells, without hyaline inner

Bischler et al. 118 ring, radial walls thin. Air chambers in 1-2 layers, high and narrow. Ventral scales purplish, lanceolate, acute apically, ending usually in row of 2 cells, without papillae and oil-cells, reaching or projecting beyond thallus margins. Dioecious? Antheridia in irregular, large groups, scales absent. Archegonia in cushions, bounded by a few, minute purplish or partly hyaline scales. Receptacle of archegoniophore 2-lobed, with purplish-hyaline scales. Spores 42-54 µm diameter, with 5-6 rounded protuberances across diameter Elaters with 2-3 helical bands. Distribution and ecology. Known from U. S. A. (Arizona, Texas, perhaps New Mexico (Schuster, 1992b)), Mexico (Mexico, Nuevo León). In Mexico the species grows in sheltered crevices of boulders and cliffs in arid scrub vegetation, between 1500-2800 m. Specimens examined. MEXICO. MEXICO: Television rebroadcast station on E slope of Cerro Gordo, W of Santiago, Tolman, just N of Teotihuacan, Whittemore 4096 (Hb. Whittemore). NUEVO LEÓN: Puente de Dios, ca. 8 km N of Galeana on road to Rayones, Whittemore et al. 2469 (Hb. Whittemore). The species has been reported from Mexico as A. hyalina (Whittemore, 1987), but in the latter, holarctic species the radial walls of the pores are strongly thickened. Athalamia pygmaea is more close to A. spathysii, occurring in the Mediterranean area and sharing with A. pygmaea hardly thickened radial walls in the epidermal pores. The thalli are more delicate in A. spathysii, with less developed ventral tissue, and the spore wall ornamentation is slightly different.

10. SAUTERIA Nees, Naturgesch. eur. Leberm. 4: 139. 1838. Type. Sauteria alpina (Nees) Nees (=Lunularia alpina Nees). Genus dedicated to A. Sauter, 1800-1881, medical doctor of Mittersill (Germany). Thallus 3-5 mm wide. Branching dichotomous and by apical adventitious branches after initiation of archegoniophores; ventral branches rare. Epidermal pores simple, with 1 or 2-3 rings of 5-8 cells, radial walls thin or thickened. Air chambers in 1-3 layers. Basal tissue without mucilage cavities. Ventral scales with oil-cells and marginal papillae, with 1(-2) filiform appendages, not constricted basally. Monoecious. Antheridia in irregular groups, dorsal on main thallus behind archegoniophore, or on small ventral branches; scales associated with androecia lacking. Archegonia 1(-2) per archegonial cavity, in terminal cushions, bounded by scales. Archegoniophore stalk terminal, without assimilatory strip, with 1 rhizoid furrow and scales at top; receptacle 2-7-lobed, with simple or without epidermal pores on dorsal side. Calyptrae 2-3-layered after fertilization. Sporophytes one per involucre; mature sporophyte with slightly elongating seta. Capsules splitting into 4-6(-8) irregular valves to 1/2 of capsule length. Spores approx. 2000 per capsule, 45-80 µm diameter, proximal and distal faces with rounded, tuberculate protuberances, trilete scar distinct. Elaters with 2-4 helical bands. Gametophytic chromosome number as far as known n = 36.

Bischler et al. 119 A genus of about 5 species of rock crevices and rock outcrops, mostly arctic-alpine in distribution, with a single neotropical representative.

1. Sauteria chilensis (Lindenb. ex Mont.) Grolle, J. Hattori Bot. Lab. 58: 200. 1985. Grimaldia chilensis Mont., in A. D. Orbigny, Voy. Amér. Mérid. 7, Bot. 2: 56. 1839. Neesiella chilensis (Lindenb. ex Mont.) Steph., Sp. hepat. 1: 95, 1898. Type (Grolle, 1975). Chile. Valparaiso: Quillota, Bertero 1129 (lectotype, PC; isolectotypes, BM, G, STR).

Figs. 108, 109, 136: C

Sauteria berteroana Mont., in Gottsche, Lindenb. & Nees, Syn. hepat. 541. 1846. Sauteria berteroana Mont., in A. D. Orbigny, Voy. Amer. Mérid. 7, Bot. 2: 56. 1839, nom. nud. Grimaldia debilis Bisch., in Gottsche, Lindenb. & Nees, Syn. hepat. 552. 1846. Type. Chile. Valparaiso: Mt. La Leona, Bertero 354 (syntype, PC). Thallus 1.5-2.5 mm wide, dark green with purplish borders. Branching dichotomous and by apical adventitious branches. Epidermal cells thin-walled. Epidermal pores 28-38 µm diameter, with single ring of 4-7 cells, radial walls thickened. Air chambers in single layer, narrow. Basal tissue with oilcells. Ventral scales purplish or hyaline, with oil-cells; appendage long, acute, 2-3 cells wide basally, ending in row of 4-10 cells. Monoecious. Antheridia in irregular groups behind archeoniophore. Archegonia bounded by hyaline scales. Archegoniophore with 3-5-lobed receptacle, with simple epidermal pores bounded by a ring of 5-6 cells. Involucres hyaline. Spores brown, 45-60 µm in diameter, with rounded protuberances on both faces. Elaters with 3-4 helical bands. Distribution and ecology. A rare species, known from Ecuador (Galápagos (Isabela), Peru (Loreto (Bryan, 1929), Bolivia (s. loc., Herzog, 1921), Chile (Valparaiso, O'Higgins (Montagne, 1850)), and Argentina (Catamarca (Hässel, 1963), Salta, Tucuman (Grassi, 1976)). It has been collected on bare, gravelly, volcanic soil among grassy vegetation or under bushes in rather dry and cool, high mountain environments, in the Andes from 2150 to 4000 m, and at lower elevations (1400 m) in pampa vegetation on the Galápagos islands (Gradstein et al., 2001). Specimens examined. ECUADOR. GALÁPAGOS ISLANDS: Isabela, Volcán Cerro Azul, Gradstein H 447 (U). CHILE. s.loc., Bertero s.n. (PC) ; s. loc., Gay 9260 (PC). ARGENTINA. SALTA: Cafayate, Cerro Santa Teresita, Vervoorst 5049 (BA).

8. AYTONIACEAE Cavers, New Phytologist 10: 42. 1911. Type. Aytonia J. R. & G. Forst., nom. rejic. (=Plagiochasma Lehm. & Lindenb. nom. cons.). Genus dedicated to John Ayton, hortulanus of Kew Gardens. Thallus perennating sometimes with tuberous apices; branching dichotomous, ventral and/or

Bischler et al. 120 apical. Epidermal pores simple, with (1-)2-6 concentric rings of 4-12 cells, with or without hyaline inner ring. Air chambers in (1-)2 to several layers, empty. Ventral scales in 2 rows. Asexual reproduction by specialized propagules lacking. Monoecious, rarely dioecious. Antheridia in irregular groups or in cushions, dorsal behind gynoecia, or terminal, or on small ventral branches. Archegonia in cushions, dorsal, or terminal on leading thallus segments, or on short ventral branches. Archegoniophore stalk developing after fertilization; receptacle with compound epidermal pores on dorsal side. Calyptrae 2-4-layered after fertilization. Involucres cup-shaped or bilabiate. Pseudoperianths splitting into linear segments, or absent. Sporophytes with hardly or slightly elongating seta. Capsules opening by a lid, wall without annular thickenings. Spore : elater ratio 4 : 1. Elaters with helical bands, or entirely thickened. Gametophytic chromosome number x = 9. The family Aytoniaceae comprises five genera, all of which are represented in the Neotropics. The family was subdivided into two subfamilies, Aytonioideae and Reboulioideae Grolle, by Grolle (1972a).

KEY TO THE NEOTROPICAL GENERA OF AYTONIACEAE 1. Antheridia dorsal or at bottom of apical adventitious branches, in cushions bounded by numerous, conspicuous scales. Archegoniophores dorsal (not terminal); stalk without rhizoid furrow. Involucres large, globose, opening sidewards ........................................................................... 14. Plagiochasma 1. Antheridia dorsal, terminal or on small ventral branches, in cushions or in irregular groups, scales few or absent. Archegoniophores terminal on main thallus, at base of apical adventitious branches or terminal on ventral branches; stalk with one rhizoid furrow. Involucres opening obliquely downwards ............................................................................................................................................................2 2. Each archegonium bounded by a large pseudoperianth splitting into lanceolate segments ........ .................................................................................................................................... 11. Asterella 2. Pseudoperianths absent ..............................................................................................................3 3 .Thallus scales with filiform appendages. Involucres strongly bilabiate...............................4 4. Thalli without tubers or tuberous apices. Epidermal cells with bulging trigones. Radial walls of epidermal pores usually strongly thickened. Antheridia in cushions, terminal on main thallus, or on small ventral branches, or in irregular groups dorsal behind archegoniophore, bounded usually by some minute scales. Female receptacle lobed. Capsule opening by a lid breaking in fragments. Spores with tuberculate areoles on distal face ....... .......................................................................................................................... 15. Reboulia 4. Thalli often with apical tubers. Epidermal cells without trigones. Radial walls of epidermal pores thin. Antheridia dorsal, in irregular, loosely aggregated groups behind

Bischler et al. 121 archegoniophore; no scales. Female receptacle hardly lobed. Capsule opening by a lid shed unbroken. Spores with smooth areoles or with rounded on distal face ................................ ................................................................................................................. 12. Cryptomitrium 3. Thallus scales usually with lanceolate appendages. Involucres cup-shaped ....... 13. Mannia

ASTERELLA

by D. G. Long

11. ASTERELLA P.Beauv. in Cuvier (ed.), Dict. Sci. Nat. 3: 257. 30 jan 1805 '1804'. Lectotype (Zijlstra, 1993). Marchantia tenella L. [= A. tenella (L.) P.Beauv.] (typ. cons.). From Aster = star, ella = little, referring to the star-like carpocephala with radiating pseudoperianths. Fig. 158: K-R Fimbraria Nees, Horae Phys. Berol. 44. 1820, hom. illeg. (Art. 53.1) non Fimbriaria Stackhouse 1809 (Rhodophyta). Lectotype (Evans, 1920). Fimbraria marginata Nees. [= Asterella marginata (Nees) S.W.Arnell]. Fimbriaria Nees, mut. Steudel, Nomencl. Bot. ed. 1, 2: 165. 1824, orth. var., hom. illeg. (Art. 53.1), non Fimbriaria Stackhouse 1809 (Rhodophyta). Type. Fimbraria Nees. Hypenantron Corda, in Opiz (ed.), Naturalientausch [12] (Beiträge zur Naturgeschichte. [1]): 648. after 21 Sept. 1829(–1830). Type. Hypenantron ciliatum Corda. [= Asterella saccata (Wahlenb.) A.Evans]. Thallus prostrate, sometimes with pungent 'fishy' smell, green, often purplish-pigmented especially beneath, thin and delicate or thick and leathery. Branching of terminal apical adventitious branches, terminal branches or stipitate-based ventral branches. Epidermal pores slightly elevated, simple, with 4–8 radial rows of cells in concentric rings, radial walls of cells sometimes somewhat thickened but not stellate. Assimilatory tissue of 1–several layers of overlapping air chambers which are empty or subdivided by secondary walls. Ventral tissue often mycorrhizal; oil-cells scattered. Rhizoids smooth and pegged. Ventral scales in two rows on midrib, colourless or violet to purplish pigmented,

Bischler et al. 122 asymmetric, with 1 or 2 lanceolate to ovate appendages often somewhat constricted at base, with scattered oil-cells in body and sometimes in appendage, margin with few to many slime-papillae. Asexual reproduction by specialized propagules lacking, but sometimes with tuberous persistent branch tips. Monoicous or dioecious. Antheridia scattered or in cushions on main thallus or main or reduced branches. Archegonia in cushions bounded by scales, 2-4 per archegonial cavity, terminal on main thallus or on large or reduced ventral branches. Archegoniophore stalk elongating after fertilization, with or without air chambers, with a single rhizoid furrow, with or without scales at top and bottom. Receptacle of mature archegoniophore subglobose when young, becoming conical, ovoid or inversely saucer-shaped when mature, 2–6 lobed, with compound pores. Involucres flap-like, usually hidden beneath lobes, Involucres opening obliquely downwards. Calyptrae 3-4-layered. Sporophytes one per involucre, with bulbous foot and scarcely elongating seta, enclosed by a colourless or pigmented pseudoperianth consisting of a unistratose basal cup and 5-18 lanceolate lobes attached at apex when young, remaining attached or becoming free at maturity. Capsules globose, dehiscing when ripe in upper 1/3 to 1/4 as an irregular or regular, discoid cap, wall without thickenings. Spore : elater ratio 4 : 1. Elaters with 1-3 helical bands. Spores 2000-4000 per capsule, 55-150 µm diameter, yellow, orange, brown or black, proximal and distal surfaces with similar or dissimilar sculpturing, areolate, alveolate or ridged, sometimes with equatorial pores, trilete mark distinct. Gametophytic chromosome number mostly n = 9. A genus of 80 to 90 species with an almost cosmopolitan distribution from the tropics to the Arctic, but with most species in temperate zones, especially in the mountains of Europe, Asia and America. In the New World eighteen species are accepted, of which 8 occur in North America and eleven in Central and South America; only one species, A. echinella, occurs in both North and Central America, and one, A. chilensis (Nees & Mont.) A.Evans is found only in southern South America outside the Flora Neotropica area. Five species have rather wide distributions in tropical America, A. dominicensis S. Arn., A. venosa (Lehm. & Lindenb.) A.Evans, A. macropoda (Spruce) A.Evans, A. elegans (Spreng.) Trevis., and A. lateralis M.Howe; the remainder are much more localised particularly in Mexico. None of the neotropical species occurs in other continents; most are found in mountainous regions except A. venosa (Lehm. & Lindenb.) A.Evans, A. elegans (Spreng.) Trevis. and A. echinella (Gottsche) Underw. which occur in the tropical lowlands. Some species are xeromorphic and drought-tolerant; others are hygromorphic and drought-intolerant. The xeromorphic species have inrolling thallus margins, while the hygromorphic species sometimes perennate by persistent tuberous thallus tips. Asterella species mostly grow on soil or rock, rarely on humus and never on wood. Many are weedy and colonise road-cuttings and unstable habitats. Substrates may be acidic or calcareous. They are found in grasslands, savannahs, forests, often close to water, some are found on old lava flows; in the tropics they are rare in lowland forests but frequent in the temperate and alpine zones, from sea level up to 4000 m. Carpocephala and sporophytes are commonly produced; gemmae are

Bischler et al. 123 absent.

KEY TO THE SUBGENERA OF ASTERELLA 1. Mature carpocephalum usually umbrella-shaped or flat; pseudoperianth lobes free at maturity; spores yellow to orange-yellow, proximal and distal surfaces with dissimilar sculpturing.................. .......................................................................................................................... 11a. subgen. Asterella 1. Mature carpocephalum hemispheric to conical, rarely umbrella shaped (A. macropoda); pseudoperianth lobes remaining firmly joined at apex; spores brown, red-brown to black, rarely yellowish, proximal and distal surfaces with similar sculpturing ................................................…. 2 2. Branching ventral and terminal; androecia on ventral, often reduced branches; mature carpocephalum hemispheric, rarely umbrella-shaped, shortly to deeply lobed; stalk slender with scattered scales at least when young; pseudoperianth not constricted at base of lobes; spores brown or red-brown or purplish-black, rarely yellowish............................................................................... .........................................................................................................11b. subgen. Phragmoblepharis 2. Branching mostly terminal; androecia on main thallus behind carpocephalum stalk, or antheridia on separate terminal branch, or dioecious; mature carpocephalum hemispheric to conical, almost unlobed; stalk stout, naked; pseudoperianth constricted at base of lobes; spores dark brown to black ....................................................................................................................... 11c. subgen. Saccatae

11a. ASTERELLA subgen. ASTERELLA Thalli hygromorphic, sometimes with seasonal xeromorphic branches with inrolling thallus margins, usually strongly aromatic; vegetative branches terminal dichotomies and terminal adventitious branches (ventral branches rare in neotropical species); assimilation tissue of one to several layers of spreading, overlapping air chambers; ventral scale appendages 1(-2), oblong to lanceolate, rarely ovate, acute to obtuse, constricted or not at base, margins entire, rarely toothed. Androecia borne on main thallus behind base of carpocephalum stalk, or on short terminal branches in neotropical species, of scattered ostioles or weakly cushion-forming; carpocephalum umbrella-shaped to almost flat, rarely conical or hemispheric, lobes shallow to deep; involucre with free margin entire (in neotropical species); pseudoperianth not compressed or constricted, lobes free at apex, divided approximately to half. Capsule lid fragmenting irregularly; spores yellow or orange-yellow, regularly areolate distally, proximally dissimilar, trilete, with distinct proximal pole; elaters mostly 2-spiral.

Bischler et al. 124 This subgenus is defined by its hygromorphic thalli, free pseudoperianth lobes and yellow spores with dissimilar proximal and distal areolate sculpturing. It comprises two sections and thirteen species. Section Asterella, of which the temperate North American A. tenella is the only representative, is not known in the Neotropics.

11b. ASTERELLA subgen. ASTERELLA sect. BRACHYBLEPHARIS (Gottsche et al.) D.G.Long, J. Bryol. 22: 113. 2000. Asterella subgen. Brachyblepharis (Gottsche et al.) Grolle, Feddes Rep. 87: 246, 1976. Fimbriaria subgen. Brachyblepharis Gottsche et al., Syn. hepat. 569. 1846. Type. A. leptophylla (Mont.) Grolle. Characters as for subgenus, but with carpocephalum flat or umbrella-shaped, not hemispheric when mature; involucre with entire free margin. The section contains twelve species and is widely distributed in the montane tropics and temperate regions: one species in Australasia, five in eastern Asia, three in tropical Africa, one in Macaronesia and SW Europe and three in Central and South America.

KEY TO THE SPECIES OF SUBGEN. ASTERELLA SECT. BRACHYBLEPHARIS 1. Plants with androecia on short terminal branches (sometimes very short and appearing lateral)..... .................................................................................................................................1. A. dominicensis 1. Plants with androecia borne on main thallus behind base of carpocephalum stalk ...... 2. A. venosa

1. Asterella dominicensis S.W.Arnell, Bryologist 61: 140. 1958. Type. Dominican Republic “Hispaniola”. Santo Domingo, Cordillera Central, prov. Azua, San Juan, Lomas de la Mediana, Sabana Nueva, edge of brook, ca. 1950 m.s.m., 16 sep 1929, Ekman 13584 (holotype, S). Figs. 110, 116 Fimbriaria pringlei Steph., Rev. Bryol. 36: 139. 1909, hom. illeg., non (Underw.) Steph. 1899. Type. Mexico. Distrito Federal: Cañada, valley of Mexico, 2800 m, 15 Oct 1908, Pringle 15347 (holotype, G-14956); synonymised by Grolle (1989). Asterella evansii del Rosario, Rhodora 68: 470. 1966, hom. illeg., non Kachroo 1958. Type. Mexico. Distrito Federal: Cañada Santa Magdalena, 15 Oct 1908, Barnes & Land 441 (holotype, NY); synonymised by Grolle (1989).

Bischler et al. 125 Thallus flat, delicate (hygroshoots) or narrower and somewhat leathery (xeroshoots), 2.5-5 mm broad, green, often tinged purplish below and on margins; branching mostly of terminal dichotomies and terminal adventitious branches. Epidermal cells thin-walled in hygroshoots, thick-walled and with trigones in xeroshoots. Epidermal pores with 3-4 concentric rings of cells in 5-8 rows. Air chambers large, in 1-2 layers. Ventral scales wine-red, asymmetric, with scattered oil cells and 1 marginal papilla, with 1 lanceolate appendage 2-4 cells broad at base, ending in 2-3 uniseriate cells. Antheridia in small cluster surrounded when young by scales, on long or short (sometimes very short and appearing lateral) terminal branches, sometimes several along margin of main thallus. Archegoniophore borne in apical notch on main thallus, stalk delicate, 10-15 (-35) mm long; receptacle conical when young, becoming umbrella shaped when mature, shortly 3-5-lobed. Involucre flap-like, entire. Pseudoperianth usually hyaline, divided into 7-12 lanceolate lobes which become free at apex. Spores yellow, 60–85 µm diameter, sculpturing dissimilar on distal and proximal faces, distally with 4–6 large areoles across face, areoles alveolate within, muri rounded in section, proximal facets alveolate, wing weakly differentiated. Elaters with 2 helical bands. Gametophytic chromosome number unknown. Distribution and ecology. Asterella dominicensis is known from Central America (Mexico, Guatemala, Costa Rica), the Dominican Republic in the Greater Antilles, and Venezuela, but is probably commoner than records suggest. It grows in moist shaded habitats on soil, in rock crevices and under boulders, on damp banks often near streams, in montane forests especially Pinus and Abies, mostly between 2000 and 3000 m altitude. Its appearance and micro-habitat preferences are similar to that of the next species but it occurs in cooler forests at higher altitude. Selected neotropical specimens examined. MEXICO. DURANGO: along Highway 40 about 9 miles west of La Ciudad, Bowers et al. 5238-a (E, F). D. F.: Parque Nacional Desierto de los Leones, Long 29592 (E).MEXICO: Sacro Monte, Amecameca, Barnes & Land 339 (F). MICHOACÁN: 25 km east of Morelia (at km 285 on highway 15), Iwatsuki & Sharp 5139a (JE). MORELOS: El Tepozteco, Ortega s.n. (MEXU). PUEBLA: toward Las Cuevas in Ixtaccihuatl above Huejotzingo, Sharp 4316 (F). QUERÉTARO: Sierra Gorda 2 km E of Pinal de Amoles, Long & Delgadillo 29747 (E).VERACRUZ: Pedregal de las Vigas above La Joya near Jalapa, Sharp et al. 2844 (MEXU, JE). GUATEMALA. 55 km N of Antigua, Volk 6/0711b (JE). COSTA RICA. PUNTARENAS: upper Río Burú, Gómez et al. 21701 (MO). DOMINICAN REPUBLIC. Cordillera Central, headwaters of Bao River, Liogier 3 (NY). VENEZUELA. MÉRIDA: Laguna Negra, Páramo de Macuchies, Chardón 25 (MICH).

Bischler et al. 126 Notes: some plants from Mexico have spores with the distal areoles less well-developed but in other respects the plants are typical.

2. Asterella venosa (Lehm. & Lindenb.) A.Evans, Contr. U. S. Natl. Herb. 20: 286. 1920. Fimbriaria venosa Lehm. & Lindenb., Nov. Stirp. Pug. 4: 29. 1832. Type (Grolle, 1989). Brazil. Without locality, Sellow s.n. (lectotype, S; isolectotypes, BM, STR, W).

Figs. 111, 116, 158: K-R

Fimbriaria lindmannii Steph., Bih. Kongl. Svenska Vetensk.-Akad. Handl. 3, 23 (2): 17. 1897. Type (Grolle, 1989). Brazil. Rio Grande do Sul: Porto Alegre, Ilha dos Banhos, at terram, ripae humidum, Lindman 83 (holotype, G-27250; isotypes, FH, S, UPS), synonymised by Hässel de Menéndez (1962). Thallus flat, delicate, 2.5-4 mm broad, green often purplish below and on margins; branching mostly of terminal dichotomies and occasionally terminal adventitious branches. Epidermal cells thin-walled, with small trigones. Epidermal pores with 2-3 concentric rings of cells in 3-6 rows. Air chambers large, in 1-3 layers. Ventral scales hyaline or wine-red, asymmetric, with scattered oil cells, apparently without marginal papillae, with 1 oblong-elliptic appendage, constricted and 2-4 cells broad at base, ending in a short bluntly pointed apex. Antheridia in a cluster behind base of stalk, without scales. Archegoniophore borne in apical notch on main thallus, stalk delicate, 4-15 mm long; receptacle flat or umbrella-shaped when mature, very shortly 3-6-lobed. Involucre flap-like, entire. Pseudoperianth usually hyaline, divided into 6-10 lanceolate lobes which become free at apex. Spores yellow, 60–75 µm diameter, sculpturing dissimilar on distal and proximal faces, distally with 4–7 large areoles across face, areoles alveolate within, muri rounded in section, proximal facets alveolate, wing weakly differentiated. Elaters with 2 helical bands. Gametophytic chromosome number unknown. Distribution and ecology. Asterella venosa is widespread in the southern neotropics, particularly in Brazil (many records, see Yano, 1981a), Uruguay and Argentina (Hässel de Menéndez, 1962). It is absent from the central neotropics and is rare in the northern part (only Mexico). The specimen from Suriname may be mislabelled. It typically grows on moist shady soil under herbs, amongst rocks and in crevices on cliffs, often along paths and streamsides, in moist broad-leaved and Pinus forests, between 200 and 1285 m. Habitats are similar to those of A. dominicensis but it is more typical of lowland forests. Selected neotropical specimens examined. MEXICO. MICHOACAN: about 2 km south of Patzcuaro, Clayton B-309 (F). PUEBLA: banks along Avenida Hidalgo and path to barranco, Teziutlan, Barnes & Land 555 (F, NY). VERACRUZ: Río Jamapa between Huatusco and

Bischler et al. 127 Coscomatepec, Long 29679 (E). SURINAME: without locality, Wullschlägel 1265 (JE). BRAZIL MINAS GERAIS: Caldas, 1873, Mosén s.n. (BM, FH, G-14964, JE, S, US). PARANÁ: Catartas do Iguaçu, ca. 20 km S of Foz do Iguaçu, on Rio Iguaçu border between Brazil and Argentina, Vitt 21319 (JE). RIO GRANDE DO SUL: Mun. Bom Jesus, on margin of Antas River, Vital 2019 (JE). SÃO PAULO: Mun. Eldorado, 50 m from the Caverna do Diabo entrance, Vital 1443 (JE). PARAGUAY. Perayubi, on river banks, Balansa 1279 (PC). ARGENTINA. BUENOS ARES: Cap. Federal, Punta Chica, 2 Oct 1956, C. de Donterberg 9409 (NY). URUGUAY. MINAS: Penitente, Herter 87865 (PC). The related A. chilensis (Nees & Mont.) A.Evans is not known from the Flora Neotropica area but occurs in Chile and southern Argentina (Hässel de Menéndez, 1962); it is similar to A. venosa in its sexual condition. It differs, however in its lanceolate acute appendages which are not constricted at the base and end in a long 1-2-seriate filiform apex (Hässel de Menéndez, 1962; Grolle, 1989).

11c. ASTERELLA subgen. PHRAGMOBLEPHARIS Grolle, Feddes Rep. 87: 246. 1976. Type. A. elegans (Spreng.) Trevis. Thalli xeromorphic or hygromorphic, aromatic or not; vegetative branches mostly ventral or ventral and terminal; assimilation tissue of 1(–3) layers of narrow vertical or irregular spongy air chambers, with or without lower spongy layer; ventral scale appendages 1–2 (–3), linear, lanceolate or ovate, constricted or not at base, apex subulate, acute or obtuse, margin entire or crenate. Androecia cushionlike, short, on ventral, often reduced branches; carpocephalum hemispheric, rarely umbrella-shaped, shortly to deeply lobed, stalk slender with scattered scales at least when young; involucre with free margin entire (in neotropical taxa); pseudoperianth not flattened or constricted, divided approximately to half, lobes remaining united at apex. Capsule lid fragmenting irregularly; spores yellow to brown, red-brown or purplish-black, with regularly and completely areolate or alveolate ornamentation similar on proximal and distal surfaces, trilete, with distinct proximal pole; elaters (1–)2-spiral. Subgenus Phragmoblepharis is the largest in the genus with over 20 accepted species worldwide. It is heterogeneous in several characters (e.g., involucre division, spore colour), but is united by its xeromorphic thalli with ventral and often also terminal branching, hemispheric carpocephala, united

Bischler et al. 128 pseudoperianth lobes and areolate spores with similar sculpturing on both surfaces. Grolle (1976) segregated some species as Sect. Pappiae and Sect. Lindenbergianae. Subdivision into sections is necessary but must be based on a broader survey of species worldwide. This is likely to result in recognition of four or more sections. The neotropical species all appear to be related to A. elegans, the type of the subgenus, and should therefore be placed in sect. Phragmoblepharis.

KEY TO THE SPECIES OF SUBGEN. PHRAGMOBLEPHARIS

1. Thalli 5–8 (–9) mm broad; ventral scale appendages lanceolate to ovate, acute to obtuse, + constricted at base; receptacle umbrella-shaped......................................... 6. A. macropoda 1. Thalli 1.5–5 (–6) mm broad; ventral scale appendages narrowly lanceolate or subulate, not constricted at base; receptacle hemispheric ................................................……………………….. 2 2. Spores without areoles but with 25-30 minute alveolae across distal face........... 7. A. versicolor 2. Spores areolate with 4-7 large areoles across distal face................ …………………………… 3 3. Carpocephala with finger-like tubercles 0.5–0.8 (–1) mm tall, 2-3 x taller than broad............... ..................................................................................................................................3. A. echinella 3. Carpocephala with rounded tubercles less than 0.5 mm tall to almost smooth, tubercles not taller than broad ............................................... ………………………………………………………. 4 4. Thalli hygromorphic with margins not inrolling when dry; carpocephala covered with low rounded tubercles .................................................................................................... .4. A. elegans 4. Thalli xeromorphic with margins incurved when dry; carpocephala covered with low rounded tubercles to almost smooth................................ ………………………………… 5. A. lateralis

3. Asterella echinella (Gottsche) Underw., Bot. Gaz. 20: 62. 1895. Fimbriaria echinella Gottsche, Kong. Danske Vidensk. Selsk. Skrift. ser. 5, 6: 367. 1868. Type. Mexico. Veracruz, Orizaba, Müller 2317 (isosyntype, G-15323); ibid., Müller 3317 (isosyntype, NY); ibid., Müller 2245 (isosyntype, NY); ibid., Müller 2327 (isosyntype, US).

Figs. 113, 116

Thallus flat, 2–3 (-4) mm broad, green but sometimes purplish beneath and on margins; branching

Bischler et al. 129 by apical adventitious branches and ventral intercalary branches, rarely by terminal dichotomies. Epidermal cells with weakly thickened walls. Epidermal pores of 2-3 concentric rings of 6-8 rows of cells. Air chambers irregular, spongy, in several layers. Ventral scales wine-red or dark purple, with scattered oil cells and marginal papillae, with 1-2 linear-lanceolate appendages, not constricted and 36 cells wide at base, margin entire, apex subulate. Antheridia in a small well-defined cushion surrounded by minute scales, on short stipitate-based ventral branch, more rarely on larger branch. Archegoniophore borne in apical notch on main thallus or on large ventral branch, stalk slender, 5-15 mm long, sparsely scaly but with beard of long whitish scales at apex; carpocephalum hemispheric, with (2-)4 short descending lobes, covered with tall finger-like tubercles 0.5–0.8 (–1) mm tall, 2-3 x as long as broad. Involucre an entire flap. Pseudoperianth lobes 8-10, hyaline or brownish, sometimes purplish, remaining united at apex. Spores brown or red-brown, 65–105 µm diameter, sculpturing similar on distal and proximal faces, distally with 5–6 large areoles across face, areoles + smooth within, muri flat in section, alveolate, wing moderately differentiated, sculptured. Elaters with 1–2 helical bands. Gametophytic chromosome number unknown. Distribution and ecology. Asterella echinella has a restricted distribution in Central America, Mexico, Guatemala, Belize and Honduras. It also extends to Arkansas and Texas in the southern USA (Schuster, 1992b). Ecologically it is similar to A. elegans in its preference for wet limestone outcrops along rivers, in ravines and by waterfalls, sometimes associated with the limestone fern Adiantum capillus-veneris, in warmer forests from 200 to 1240 m altitude. According to Whittemore (in lit.), it often grows in places submerged in fast-moving water at high water level which are at other times of the year very dry. Asterella echinella is closely allied to A. elegans, but differs in the usually smaller plants with narrower thalli, and in the tall finger-like tubercles on the receptacle. The two have very similar ecological requirements. This relationship was discussed by Evans (1920) and Schuster (1992). Some specimens of A. lateralis can be confused with A. echinella (see note under that species). Selected neotropical specimens examined. MEXICO. CHIAPAS: along Rio Lacantún upstream of Chajal, Gradstein 8175, Bryophyta Neotropica Exsiccata 389 (E, G, GOET, JE, MO, NY, S). NUEVO LEÓN: 22 miles SE of Monterrey, Horse-tail Falls, McGregor 4740 (NY). QUERÉTARO: 15 miles west of Jalpan at the falls, towards Querétaro, Sharp 3692 (JE); above Jalpan, Webster et al. 189 (FH). SAN LUÍS POTOSÍ: Upper Falls, El Salto, E of Ciudad Maiz, 1962, Sharp s.n. (MEXU, NY). TAMAULIPAS: 4 mi N of Magueyes, Hw 85, s. coll. 16289 (NY). VERACRUZ: Teocelo Canyon, just south of Xico and Jalapa, Hale & Soderstrom 21167 (US). GUATEMALA. ALTA VERAPAZ: Rio Cobán, 5 km NE of Cobán, Richards, Williams & Williams 2869 (F, H, NY, U).

Bischler et al. 130 BELIZE. CAYO: Monkey Tail River 5 miles E of Las Cuevas Field Station, Collins & Ensoll 5 (E). HONDURAS. CAMAYAGUA: Ojos de Agua 30 km N of Camayagua, Nelson et al. 6831 (MO).

4. Asterella elegans (Spreng.) Trevis., Rend. R. Ist. Lombardo Sci. Lett. ser. 2, 7: 785. 1874. Fimbriaria elegans Spreng., Syst. Veg. ed. 16, 4(1): 235. 1828. Type. [Santo Domingo] Hispaniola, 1820, Bertero s.n. (isotypes, G-15330, FH, JE).

Figs. 112, 117

Fimbriaria cubanensis Lehm. ex Mont., in Ramon de la Sagra, Hist. l’Ile de Cuba 489. 1842. Type. Cuba. Ad ripas fluminis Carima, 1839, Otto s.n. (isotypes, G-15322, S); synonymised by Evans (1920). Fimbriaria elegans Spreng. var. cubensis Gottsche, in Gottsche, Lindenb. & Nees, Syn. hepat. 565. 1846. Type. Fimbriaria cubanensis Lehm. ex Mont. Fimbriaria cubensis (Gottsche) Steph., Sp. hepat. 1: 125, 1899. Asterella austinii Underw., Bot. Gaz. 20: 64. 1895. Type. Cuba, Wright 136c (isotypes, G-15335, FH, MICH, NY); synonymised by Evans (1920). Asterella wrightii Underw., Bot. Gaz. 20: 64. 1895. Type. Cuba, Wright s.n. (holotype, NY; isotypes, G-15371, BM, US); synonymised by Evans (1920). Asterella reticulata A.Evans, Contr. U. S. Natl. Herb. 20: 302. 1920. Type. Cuba. Guantanamo, Monte Verde, Yateras, 19 iii 1860, Wright s.n. (holotype, YU, (n.v.); isotype, US); syn. nov., proposed by Grolle, in litt. Thallus flat, somewhat hygromorphic, 2–6 mm broad, green but purplish beneath and on margins; branching by apical adventitious branches and ventral intercalary branches, rarely by terminal dichotomies. Epidermal cells with weakly thickened walls. Epidermal pores of 3-4 concentric rings of 6-8 rows of cells. Air chambers irregular, spongy, in several layers. Ventral scales wine-red or purple, with scattered oil cells and short-lived marginal papillae, with 1(-2) linear to narrowly lanceolate appendages, gradually expanding and not constricted at base, margin entire, apex acuminate. Antheridia in a small n-defined cluster surrounded by scales on short stipitate-based ventral branch. Archegoniophore borne in apical notch on main thallus or on medium to large (rarely small) ventral branch, stalk slender, purplish, 5-17 mm long, with a few long whitish scales at apex; carpocephalum hemispheric, with (2-)4 short descending lobes, covered with low rounded tubercles. Involucre an entire flap. Pseudoperianth lobes 8-12, hyaline or brownish, remaining united at apex. Spores brown or red-brown, 75–120 µm diameter, sculpturing similar on distal and proximal faces,

Bischler et al. 131 distally with 5–7 large areoles across face, areoles + smooth within, muri flat in section, alveolate, wing moderately differentiated, sculptured. Elaters with 2 helical bands. Gametophytic chromosome number unknown. Distribution and ecology. Asterella elegans is a widespread and abundant species throughout the West Indies. There are several reports of A. elegans from Central America e.g. Guatemala but these mostly refer to A. echinella. Records from Mexico (e.g. Stephani, 1898-1900) are based on misidentifications of A. echinella and A. lateralis. It has a specialised ecology being restricted to wet limestone rocks, usually close to rivers, streams, wet rocks in ravines and by waterfalls (a rheophyte), sometimes on coral, often on outcrops of tuffs, usually in tropical or broad-leaved forests at low altitudes from sea level up to 1130 m. It often grows in extensive patches. Selected neotropical specimens examined. BAHAMA ARCHIPELAGO. North Andros, just north of Love Hill, Correll et al. 49333 (NY). CUBA. LAS VILLAS: Prov. Cienfugos, Trinidad Mts., Las Vegas de Mataguá, Morton 10775 (US, JE). ORIENTE: Mt. Libano, Maxon s.n., Kryptogamae exsiccatae 1689 (BM, H, JE, L, NY, G-15344, G-15331, S, US). PINAR DEL RIO: Rio Mestanza, Britton et al. 10160 (F, NY, U); Rio Mantaneales north of Candelaria, Britton et al. 14137 (NY). SANTA CLARA: Trinidad Mountains, Arroyo Grande, Britton 5483 (NY). SANCTI SPIRITUS: Las Veguetas, Banao, Leon & Caiderin 3517 (JE). JAMAICA. Road from Silver Hill Gap to Hardware Gap, Maxon & Phillip 1222 (F, FH, MICH, NY, S, US). HAITI. Vicinity of Cabaret, Baie des Moustiques, Leonard & Leonard 11957 (F, FH, JE, NY, US). DOMINICAN REPUBLIC. MONTE CRISTI: Cabo del Morro, El Morro, Buck 4889 (F, NY). PEDERNALES: ca. 37 km along road from Cabo Roja, from junction of road from Oviedo to Pedernales, Buck 4435 (NY). TACHUELA: Cordillera Central, Alto de Los Tramojos to El Cercado 2 km, Zanoni et al. 18166c (NY). PUERTO RICO. Rio Abajo Guajataca Project, Utuado, 20 iv 1941, Pagán 2157 (MICH, MO, NY).

5. Asterella lateralis M.Howe, Bull. Torrey Bot. Club 25: 189. 1898. Type. Mexico. Sinaloa, foothills of Sierra Madre, Colomas, 900 m, 18 vii 1897, Rose s.n. (holotype, NY; isotypes, G-23823, NY, US).

Figs. 114, 117

Bischler et al. 132 Fimbriaria atrispora Steph., Sp. hepat. 1: 106. 1899. Type. Mexico. Schaffner in herb. Levier 136 (lectotype nov., G-14950 p.p.); syn. nov. Fimbriaria arsenii Steph., Sp. hepat. 6: 11. 1917. Type (Grolle, 1989). Mexico, Arsén 7767 (lectotype, G-14949); synonymised by Grolle (1989). Fimbriaria mexicana Steph., Sp. hepat. 6: 15. 1917. Type. Mexico. Puebla, Santa Barbara, 19 xi 1910, Nicolas s.n. (lectotype nov., G-14952; isolectotypes, FH, JE); syn. nov. (synonymy also proposed by R. Grolle, in sched.) Fimbriaria elegans Spruce, Trans. Proc. Bot. Soc. Edinburgh 15: 563. 1897, hom. illeg. non Spreng. 1828. Thallus flat when moist, xeromorphic, becoming strongly inrolled when dry, 1.5–5 (–6) mm broad, green above, dark purple beneath and on margins; branching mostly by ventral intercalary branches. Epidermal cells with somewhat thickened walls. Epidermal pores of 3-4 concentric rings of 7-9 rows of cells. Air chambers irregular, spongy, in several layers. Ventral scales wine-red or purple, with scattered oil cells and marginal papillae, with 1-2 narrowly lanceolate appendages, not constricted at base, margin entire, apex subulate. Antheridia in a small cluster without scales on short stipitate-based ventral branches. Archegoniophore borne in apical notch on small obcordate stipitate-based ventral branch, stalk slender, 10-25 mm long, with a few whitish hanging scales at apex; carpocephalum hemispheric, with 2-4 descending lobes, covered with low tubercles to almost smooth. Involucre an entire flap. Pseudoperianth lobes (8-)10-12, hyaline, remaining united at apex. Spores brown or redbrown (light brown according to Whittemore, in lit.), 105–135 µm diameter, sculpturing similar on distal and proximal faces, distally with 4–5 large areoles across face, areoles + smooth within, muri flat in section, alveolate, wing strongly differentiated, almost smooth. Elaters with 2 helical bands. Gametophytic chromosome number unknown. Distribution and ecology. Asterella lateralis is a widespread species in Mexico and it occurs in Guatemala, Costa Rica, Panama, Colombia and Ecuador. Like A. elegans and A. echinella it appears to prefer calcareous substrates, but is a xerophytic species, growing in drier habitats, on shady banks, limestone walls, cliffs, in Quercus and Pinus forests and at higher altitudes, from 900 to 2830 m. A. lateralis is usually distinctive amongst neotropical species in having strongly xeromorphic thalli with both androecia and archegoniophores borne on short ventral branches, which are often numerous along the sides of the main thallus; the spores are also very distinctive in having a broad smooth wing without areoles (in related species the areoles on both surfaces usually extend to the outer margin of the spore wing). It can be confused both with A. echinella and A. macropoda. A. echinella differs in its more hygromorphic, usually flat thalli, its archegoniophores often borne on main thalli or on short

Bischler et al. 133 to long ventral branches and its carpocephala which have tall warts 2-3 x as long as broad; A. macropoda differs in having usually flat thalli up to 9 mm broad, acute to obtuse scale appendages, archegoniophores borne both on the main thallus and on small to large ventral branches, receptacles umbrella shaped and spores with areoles extending to the wings. The warts on the carpocephala of A. lateralis are variable in development, in some specimens the surface is almost smooth, in others it is strongly warty but the warts are scarcely taller than broad. A lateralis is very closely related to A. bolanderi (Austin) Underw. from California and the two may be better treated as subspecies; the only consistent difference is in the spores which in A. bolanderi have areoles extending to the equator, without a smooth wing as in A. lateralis. The type of Fimbriaria atrispora Steph. (G-14950), as annotated by Grolle in 1971, consists of a mixture of plants of A. lateralis (plants with antheridia on short stipitate-based ventral branches) and A. pringlei (plants with antheridia on separate terminal branch), and Stephani’s description is based on both elements. The A. lateralis element is selected above as lectotype. Selected neotropical specimens examined. MEXICO. BAJA CALIFORNIA SUR: Sierra de Laguna, 15 miles E of Todos Santos, Bowers et al. 5173-a (E, F). CHIAPAS: above Paval north of Mapastepec, Sharp 4430 (E, F). D. F.: Parque Nacional Desierto de los Leones, near Exconventa, Long 29598 (E). HIDALGO: 5 km W of Jacala, Sharp 81 (E, F). JALISCO: Rio Blanco, 16 km from Guadalajara, Barnes & Land 220 (NY, F). MEXICO: Amecameca, Pringle Pl. Mex. 10674 (JE, FH, MEXU, MICH, US). MICHOACÁN: between Chilchota and Zacupa on Mex. Hwy. 15, Weber & Charette B9852 (NY, US). MORELOS: Parque, near Cuernavaca, Pringle 15320 (MEXU, US). NAYARIT: Sierra Madre, Tepic, 1897, Rose s.n. (NY, G-27246, isosyntype of A. lateralis). PUEBLA: Tezuitlan, banks along Avenida Hidalgo and path to barranco, Barnes & Land 557 (F, NY). SAN LUÍS POTOSÍ: San Luis Potosí, 1876, Schaffner s.n. (NY). SINALOA: type of A. lateralis. VERACRUZ: Cordoba, 1888, Farlow s.n. (NY). GUATEMALA. SACATEPÉQUEZ: Cuesta de las Cañas, above Antigua, Standley 58935 (MICH, F, NY). SAN MARCOS: along Quebrada Canjulá, between Sibinal and Canjulá, Volcán Tacaná, Steyermark 36027 (F). QUEZALTENANGO: Volcán Zunil, Steyermark 34610 (F, JE). COSTA RICA. SAN JOSÉ: vicinity of Santa Maria de Dota, Standley & Valerio 43413 (F, JE, NY, US). PANAMA. CHIRIQUÍ: Alto Quiel, Boquete, Salazar Allen 3996 (PMA). COLOMBIA. CAUCA: Río Piendamó, between Popayán and Piendamó, Cuatrecasas 13859 (US).

Bischler et al. 134 ECUADOR. TUNGURAGUA: Baños, Spruce s.n. (BM, E, NY, US, G-15343, G-15332, G-15327, JE-H3566).

6. Asterella macropoda (Spruce) A.Evans, Bull. Torrey Bot. Club 46: 472. 1919. Fimbriaria macropoda Spruce, Trans. Proc. Bot. Soc. Edinburgh 15: 564. 1885. Type. Ecuador. Pichincha: “Andes Quitenses”, Mt. Pichincha, 3400 m, Spruce s.n. (holotype, MANCH (n.v.); isotypes, BM, E, G-15350).

Figs. 112, 116

Asterella boliviana (Steph.) A.Evans, Bull. Torrey Bot. Club 46: 477. 1919. Fimbriaria boliviana Steph., Sp. hepat. 6: 11. 1917. Type. Bolivia. Without locality, Bang s.n. (holotype, G-15317; isotype, NY); synonymised with A. fissisquama by Hässel de Menéndez (1962). Asterella fissisquama (Steph.) Hässel, Opera Lilloana 7: 113. 1962. Fimbriaria fissisquama Steph., Biblioth. Bot. 87: 172, 1916. Type. Bolivia.” Zwischen San Mateo und Sunchal”, 2900 m, Apr 1911, Herzog 4445 (holotype, G-15221; isotype, L); syn. nov. Fimbriaria mandonii Steph., Sp. hepat. 1: 133. 1899. Type. Bolivia. Mte. Sorata, Mandon s.n. (holotype, G-15356); synonymised by Evans (1919). Fimbriaria canelensis Spruce, Trans. Proc. Bot. Soc. Edinburgh 15: 564. 1885. Type. Ecuador. “Andes Quitenses”, Canelos, 1200 m, Spruce s.n. (holotype, MANCH (n.v.); isotypes, BR, E, G15319); synonymised by Evans (1919). Thallus flat, 5–8 (–9) mm broad, green but purplish beneath and on margins; branching by apical adventitious branches and ventral intercalary branches, rarely by terminal dichotomies. Epidermal cells with somewhat thickened walls. Epidermal pores of 3-4 concentric rings of 7-8 rows of cells. Air chambers irregular, spongy, in several layers. Ventral scales wine-red or purple, with scattered oil cells and short-lived marginal papillae, with 1-2 lanceolate to ovate appendages, + constricted and 710 cells wide at base, margin crenate, apex acute to obtuse. Antheridia in a small cluster without scales on short stipitate-based ventral branch. Archegoniophore borne in apical notch on main thallus or on large ventral branch, stalk slender, 10-40 mm long, with dense beard of long purplish scales at apex; carpocephalum umbrella-shaped with central boss, mostly with 4 deep lobes, these + truncate at apex and with low tubercles. Involucre an entire flap. Pseudoperianth lobes 8-12, hyaline or purplish, remaining united at apex. Spores brown or red-brown, 80–105 µm diameter, sculpturing similar on distal and proximal faces, distally with 5–6 large areoles across face, areoles + smooth within, muri flat in section, alveolate, wing moderately differentiated, sculptured. Elaters with 2 helical bands. Gametophytic chromosome number unknown.

Bischler et al. 135 Distribution and ecology. Asterella macropoda is a tropical-montane species found throughout the Andes from Central America (Costa Rica and Panama) south to Argentina (Hässel de Menéndez, 1962). It appears to be the most abundant Asterella species in Ecuador, Colombia and Bolivia. It often grows in disturbed habitats and has a broad ecological range, typically growing on soil or humus on shady banks, streamsides, in rock crevices and in grassy areas or under shrubs, in forest (e.g. Weinmannia forest) and at or above the tree-line in montane grassland and scrub, from (800m, Argentina) 1400 to 4000 m in most of its range. Records of the European A. lindenbergiana (Corda ex Nees) Arnell from Colombia refer to this species, e.g. Gottsche (1864), Gradstein & Hekking (1979). It is particularly distinctive in its broad flat thalli and deeply-lobed umbrella-shaped carpocephala. Selected neotropical specimens examined. COSTA RICA. CARTAGO: Cerro de la Muerte along Panamerican Highway, river valley in moist subpáramo ± at timber line, Gradstein 9728 (E, GOET). PANAMA. Approx 2 km SW of Itamut camp, Fabrega Bocas del Toro, Gómez et al. 22638 (U). COLOMBIA. ANTIOQUIA: 9.5 km E of Sonsón, on road to Argelia/ Nariño, Churchill & Sastre-de Jesus 12925 (MO). BOYACÁ: Nevado del Cocuy, Guican, salto de San Pablino, Bischler 2981 (JE, G52015). CAUCA: La Colonia, on road to El Tambo, 2300 m, 29 Jul 1958, Bischler 431, 433 (JE). CUNDINAMARCA: Tocarema, 1859, Lindig s.n. (BM); Río Colorado, 5 km S of lake Chisacá, Bischler 2306 (JE, G-52058). RISERALDA: Santa Rosa de Cabal, trail Termales to Finca la ‘Sierra’, Aguirre & Gradstein 1268 (U). SANTANDER: Páramo de las Puentes, above La Baja, Killip & Smith 18247 (JE, US). TOLIMA: Santa Isabel, Finca El Ochoral, Quebrada Las Damas, Aguirre & Gradstein 1568 (U). VENEZUELA. MÉRIDA: Sierra Nevada bei Mérida, Teleferico to Pico Espejo, near Estación La Aguada, Schäfer-Verwimp & Verwimp 12322 (JE, E). ECUADOR. LOJA: Cajanuma, Parque Nacional Podocarpus, along trail near Refugio, Gradstein 10014 (GOET). PICHINCHA: road from Lloa to Rio Cristal, south of Mt. Pichincha, Gradstein & Frahm 6702 (G-416644, U). TUNGURAGUA: Baños, along trail up Vulcán Tungurahua, 1989, Herot s.n. (G). PERU. LA LIBERTAD: Prov. Otuzco, Usquil, towards Otuzco, Hegewald 7174 (JE). BOLIVIA. CHUQUISACA: Prov. Luis Calvo, Serranía Iñao, 10 km NE of Monteagudo, Lewis 84990 (F). COCHABAMBA: Chapare, 2-3 km from Corani Dam by path to Corani Pampa, Wood 11092-a (E). LA PAZ: 9.5km NE of Chuspipata Solomon 8104 (MO). ARGENTINA, CORDOBA: Dept. Punilla, Sierra Grande de Cordoba, below Estafeta Casa Nueva

Bischler et al. 136 (Pedernera), 3 May 1936, C. C. Hosseus in Musci Americani Ser. III nr. 2053 (NY).

7. Asterella versicolor A.Evans, Contr. U. S. Natl. Herb. 20: 307. 1920. Type. Mexico. Jalisco, west end of the Sierra de San Esteban, Guadalajara, on rocks, 1600 m, 28 sep 1908, Barnes & Land 192 (holotype, YU (n.v.); isotypes, NY, US).

Figs. 115, 117

Thallus flat when moist, 1.5–5 (–6) mm broad, xeromorphic, becoming inrolled when dry, green but purplish beneath and on margins; branching mostly by ventral intercalary branches, sometimes by apical adventitious branches. Epidermal cells with somewhat thickened walls. Epidermal pores of 2-3 concentric rings of 5-7 rows of cells. Air chambers irregular, spongy, in several layers. Ventral scales purplish, with scattered oil cells and marginal papillae, with 1(-2) narrowly lanceolate appendages, narrow but not constricted at base, margin entire, apex subulate. Antheridia in an irregular cluster without scales on short ventral branch. Archegoniophore borne in apical notch on small ventral branch, stalk slender, 15-20 mm long, with scattered scales and beard of longer scales at apex; carpocephalum hemispheric, with 4 descending lobes with low tubercles. Involucre an entire flap. Pseudoperianth lobes 10-12, hyaline or purplish, remaining united at apex. Spores brown or purplishbrown, 100–115 µm diameter, sculpturing similar on distal and proximal faces, areoles absent, distally with 25–30 minute alveolae across face, wing moderately differentiated, sculptured. Elaters with 2 helical bands. Gametophytic chromosome number unknown. Distribution and ecology. Asterella versicolor is a poorly-known taxon described from Mexico (Evans, 1920, 1923). Only a few specimens agreeing with the type have been traced, from Costa Rica and Panama. Little information is available on ecology; the type grew on rocks at 1600 m altitude; the collections from Costa Rica and Panama grew on soil at 1500-1600 m and 1000-1400 m respectively. Although Evans compared this species with A. elegans and the European A. lindenbergiana (Corda ex Nees) Arnell, vegetatively it is very similar to A. lateralis and differs significantly only in its spores which lack areoles and are ornamented by finer alveoli. However, in other Asterella species such as A. dominicensis, development of areoles is sometimes suppressed and it is likely that A. versicolor is an atypical form of A. lateralis. Neotropical specimens examined. MEXICO. JALISCO: type. MORELOS: near Cuernavaca, Pringle 10667 (G-15326). COSTA RICA. ALAJUELA: San Miguel Oeste, Naranjo, Cerro Espiritu Santo, finca Napoleon Pérez, Gerardo Herrera 52B (MO).

Bischler et al. 137 PANAMA.CHIRIQUÍ: Alto Quiel, Boquete, Salazar Allen 3989 (PMA).

11d. ASTERELLA subgen. SACCATAE (Grolle) D.G.Long, J. Bryol. 22: 113. 2000. Asterella sect. Saccatae Grolle, Feddes Repert. 87: 171. 1976. Type. Asterella saccata (Wahlenb.) A.Evans Thalli xeromorphic or hygromorphic, without strong smell; vegetative branches mostly terminal; assimilation tissue with a single layer of air chambers or with ‘spongy tissue’ below; ventral scale appendages 1 or 2 per scale, lanceolate, not constricted, entire or toothed. Antheridia on main thallus behind base of archegoniophore stalk, on separate terminal branch, or dioecious; androecia scattered or aggregated in cushions, on main thallus or large terminal branches; carpocephalum stalk ridged or deeply lobed in type specimen, sometimes with vestigial air chambers, receptacle tall-hemispheric or rounded-conical; shortly-lobed or almost unlobed, stalk stout, naked; involucre with median cleft; pseudoperianth laterally compressed when young, transversely constricted at base of lobes, lobes united at apex, divided approximately to half-way. Capsule lid disc-like or fragmenting irregularly; spores dark brown or black, with ridged ornamentation similar on proximal and distal surfaces, trilete, with distinct proximal pole; elaters 2-spiral. Subgenus Saccatae as defined currently contains seven highly distinctive and geographically isolated species: A. saccata (Wahlenb.) A.Evans, the type of the subgenus, is a disjunct arctic-alpine species in Europe and Asia, A. muscicola (Steph.) S.Arnell is endemic to southern Africa, A. grollei D. G. Long is found in Nepal and China and A. palmeri (Austin) Underw. is endemic to California, in addition to the three neotropical species treated below. Some members of this subgenus may be artificially grouped together and the subgenus may require redefinition (or subdivision) in future. Of the three neotropical species, A. alpina is most closely allied to A. saccata. The other two species show similar characters of the pseudoperianth but show significant differences in other characters such as sexual condition and spore ornamentation.

KEY TO THE SPECIES OF SUBGEN. SACCATAE

1. Antheridia on main thallus behind archegoniophore stalk; carpocephalum conic …………….. 8. A. alpina 1. Antheridia on separate terminal branch or dioecious; carpocephalum hemispheric.......................2 2. Thallus 3.5-5 mm broad; antheridia on separate terminal branch; spores irregularly alveolate, alveolae 2–4 µm diameter ………………………… ....................................................9. A. pringlei

Bischler et al. 138 2. Thallus 8-15 mm broad; sexual condition dioecious; spores areolate, areoles ca. 10 µm diameter ………………………………………………... .......................................................... 10. A. rugosa

8. Asterella alpina (Steph.) D.G.Long, in Gradstein et al., J. Hattori Bot. Lab. 93: 9. 2003. Fimbriaria alpina Steph., Sp. hepat. 1: 137. 1899. Type. Bolivia. Tica, Mandon 21 (holotype, G15314).

Figs. 118, 120

Thallus thick, concave above, xeromorphic, becoming inrolled when dry, ca. 3 mm broad, green above, dark purple beneath and on margins; branching mostly by terminal dichotomies. Epidermal cells with thin walls. Epidermal pores of 2 concentric rings of 5 rows of cells. Air chambers tall and narrow. Ventral scales dark wine-red, with scattered oil cells and marginal papillae, with 1 narrowly lanceolate appendage, abruptly narrowed but not constricted at base, margin entire, apex subulate. Antheridia in a short elongate group without scales on main thallus behind base of archegoniophore. Archegoniophore borne in apical notch on main thallus, stalk stout, 5-20 mm long, smooth, without scales, ribbed longitudinally, in type specimen with deep groove and without air chambers; carpocephalum conic, unlobed, covered with low tubercles. Involucre not seen. Pseudoperianth constricted and compressed at base of lobes, lobes ca. 8, hyaline, remaining united at apex. Spores (only two, probably immature, seen), dark brown, 95–110 µm diameter, sculpturing similar on distal and proximal faces, without areoles, distally with 18–20 minute alveolae across face, wing absent. Elaters with 2 helical bands. Gametophytic chromosome number unknown. Distribution and ecology. Asterella alpina is only known from the type from Tica, Bolivia, which lacks any ecological information. Neotropical specimens examined. BOLIVIA. known only from type. The above description is based partly on the description of Stephani (1898-1900) and on the very fragmentary and inadequate type specimen. The distinctive shape of the carpocephalum and the structure of the pseudoperianth indicates a relationship with the northern hemisphere arctic-alpine A. saccata (Wahlenb.) A.Evans which differs in having a conspicuous beard of hyaline scale appendages at the apex of the thallus but otherwise is very similar, as far as can be evaluated from the very fragmentary material and probably immature spores of A. alpina. The Californian A. palmeri (Austin) Underw. is also superficially similar but differs in having air chambers in the carpocephalum stalk and in the broad, smooth ridges of the spores.

9. Asterella pringlei Underw., Bot. Gaz. 20: 64. 1895. Fimbriaria pringlei (Underw.) Steph., Sp.

Bischler et al. 139 hepat. 1: 109. 1899, non F. pringlei Steph. 1909 (= A. dominicensis). Type. Mexico. Jalisco, near Guadalajara, wet cliffs, 11 ix 1890, Pringle s.n. (holotype, NY; isotype, G-14955. Figs. 117, 119 Fimbriaria bakeri Steph., Sp. hepat. 6: 11. 1917. Type. Colombia. Santa Marta, near Bonda, xii 1898, Baker s.n. (holotype, G-15315; isotype, JE); syn. nov. (synonymy also proposed by R. Grolle, in sched.) Fimbriaria stahlii Steph., Sp. hepat. 1: 127. 1899. Type. Mexico. Veracruz, Orizaba, Stahl s.n. (lectotype nov., G-14957; isolectotype, FH); syn. nov., (synonymy also proposed by R. Grolle, in sched.). Asterella stahlii (Steph.) Tatuno, Cytologia 25: 219. 1960, nom. inval. Thallus flat or slightly concave when moist, somewhat leathery, but not becoming strongly inrolled when dry, 3.5–5 mm broad, green above, purple beneath and on wavy margins; branching mostly regularly terminal dichotomies, occasionally by apical adventitious branches. Epidermal cells with thin-walled cells without trigones. Epidermal pores of 2 concentric rings of 4-6 rows of cells. Air chambers of a single layer of larger chambers above, spongy below with several layers of smaller chambers. Ventral scales purple, with 1-3 oil cells, with marginal papillae, with 1(-2) large hyaline appendages, + lanceolate but variable in shape, not constricted at base, margin of appendages entire or with 1-5 irregular teeth which also bear papillae, apex subulate. Antheridia on separate terminal branch, sometimes also on main thallus behind base of archegoniophore stalk, scattered in a loose elongate cluster of tall ostioles, not surrounded by scales, on separate terminal branch or occasionally behind base of archegoniophore. Archegoniophore borne in apical notch on main thallus, limiting growth; stalk slender, rigid, 10-15 mm long, naked and without beard at apex, in type specimen deeply cruciform and with vestigial air chambers in wings; carpocephalum rounded-hemispheric, covered with rounded tubercles, lobes 4, short, descending. Involucre broad, thin, inflated, deeply cleft, + 2-lipped. Pseudoperianth constricted at base of lobes, lobes 12-16, hyaline, remaining united at apex. Spores black or black-brown, 80–105 µm diameter, sculpturing similar on distal and proximal faces, without areoles, with ca. 20 minute strongly thickened alveolae across distal face, alveolae 2–4 µm diameter, wing weakly differentiated, sculptured. Elaters with 1–2 helical bands. Gametophytic chromosome number n = 9 (Tatuno 1960 for Asterella stahlii). Distribution and ecology. Asterella pringlei occurs in Mexico, Guatemala and Colombia, and has been reported also from Costa Rica by Cole (1984). It is rare throughout its range. It grows in moist shaded habitats usually associated with rocks, cliffs or lava flows, in broad-leaved forests of Quercus, Platanus, Liquidambar, etc. between 1100 and 2300 m altitude. Selected neotropical specimens examined. MEXICO. CHIAPAS: between Finca Cuxtepeque and

Bischler et al. 140 Finca Cabañas, Municipio Angel Albino Corzo, Breedlove & Daniel 71090 (MEXU, MO, NY). JALISCO: Guadalajara, Barranco de Oblatoz, Barnes & Land 121 (F, NY). MEXICO: Pedregal, Teotihuacan, Düll 409 (JE). MORELOS: near Cuernavaca, 1905, Rose et al. 10242 (NY, JE, US). NAYARIT: Tepic, Orcutt 1136 (US). VERACRUZ: near Orizaba, Barnes & Land 671 (F, JE, NY). GUATEMALA. JALAPA: vicinity of Jalapa, damp shaded bank, Standley 76640 (JE, F). SANTA ROSA: along road south-east of Barberena, Standley 77800 (JE, F). COLOMBIA. SANTA MARTA: type of Fimbriaria bakeri.

10. Asterella rugosa A.Evans, Contr. U. S. Natl. Herb. 20: 289. 1920. Type. Mexico. Distrito Federal, La Cima, 3000 m, 14 x 1908, Barnes & Land 413 (holotype, YU, n.v.; isotypes, NY, US. Figs. 118, 120 Thallus thin and broad, flat, hygromorphic, not inrolling when dry, 8–15 mm broad, green above, purple beneath and on undulate margins; branching fan-like, by repeated apical adventitious branches and terminal branches. Epidermal cells with thin walls and small trigones. Epidermal pores of 3-4 concentric rings of 5-7 rows of cells. Air chambers irregular, spongy, in several layers. Ventral scales purple, with scattered oil cells and marginal papillae, with 1(-2) linear appendages, not constricted at base, margin often with 1-2 teeth, apex acuminate. Dioecious. Antheridia in distinct elongate-elliptic clusters without scales along middle of main thallus and main branches, often several clusters in a line. Archegoniophore borne at apex of main thallus but lobes at each side of stalk continuing fan-like growth, stalk short, stout, curved, 5-10 mm long, naked but with a few whitish scales at apex, in type specimen deeply ribbed and with vestigial air chambers in wings; carpocephalum hemispheric, unlobed, with low tubercles, bearing up to 4 pseudoperianths. Involucre not seen. Pseudoperianth lobes 12-14, hyaline or brownish, remaining united at apex. Spores (only four seen) black-brown, 100–120 µm diameter, sculpturing dissimilar on distal and proximal faces, distally with smooth narrow ridges forming incomplete areoles ca. 10 µm diameter, areoles and muri smooth, proximal facets with scattered large pores, wing weakly differentiated, sculptured. Elaters with 2 helical bands. Gametophytic chromosome number unknown. Distribution and ecology. Asterella rugosa is a highly localised endemic to Mexico which has been collected only a few times. It is restricted to shady humid recesses in lava flows, amongst scrub and in Abies/ Cupressus forest, from 2250 to 3080 m altitude. A very distinctive species, characterised by the apparently dioecious sexual condition, the very broad flat thalli with repeated fan-like branching, and the archegoniophores which do not limit

Bischler et al. 141 vegetative growth, which continues from the lobes on both sides of the stalk. However, the species is inadequately known due to lack of good herbarium specimens for study. Its relationships are not clear and it is placed in subgen. Saccatae with some reservations. Neotropical specimens examined. MEXICO. D. F.: Ciudad Universitaria, Pedregal de San Angel, Delgadillo 1660 (MEXU); road to Popocatepetl, ca. 14 km from Amecameca, Düll 2/365 (JE).

12. CRYPTOMITRIUM Austin ex Underw., Bull. Illinois State Lab. Nat. Hist. 2: 36. 1884. Platycoaspis Lindb., in Lindberg & Arnell, Kongl. Svenska Vet.-Akad. Handl., N.F. 23, 5: 11. 1889, nom. illeg. Type. C. tenerum (Hook.) Austin ex Underw. (Marchantia tenera Hook.). Name from crypto = hidden, mitra = helmet, referring to the female receptacle. Thallus 2.7-12 mm wide, green, often tinged with purple, perennating with apical tubers. Epidermal cells without thickened angles. Epidermal pores with 3-5 rings of 6-12 cells, radial walls thin. Air chambers in (1-)3-4 layers. Basal tissue without mucilage cavities. Ventral scales with oilcells and marginal papillae and 1-2(-3) filiform appendages, not constricted basally. Monoecious. Antheridia in irregular groups behind archegoniophore; scales associated with androecia lacking. Archegonia in terminal cushions, 1-4 per archegonial cavity. Archegoniophore stalk terminal, without assimilatory strip, with 1 rhizoid furrow, with scales at top, sometimes at bottom; receptacle very shallowly 2-6-lobed. Calyptrae 3-layered after fertilization. Involucres bilabiate. Sporophytes one per involucre. Capsules opening to 1/3 of capsule length by a lid shed unbroken. Spores approx. 1000 per capsule, 35-95 µm diameter, proximal and distal faces with more or less complete, smooth areoles and smooth ridges, or with rounded protuberances, trilete scar distinct. Elaters with 2-3 helical bands. Gametophytic chromosome number n = 9. A genus of 3 species of rock outcrops and rock crevices, with disjunct distribution in the New World, SE Africa and the western Himalayas.

1. Cryptomitrium tenerum (Hook.) Underw., Bull. Ill. State Lab. Nat. Hist. 2: 36. 1884. Marchantia tenera Hook., in Kunth, Syn. Pl. 1: 45. 1822. Duvalia tenera (Hook.) Gottsche, in Gottsche, Lindenberg & Nees, Syn. hepat. 554. 1846. Platycoaspis tenera (Hook.) Lindb., in Lindberg & Arnell, Kongl. Svenska Vetensk. Akad. Handl. 23, 5: 11. 1889. Type (Frye & Clark , 1937). Mexico. Michoacan: Ario, Humboldt s.n. (BM).

Figs. 121, 123, 136: D

Duvalia gayana Mont., Ann. Sci. Nat., Bot. sér. 3, 4: 354. 1845. Type. Southern Chile, St. Antoine, 1839, Gay s.n. (PC). Duvalia brevipedunculata Mont., in Gottsche, Lindenb. & Nees, Syn. hepat. 555. 1846. Type. Chile. Gay s.n. (n.v.).

Bischler et al. 142 Thallus 4-8 mm wide, yellowish green with purplish margins. Branching dichotomous and by apical adventitious branches, especially after initiation of archegoniophores. Epidermal cells thinwalled. Epidermal pores 120-150 µm diameter, with 3-4 rings of 6-9 cells, radial walls hardly thickened. Air chambers in 1-4 layers. Ventral scales purplish, with oil-cells, without marginal papillae, with single, filiform appendage, 1-3 cells wide basally, with row of 4-12 cells apically. Monoecious. Archegoniophore receptacle nearly disciform, very shallowly 2-6-lobed, with compound epidermal pores on dorsal side. Spores 60-65 µm diameter, yellow-brown, with smooth areoles. Elaters with 3-4 helical bands. Distribution and ecology. Cryptomitrium tenerum is known from U. S. A. (California, Washington), Mexico (Querétaro), Guatemala (Haupt, 1942), Costa Rica (Haupt, 1942), Chile (Aisén (Montagne, 1850), O'Higgins, Valparaiso), and Argentina (Salta (Hässel, 1963)). The species is common in California, but rare in tropical America. It has been collected on mountain rocks in rather mesic, seasonal climates, in forest on limestone, between 1500-2500 m. Specimens examined. MEXICO. Ario, Humbolt 236 (BM). QUERÉTARO: Sierra Gorda 2 km E of Pinal de Amoles, Long & Delgadillo 29748 (E). CHILE. O'HIGGINS: Pelequén, Dusén 149, 150 (G). VALPARAISO: 1896, Dusén s.n. (BM, G); Viña del Mar, Dusén 132 (G).

13. MANNIA Opiz, Naturalientausch 12 [Beitr. Naturg.] 646. 1829. nom. cons. Type. Mannia michelii Opiz nom. illeg. (Grimaldia dichotoma Raddi nom. illeg., Marchantia androgyna L., Mannia androgyna (L.) A.Evans). Genus dedicated to the Bohemian botanist and physician W. B. Mann (1799-1839). Grimaldia Raddi, Opusc. Sci. 2: 356. 1818, nom. illeg. Type. Grimaldia dichotoma Raddi, nom. illeg. Duvalia Nees, Ges. Naturf. Freunde Berlin, Mag. 8: 271. 1818, nom. illeg. Neesiella Schiffn., in Engler & Prantl, Natürl. Pflanzenfam. 1, 3: 32. 1893. Type. Duvalia rupestris Nees (Neesiella rupestris (Nees) Schiffn.). Cyathophora S.Gray, Nat. Arr. Brit. Pl. 1: 678. 1821. nom. rej. Type. Cyathophora angustifolia S. Gray nom. illeg. (Marchantia androgyna L. = Mannia androgyna (L.) A.Evans). Thallus 1.5-4 mm wide, often tinged with purple. Branching dichotomous, ventral, sometimes by apical adventitious branches. Epidermis rupturing in some species in older parts of thalli. Epidermal pores with (1)2-3 rings of 5-9(-10) cells, radial walls thin or thickened. Air chambers in 2 to several layers. Basal tissue without mucilage cavities. Ventral scales usually with oil-cells and marginal papillae, with 1-3 lanceolate appendages, not constricted basally. Monoecious or dioecious. Antheridia and archegonia as in the family, archegonia 1(-2) per archegonial cavity. Archegoniophore

Bischler et al. 143 stalk terminal, without assimilatory strip, with 1 rhizoid furrow and scales at top and bottom, or at bottom alone; receptacle shallowly (2-)3-5-lobed. Calyptrae 2-3-layered after fertilization. Involucres cup-shaped. Sporophytes one per involucre, bending obliquely downwards. Capsules opening to 1/3 of capsule length by a lid shed unbroken. Spores 2000-3000 per capsule, 50-90 µm diameter, distal and proximal faces with tuberculate areoles and tuberculate ridges, or distal face with rounded protuberances, proximal face reticulate-tuberculate, or tuberculate, trilete scar distinct. Elaters with 24 helical bands. Gametophytic chromosome number n = 9. A genus of 15 drought-tolerant species growing on rocky soil or in rock crevices. The genus is nearly worldwide in distribution, reaching northwards up to the Arctic but being absent from Southeast Asia and Australasia. The species often have fragmented ranges, some being arctic-alpine, others mainly Mediterranean. Two species are known from the Neotropics, M. californica and M. hegewaldii sp. nov. Grolle (1983b) subdivided the genus into two subgenera, Mannia (= subgen. Xeromannia, Grolle 1983. Type. M. androgyna (L.) A.Evans) and Arnelliella (C. Massal.) Grolle (= subgen. Mannia, Grolle 1983. Type. M. triandra (Scop.) Grolle), distinguished by the structure of the assimilatory layer. This subdivision of the genus was not accepted by Schuster (1992b). Mannia californica would belong to subgen. Mannia, M. hegewaldii, to subgen. Arnelliella.

KEY TO THE NEOTROPICAL SPECIES OF MANNIA 1. Ventral scales blackish purple. Epidermal cells thick-walled. Antheridia dorsal, in irregular groups without scales. Archegoniophores on short ventral branches. Spores with rounded protuberances on distal face ................................................................................................................... 1. M. californica 1. Ventral scales purplish with hyaline margins. Epidermal cells thin-walled. Antheridia in terminal cushions, bounded by a few scales. Archegoniophores terminal on main thalli. Spores with large, tuberculate areoles on distal face ............................................................................... 2. M. hegewaldii

1. Mannia californica (Gottsche ex Underw.) Wheeler, Bryologist 37: 88. 1935. Grimaldia californica Gottsche ex Underw., Bot. Gaz. 13: 114. 1888. Type. U. S. A. California: Yosemite Valley, Bridal Veil Falls, Jun 1866, Bolander s.n. (NY).

Figs. 122, 123

Thallus 1.5-2.5(-4) mm wide, greyish green with purplish margins. Branching dichotomous, ventral, and by apical adventitious branches. Epidermal cells thick-walled. Epidermal pores with 2-3 rings of 5-7(-9) cells, radial walls thickened. Air chambers in 2-5 layers. Ventral scales blackish purple, with oil-cells and marginal papillae, with 1-2 acuminate-lanceolate appendages. Monoecious. Antheridia in irregular groups on dorsal side of thallus, without scales. Archegonia in cushions on

Bischler et al. 144 short ventral branches, bounded by purplish scales. Archegoniophore receptacle hemispherical, 2roughened dorsally by projecting epidermal pores, shallowly 3-5-lobed. Spores 55-75 µm diameter, dark violet-purple, with rounded and tuberculate protuberances on distal face. Elaters with 2-4 helical bands, minutely roughened. Distribution and ecology. Mannia californica is known from the U.S. A. (Arizona, Arkansas, California, North Carolina, Tennessee, Texas) and two localities in Mexico (Nuevo León, San Luis Potosí). The record from Ceralvo Is. (Crum & Steere, 1959) belongs to Targionia hypophylla. The species has been collected from soil over acidic rocks, rock outcrops or in rock crevices, and roadsides, in exposed situations or under open forest, from low elevation to 3300 m. Specimens examined. MEXICO. NUEVO LEÓN: Cerro Potosi, Mc Gregor & Del Rosario 329 (NY). SAN LUIS POTOSÍ: 30 mi. E San Luis Potosí, Hwy 86, Mc Gregor & Del Rosario 629 (NY). In sterile condition, the species can be confused with Reboulia hemisphaerica. However, its thallus margins are not sinuose and the purplish thallus border and the scale appendages are wider.

2. Mannia hegewaldii Bischl. spec. nov. Type. Peru. La Libertad: Otuzco, Huancamarca, Quebrada Hornillo, Hegewald 5168 (holotype, MO; isotype, JE).

Figs. 123, 124, 160: A

Frons 2.5-3.5 mm lata, ecostata, non areolata, marginibus crispatis, purpureis. Cellulae epidermidis parietibus tenuibus. Porae epidermidis 2-3 annulis concentricis circumdatae. Cavernae 3-stratosae. Cellulae oleiferae brunneae. Squamae ventrales purpureae marginibus hyalinis papillosis, appendiculis 2-3, 6-7 cellulis latis. Monoecius. Antheridia et archegonia in pulvinis, squamis minutis purpureis circumdatis. Sporophyta in receptaculis hemisphaericis stipitatis 4-lobatis. Incolucrum cupuliforme. Capsula parietibus tenuibus. Sporae 68-76 µm diameter, brunneae, alveolatae. Elateres 2-3-spiri. Thallus 2.5-3.5 mm wide, light green, smooth, not areolate, with broad, purplish, crisped borders. Branching dichotomous and ventral. Epidermal cells thin-walled. Epidermal pores with 2-3 rings of 6-7 cells, radial walls thin. Air chambers in 3 layers. Basal tissue with brown oil-cells. Ventral scales purplish with hyaline margins, with oil-cells and marginal papillae, with 1-2 acuminate, smalllanceolate appendages, 6-7 cells wide. Monoecious. Antheridia and archegonia in terminal cushions, bounded by few, minute, purplish or light red scales. Archegoniophore stalk sometimes very long (2030 mm), naked except apically with a few, purplish scales; receptacle 17.5 mm diameter, broadhemispherical, slightly asymmetrical and roughened dorsally by projecting epidermal pores, shallowly 4-lobed. Involucres cup-shaped, hyaline, margin entire to crenulate. Capsules with yellow-brown, hardly thickened walls. Spores 68-76 µm diameter, brown, with yellow wing, distal face with large, finely tuberculate areoles. Elaters with 2-3 helical bands. Gametophytic chromosome number unknown.

Bischler et al. 145 Distribution and ecology. Only two specimens are known, both from Peru. Mannia hegewaldii was collected at high elevations (2690-3100 m), growing on soil or rock. Specimens examined. PERU. LA LIBERTAD: Prov Otuzco, Usquil, on road to Otuzco, Hegewald & Hegewald 5286 (JE). Mannia hegewaldii is closely related to M. triandra and has been attributed by Grolle (in litt.) to the latter species. However, a number of differences between the two species can be noted. The thallus surface in M. triandra is areolate, the thallus margins are hyaline, and the appendages of the ventral scales are narrower (maximally 5 cells wide). Moreover, the spores have narrower and more numerous areoles, which are coarsely (not finely) tuberculate. Since only few specimens are available, little can be said about morphological variation in M. hegewaldii. Additional, preferably living material is needed for a better understanding of the variation in this new species.

14. PLAGIOCHASMA Lehm. et Lindenb., in Lehmann, Nov. stirp. pug. 4: 13. 1832 (nom. cons.). Type. Plagiochasma cordatum Lehm. et Lindenb. (typus cons.). From plagios = lateral, chasma = opening, referring to the bilabiate involucres with lateral opening.

Fig. 125

Aytonia J.R. & G.Forst., Char. gen. pl. ed. 1 (folio) 74. 1775, nom. rejic. Type. Aytonia rupestris J.R. & G.Forst. Ruppinia L.f., Acta Med. Suec. 1: 204. 1783. nom rej. Type. Ruppinia rupestris L. f. Thallus usually 3-5 mm wide, green, often tinged with purple, ventral side from purplish to nearly black. Branching dichotomous, ventral and apical; activity of adventitious meristems beneath the apex may result in successive thallus constrictions (apical adventitious branches). Epidermal pores elevated or not elevated above epidermis, simple, with 1-4 rings of 4-10 cells, inner ring of collapsed cells present or absent, radial walls thin or thickened. Assimilatory tissue in several layers of empty air chambers. Ventral tissue often partly purplish, with numerous oil-cells, without mucilage cavities or sclerotic cells. Ventral scales in two rows, with oil-cells, with or without marginal papillae, with 1-4 lanceolate or ovate appendages, constricted or not constricted basally. Monoecious. Antheridia in cushions bounded by scales, dorsal or at bottom of apical adventitious branches. Archegonia 2-4 per archegonial cavity, in cushions bounded by scales, usually dorsal, sometimes at base of apical adventitious branches. Archegoniophore stalk dorsal (not terminal), without assimilatory strip or rhizoid furrow, with scales at top and bottom; receptacle of mature archegoniophore 1-4(-9)-lobed, with compound pores on dorsal side. Involucres bilabiate, opening sidewards. Calyptrae 2-3-layered. Sporophytes one per involucre, with bulbous foot and hardly elongating seta. Capsules globose, opening by a pluristratose, irregularly fragmenting lid to 1/3 of capsule length, walls without annular thickenings. Spore : elater ratio 4 : 1. Spores 2800-3200 per capsule, 52-118 µm diameter, yellow or

Bischler et al. 146 brown; proximal and distal faces with more or less complete, tuberculate areoles and tuberculate ridges, trilete scar distinct. Elaters with 2-3 helical bands, or entirely thickened. Gametophytic chromosome number n = 9 or 18. A genus of 16 species, distributed in warm areas. In the New World, nine species are known; all except one (the widespread P. rupestre) have limited distributions. The species are drought-tolerant and the thalli curl up laterally when dry, only the blackish ventral side and the scales remaining visible. Material dried for 2-3 years can resume growth within a few hours. The species colonize mainly rock or wall crevices, or soil over rocks, and are nearly indifferent to pH but prefer calcareous substrates. They are found in steppes, savannas, under shrub vegetation, and in open dry forests, in tropical continental areas from 1000 m to 4000 m, on islands from sea level to 2300 m.

KEY TO THE SUBGENERA OF PLAGIOCHASMA 1. Epidermal pores 8-66 per mm2, 12-35 µm diameter, bounded by 2-4 concentric rings of (5-)6-9(10) cells. Epidermis smooth or finely granulose. Ventral scales often with papillae; margins of appendages differentiated or irregularly toothed .................................14a. subgen. Plagiochasma 1. Epidermal pores 130-280 per mm2, 5-6 µm diameter, bounded by a single ring of 4-6 cells. Epidermis covered irregularly by hydrophobic material. Ventral scales without papillae; margins of appendages entire, margins not differentiated ............... .........................14b. subgen. Micropylum

14a. PLAGIOCHASMA subgen. PLAGIOCHASMA Thallus often with successive, apical adventitious branches. Epidermis smooth or finely granulose. Epidermal pores 8-66 per mm2, raised above epidermis, 12-35 µm diameter, bounded by a hyaline ring and 2-4 concentric rings of (5-)6-9(-10) cells. Scales of thallus and receptacles with papillae and with 1-3 rows of smaller, or sinuose, or obliquely oriented cells on margins, or with 1- to pluricellular teeth. Male and female receptacles often at base of apical adventitious branches. The subgenus is distributed from 40° N to 30° S. Species diversity is highest north of the equator, in the New World in the southern United States and Central America. Eight species are known from the Neotropics.

KEY TO THE SPECIES OF SUBGEN. PLAGIOCHASMA

Bischler et al. 147 1. Ventral scales with 1(-2) appendages, of length : width ratio = 0.8-2 : 1, usually constricted and/or plicate basally................................................................................................. ....................................2 2. Appendages of ventral scales ovate or rounded, length : width ratio = 0.8-1.3 : 1, 22-26 cells wide, widest near the middle; margins with several rows of sinuose cells..................................... ..............................................................................................................................2. P. crenulatum 2. Appendages of ventral scales triangular, length : width ratio = 1.4-2 : 1, less than 22 cells wide, widest near the base; marginal cells not sinuose.................................................... .......................3 3. Surface of epidermis finely granulose. Epidermal pores with 2(-3) rings of 5-8 cells. Appendages of ventral scales hyaline. Spores 70-82 µm diam .................. ..........8. P. wrightii 3. Surface of epidermis smooth. Epidermal pores with 3(-4) rings of 6-7(-8) cells. Appendages of ventral scales usually not hyaline. Spores 82-94 µm................................... ........................4 4. Epidermal cells (24-)28-35 µm long. Radial walls of epidermal pores strongly thickened. Appendages of ventral scales 15-22 cells wide, often with some pluricellular teeth on margins. Thallus usually without apical adventitious branches..............7. P. muenchianum 4. Epidermal cells 35-47µm long. Radial walls of epidermal pores hardly thickened. Appendages of ventral scales 8-10 cells wide, with entire margins. Thallus often with apical adventitious branches..................................................................................5. P. jamaicense 1. Ventral scales with (1-)2-4 appendages, of length : width ratio = 2-5 : 1, hardly constricted basally ........................................................................................................................................................... 5 5. Epidermal cells (28-)35--40 µm long, walls with small trigones. Ventral scales with 2 hardly acuminate appendages, as wide in the middle as basally, length : width ratio = 2-2.5 : 1, ending in single cell apically. Elaters without helical bands. Spores 73-82 µm diameter. ............................ ............................................................................................................................4. P. intermedium 5. Epidermal cells 28-35 µm long, walls with large, nodulose trigones. Ventral scales usually with 3 acuminate appendages, widest basally, length : width ratio = 3-5 : 1, with row of 1-5 cells apically. Elaters with helical bands. Spores 82-118 µm diameter .............................. ..................6 6. Surface of epidermis smooth. Elaters with interrupted helical bands ..................6. P. landii 6. Surface of epidermis finely granulose. Elaters with complete helical bands ................ ......7 7. Thallus 3-4 mm wide. Appendages of ventral scales with row of 3-4 cells apically, 10-14 cells wide basally; marginal cells not sinuose ............................... ...............3. P. cuneatum

Bischler et al. 148 7. Thallus 3.7-8 mm wide. Appendages of ventral scales with row of 1-2 cells apically, 5-7 cells wide basally; marginal cells sinuose ...............................................1. P. argentinicum

1. Plagiochasma argentinicum Bischl., Rev. Bryol. Lichénol. 45: 301. 1979. Type. Argentina. Catamarca: S. Ambato, Volk 6/669 (holotype, BA; isotype, PC). Figs. 1: E, 126: A-E, 127, 136: E-F Thallus 3.7-8 mm wide, yellowish or olive green, with narrow, purplish margins; apical adventitious branches few. Epidermis finely granulose on surface, epidermal cells 28-35 x 24-28 µm, with medium sized to large, sometimes nodulose trigones. Epidermal pores 9-20 per mm2, raised above epidermis, 12-21 µm diameter, bounded by 2-3 concentric rings of 6-8 cells, radial walls thickened. Ventral scales purplish, margin of 2-3 rows of light red or hyaline, sinuose cells; appendages 2-4, narrowly triangular and acuminate, purplish or light red, with row of 1-2 cells apically, widest basally (5-7 cells), not constricted, length : width ratio 3.3-4.5 : 1; margins bordered by sinuose cells, sometimes with a few teeth, without papillae. Male receptacles dorsal or at base of apical adventitious branch, rounded or reniform, with 20-30 antheridia. Scales of male and female receptacles purplish, or hyaline, or hyaline and tinged with purple, acute or apiculate, with some papillae on margins. Archegoniophores dorsal, stalk 0.5-1.3 mm long, with thickened, dark red epidermal cells. Involucres 1-3, light brown or purplish. Spores 106-118 µm diameter, light or dark brown, regularly areolate on both faces, areoles and ridges tuberculate. Elaters with 3-5 helical bands. Gametophytic chromosome number unknown. Distribution and ecology. Collected twice in northern Argentina (Catamarca, Córdoba, 30-35° S), growing in crevices of calcareous rocks in dry habitats, at 1000 m. Specimens examined. ARGENTINA. CATAMARCA: Type. CÓRDOBA: Calamuchita, San Agustín, Hosseus 163 (JE) Plagiochasma argentinicum is easily recognised by the finely granulose epidermis and by the ventral scales being bordered by sinuose cells and having 2-4 narrow appendages.

2. Plagiochasma crenulatum Gottsche, Mexik. Leverm. 266. 1863 (Kongel. Danske Selsk. Naturvidensk. Math. Afh., ser. 5, 6: 362. 1868). Aytonia crenulata (Gottsche) Underw., Bot. Gaz. 20: 66. 1895. Type. Mexico. Veracruz: Orizaba, Müller s.n. (holotype, STR p.p.; isotypes, G p.p., YU).

Figs. 126: G-N, 127, 136: M

Thallus 5.3-6.7 mm wide, light or yellowish green, with red or purplish, narrow margins. Apical

Bischler et al. 149 adventitious branches infrequent. Epidermis smooth, epidermal cells 31-40 x 24-28 µm, walls usually with large, nodulose trigones. Epidermal pores 27-66 per mm2, slightly raised above epidermis, 21-24 µm diameter, bounded by 2-3 concentric rings of 6-7(-8) cells with slightly thickened radial walls. Ventral scales purplish, bordered by 2-3 rows of light red or hyaline, sinuose cells; appendage 1, rarely 2, ovate or rounded, hyaline, sometimes light red or yellowish in the central part, length : width ratio 0.8-1.3 : 1; obtuse, acute or apiculate apically, largest in the middle, 22-26 cells wide, constricted and plicate basally, margins crenulate or toothed, with several rows of sinuose cells, some with papillae. Male receptacles dorsal, usually reniform, with 20-25 antheridia. Scales of male and female receptacles hyaline or partly purplish, acute or apiculate, with crenulate or remotely toothed margins. Archegoniophores dorsal, stalk 0.8-5 mm long, with thickened, red or brown epidermal cells. Involucres 1-3, yellow, purplish or brown. Spores 70-78 µm diameter, dark yellow or brown, areolate; areoles and ridges tuberculate. Elaters with 2-4 helical bands. Gametophytic chromosome number n = 18. Distribution and ecology. A species found in scattered localities (10°-33° N), quite common in southern Mexico. Known from central Alabama, southern Arizona, Mexico (Chiapas (Boreal, 1992), Hidalgo, Morelos, Nuevo León, Oaxaca, Puebla, San Luis Potosí, Sonar, Veracruz), Guatemala, Honduras and Costa Rica (Cole, 1984), growing in rock and wall crevices, in dry and warm areas, from 650 to 2000 m. Selected neotropical specimens examined. MEXICO. HIDALGO: Chapulhuacan, Sharp 1503a (MEXU, US). MORELOS: near Cuernavaca, Pringle 15319 (MEXU). NUEVO LEÓN: Montemorelos, near Monterrey, Lacas 465 (F); Santiago, Horsetail Fall, Frye & Frye 2668 (F, G, NICH, NY, S, US); Rio Valle Alto, Spessard 435 (MEXU). OAXACA: without locality, Galeotti 6890 (G). PUEBLA: Mun. Xochitlan, Arroyo Atena, 15 km SE Zapotitlan, Grimes 2765 (MEXU). QUERÉTARO: : Sierra Gorda, Chuveje Cascades, W of Jalpan, Long & Delgadillo 29721. SAN LUIS POTOSÍ: Tamazunchale, Frye & Frye 2977 (F, G, NY, S); above Xilitla, Sharp 5849 (MEXU). SONORA: Near Alamos, Rose et al. 13121 (BM, NY, US, YU); Huehnerache, Hartman 307 (FH, YU). VERACRUZ: Type; Orizaba, Barnes & Land 685 (F, FH, NY, YU); Metlac Gorge, Barnes & Land 640 (F, NY, YU), 659 (F, NY, YU); Córdoba, Conzatti & Gonzalez 1130 (FH, US, YU); Tuxpango, Sharp 1228 (MEXU, US); river gorge below Teocelo, Long 29695, 29697 (E). GUATEMALA. Alta Verapáz, Coban, Richards & Williams 2791A (F); Izabal, along Río Frio, Steyermark 41620 (F). HONDURAS. Comayagua, vicinity of Siguatepeque, Yuncker et al. 6461 (Hb. Fulford), Standley & Chacon 6272 (F). Plagiochasma crenulatum is a polyploid species. It is distinguished from other neotropical species by the wide, rounded or ovate appendages of the ventral scales.

Bischler et al. 150

3. Plagiochasma cuneatum A.Evans, Amer. J. Bot. 19: 627. 1932. Type. U. S. A. Texas: Hamilton's Pool, Feb 1931, McAllister s.n. (holotype, YU).

Figs. 128: A-G, 130

Plagiochasma elongatum Lindenb. & Gottsche var. ß leucolepis Gottsche, Lindenb. & Nees, Syn. hepat. 519. 1846. Type. Mexico. Veracruz: Jalapá, Deppe & Schiede 109 (isotypes, BM, G). Thallus 3-4 mm wide, light or yellowish green with narrow, purplish margin; apical adventitious branches frequent, often short. Epidermis finely granulose; epidermal cells 28-35 x 21-24 µm, with medium sized or large, often nodulose trigones. Epidermal pores 15-20 per mm2, raised above epidermis, usually 28-35 µm diameter, bounded by (2-)3(-4) concentric rings of (7-)8(-9) cells with strongly thickened radial walls. Ventral scales purplish or dark red, with border of 1-2 rows of light red or hyaline, sinuose cells; appendages usually 2-3, narrowly triangular, apex acuminate, with row of 3-4 cells apically, hyaline, contrasting with the dark scale body, widest basally, 10-14 cells wide, not constricted or plicate, length : width ratio 4-5 : 1, margins not differentiated, sometimes with some teeth and papillae. Male receptacle dorsal or at base of apical adventitious branch, usually reniform, with 20-30 antheridia. Scales of male and female receptacles hyaline or purplish, or partly tinged with purple, acuminate, with row of 3-5 cells apically, margins with scattered papillae. Archegoniophores dorsal or at base of apical adventitious branch, stalk 2.3-4 mm long, with thickened, dark red epidermal cells. Involucres 2-3, light brown or purplish. Spores 94-118 µm diameter, light or dark brown, both faces areolate, areoles and ridges tuberculate. Elaters with 2-3 helical bands. Gametophytic chromosome number n = 9. Distribution and ecology. The range of the species extends from 30° N to 15° N, from the U. S. A. (Texas), Mexico (D. F., Mexico, Nuevo León, Veracruz), Guatemala to Haiti. It is infrequent everywhere and has been collected in rock crevices of calcareous rocks or sandstone, in dry, warm climatic areas, from sea level to 2300 m. Specimens examined. MEXICO. D. F.: Desierto de los Leones, walls of El Convento, 1944, Ruiz Oronoz s.n. (MEXU); Jardín Botanico Exterior, Whittemore 4098 (MEXU). MEXICO: Barranca Malinaltenango, Pócs 9552B (MEXU). NUEVO LEÓN: 19 mi. Linares, McGregor & Del Rosario 220 (NY); 7 mi. W Iturbide, Hwy 60, Mc Gregor & Del Rosario 30 (NY).VERACRUZ: Type; type of P. elongatum var. ß leucolepis; Orizaba, Farlow 17 (NY); Jalapa, Deppe & Schiede s.n. (FH, NY, US). GUATEMALA. Zaculco, Volk 6/717 (PC). HAITI. Near Jean Rabel, Leonard & Leonard 12848 (FH, JE, NY, US); Dept. de L'Ouest, Forêt des Pins (Shada Station), Imshaug & Wetmore 2952 (NY). Plagiochasma cuneatum is easily distinguished by the finely granulose epidermis and by its dark

Bischler et al. 151 purplish ventral scales with long, acuminate, hyaline appendages.

4. Plagiochasma intermedium Lindenb. & Gottsche, in Gottsche, Lindenb. & Nees, Syn. hepat. 513. 1846. Ruppinia intermedia (Lindenb. & Gottsche) Trevis., Mem. Reale Ist. Lombardo Sci. Ser. 3, Cl. Sci. Mat. 4: 437. 1877. Aitonia intermedia (Lindenb. & Gottsche) Kuntze, Revis. gen. pl. 1: 143. 1891. Aytonia intermedia (Lindenb. & Gottsche) Underw., Bot. Gaz. 20: 66. 1895. Type (Evans, 1915. Mexico.Veracruz: Hacienda de Fovo, May 1841, Liebmann 532b (lectotype, W; isolectotype, S).

Figs. 128: H-N, 130, 136: H-I, L

Thallus 3.7-5 mm wide, yellowish green, often tinged with purple, with narrow, purplish margins; apical adventitious branches frequent. Epidermis smooth; epidermal cells (28-)35-40 x 21-24 µm, with small to medium sized trigones. Epidermal pores 46-50 per mm2, raised above epidermis, 24-28 µm diameter, bounded by (2-)3-4 concentric rings of 8-9 cells, radial walls strongly thickened. Ventral scales purplish, with border of 1-2 rows of light red or hyaline, sinuose cells; appendages 2(3), triangular, apex acute, with single cell or papilla apically, light red or purplish, as wide basally as in the centre, 10-12 cells wide, not constricted but horizontally plicate basally, length : width ratio 22.5 : 1, margins partly bordered by smaller cells, without papillae. Male receptacle usually at base of apical adventitious branch, rarely dorsal, usually reniform, with approx. 30 antheridia. Scales of male and female receptacles hyaline or partly tinged with purple, acute, with single cell apically, margins with scattered papillae. Archegoniophores usually dorsal, rarely at base of apical adventitious branch, stalk 0.5-5.7 mm long, with thickened, dark purplish epidermal cells. Involucres 2-4, yellow, purplish or brown. Spores 73-82 µm diameter, light or dark brown, with complete or incomplete areoles on both faces, ridges tuberculate. Elaters without helical bands. Gametophytic chromosome number n = 9. Distribution and ecology. The geographical range of P. intermedium is mainly neotropical and extends from 28° N to 10° N, including Mexico (Chiapas, Chihuahua, D. F., Jalisco, Mexico, Oaxaca, Puebla, San Luis Potosí, Veracruz), Guatemala, Costa Rica, and Venezuela (Aragua). The species has also been recorded from North Carolina (Schuster, 1992b). Records from other continents are erroneous and apply to other species. P. intermedium is most common in central Mexico and Guatemala, rare elsewhere, and has been collected in dry, warm climatic areas in rock and wall crevices (lava or limestone), in exposed habitats or in open forests, between 800 m and 2300 m. Selected neotropical specimens examined. MEXICO. CHIAPAS: 14 km S San Cristóbal, Düll A 125 (NY). CHIHUAHUA: National Park, Basaseachi waterfall, Whittemore 2863 (MEXU). D. F.: Desierta Vieja forest, 1865, E.B. s.n. (PC); Botanical Garden, 1979, Jovet-Ast s.n. (PC), Schwab SN 67 (F, JE, NY, PC, YU), Standinger s.n. (Hb. Volk); Desierto de los Leones, 1945, Ruiz Oronoz s.n. (MEXU); Pedregal de San Angel, Amable 1548 (FH, G, US). JALISCO: Below Experiencia, Barnes &

Bischler et al. 152 Land 146 (G, PC, YU); Etzatlan, Barnes & Land 261 (F, JE, NY, YU); Guadalajara, Barnes & Land 136 (BM, F, NY, W, YU); San Sebastian, Mexia 1363 (JE, NY, PC); Guadalajara, Pringle 700 (NY). MEXICO: Tlalpan, Amable 1543 (US, YU); Cerro de Guadalupe, Maury 3227 (F, G, NY, PC, YU). OAXACA: Rio San Cristóbal, Düll A 64 (F, JE, NY, YU). PUEBLA: Santa Barbara, Arsène s.n. (NY, PC); Tezuitlan, Barnes & Land 538 (F, JE, NY, YU); Chinantla, Lieman 438b (BM, US, W). SAN LUIS POTOSÍ: 19 mi. E San Luis Potosí, Hwy 86, Mc Gregor & Del Rosario 688 (NY); 20 mi. E San Luis Potosí, McGregor & Del Rosario 622 (G, NY, STR). VERACRUZ: Type; Orizaba, Mohr 30 (G), Müller s.n. (G, STR, US); Huatusco, Mohr 17 (FH, G, PC), Mohr s.n. (G). GUATEMALA. Dep. Suchiate, Finca El Naranjo, Svihla 45-6189 (Hb. Fulford); Chimaltenango, 5 km W of Patzun, Richards & Williams 3018 (F); Escuintla below Las Lajas, Standley 64856 (F, NY); Solola, Panajachel, Richards & Williams 3051 (F); Sta Rosa, E of Cuilapa, Standley 78282 (F); Sta Rosa, Guachipilín, Heyde & Lux 6292 (BM, F, FH, G, NY, YU); Zaculco, Volk 6/0718 (JE). COSTA RICA. N of Cartago, Standley & Valerio 49396 (NY, US, YU); S. José, near Las Pavas, Standley 36060 (YU). VENEZUELA. ARAGUA: Parque Nacional Pittier, above Las Delicias, en route to Choroni Pass, Steyermark 100511 (NY, PC). Plagiochasma intermedium is the sole neotropical species having elaters without helical bands. Additionally, shape and structure of the appendages of the ventral scales separate the species from all other neotropical representatives of the genus.

5. Plagiochasma jamaicense (Haynes) A.Evans, Bull. Torrey Bot. Club 42: 292. 1915. Aytonia jamaicensis Haynes, Bull. Torrey Bot. Club 34: 58. 1907. Type. Jamaica. Chestervale, 1000 m, Underwood 1173 (holotype, NY; isotype, YU).

Figs. 129: A-G, 130

Thallus 4.3-6.7 mm wide, light or yellowish green, with narrow, purplish margins; apical adventitious branches frequent. Epidermis smooth; epidermal cells 35-47 x 24-28 µm, with medium sized or large, sometimes nodulose trigones. Epidermal pores 10-35 per mm2, raised above epidermis, 14-17 µm diameter, bounded by 3(-4) concentric rings of (6-)8 cells, radial walls hardly thickened. Ventral scales purplish, with border of 2-3 rows of light red or hyaline, sinuose cells; appendage 1(-2), triangular, apex acute, with row of (1-)2-3 cells apically, light red or purplish, widest basally , 8-10 cells wide, constricted and horizontally plicate, length : width ratio 1.5-2 : 1; margins bordered by 1-2 rows of smaller cells, some with papillae. Male receptacle at base of apical adventitious branches, reniform, with 20-30 antheridia. Scales of male and female receptacles hyaline or purplish, or partly tinged with purple, acute, with row of 2-3 cells apically, margins with scattered

Bischler et al. 153 papillae. Archegoniophores dorsal, stalk 4-6 mm long, with slightly thickened, dark purplish epidermal cells. Involucres 1-4, yellowish or tinged with purple. Spores 82-94 µm diameter, dark yellow or brown, with complete areoles on both faces, ridges tuberculate. Elaters with 2(-3) helical bands. Gametophytic chromosome number unknown. Distribution and ecology. A rare neotropical species known from only two localities (18-20° N), one in Jamaica (1000 m) and one in Mexico (3300 m), both in relatively dry areas. Specimens examined. JAMAICA. Type; Chestervale, Underwood 1177 (NY). MEXICO. D. F.: La Cima, Pringle 10682 (G, YU). Plagiochasma jamaicense is related to P. wrightii and can be distinguished from the latter species by the larger thallus, finely granulose epidermis, longer epidermal cells, and smaller epidermal pores, less raised above the thallus surface. Moreover, the appendages of the scales are smaller and light red or purplish in colour.

6. Plagiochasma landii A.Evans, Bull. Torrey Bot. Club 42: 298. 1915. Type. Mexico. Morelos: West of the city of Cuernavaca, Barnes & Land 466 (holotype, YU; isotypes, FH, NY). Figs. 129: H-P, 134, 136: K Thallus 4.4-5 mm wide, yellowish green, often partly tinged with purple, with narrow, purplish margins; apical adventitious branches frequent, often long. Epidermis smooth; epidermal cells 33-35 x (21-)24-28 µm, with medium sized or large, sometimes nodulose trigones. Epidermal pores 8-20 per mm2, raised above epidermis, 14-24 µm diameter, bounded by 2-3(-4) concentric rings of 6- 8(-10) cells, radial walls strongly thickened. Scales purplish or orange, with border of 1-2 rows of light red, sinuose cells; appendages (1-)2-3, narrowly triangular, apex acuminate and acute, with row of 1-2 cells apically, light red or purplish, rarely hyaline, widest basally, 12-15 cells wide, not constricted or plicate, length : width ratio 3-4 : 1; margins often with some teeth, without papillae. Male receptacle usually at base of apical adventitious branches, rounded or reniform, with 20-30 antheridia. Scales of male and female receptacles hyaline, or partly tinged with purple, acute, with row of 2-3 cells apically, margins often toothed and with scattered papillae. Archegoniophores dorsal, stalk 0.2-0.4 mm long, with thickened, dark purplish epidermal cells. Involucres 2-3, light brown or purplish. Spores 82-100 µm diameter, light or dark brown, with complete or incomplete areoles on both faces, ridges more or less densely tuberculate. Elaters with 2-4, sometimes interrupted helical bands. Gametophytic chromosome number unknown. Distribution and ecology. Plagiochasma landii is an uncommon New World species distributed from 23°5' N to 11° N, known from southern U. S. A (Texas), Mexico (Chiapas (Bourell, 1992), D.

Bischler et al. 154 F., Morelos, San Luis Potosí), Guatemala, and Costa Rica. It has been collected in rock crevices in exposed or partly shaded habitats, in relatively dry areas between 1500 and 2850 m. Neotropical specimens examined. MEXICO. D. F.: Desierto de los Leones, Woronow 4 (FH, G, S, US); El Desierto, Amable 1649, 1650 (NY, US, YU), 1652 (F, US); Parque Nacional Desierto de los Leones, near Exconventa, Long 29589 (E). MORELOS: Type; near Cuernavaca, Pringle 10669 (FH, G, S, US, YU). SAN LUIS POTOSÍ: without locality, Schaffner 3 (NY, US, YU). GUATEMALA. Guezaltenango, near Alonzo, San Juan Ostuncalco, Standley 84178 (F). COSTA RICA. Cartago, Irazú, Stork s.n. (US). Although considerable variation is observed in the shape and structure of the scale appendages, in elater structure, and in spore wall ornamentation, this species cannot be split into morphologically well-characterized groups. Plagiochasma landii is most closely related to P. cuneatum and P. argentinicum and is separated from these two species by the smooth epidermis.

7. Plagiochasma muenchianum Steph., Sp. hepat. 6: 9. 1917 (not “münchianum”, fide Art. 60.6, ICBN). Type. Mexico. Chiapas: San Cristóbal, 2100 m, Münch 5607 (holotype, FI; isotypes, FH, G, NY).

Figs. 131: A-G, 132

Thallus 6-7.3 mm wide, yellowish green, sometimes partly tinged with purple, with purplish margins; apical adventitious branches infrequent. Epidermis smooth; epidermal cells (24-)28-35 x 2124 µm, usually with large, sometimes nodulose trigones. Epidermal pores 8-15 per mm2, slightly raised above epidermis, 24 µm diameter, bounded by 3(-4) concentric rings of 6-7(-8) cells, radial walls thickened. Ventral scales purplish, with border of 1-2 rows of light red or hyaline, sinuose cells; appendage 1(-2), widely triangular or ovate, apex acute, with 1-2 cells apically, light red or hyaline, widest basally, 15-22 cells wide, constricted and horizontally plicate, length : width ratio 1.4-1.6 : 1; margins with 1-2 rows of smaller cells, some with papillae, sometimes with 1-2 large, pluricellular teeth. Male receptacle dorsal, rounded to reniform, with 25 antheridia. Scales of male and female receptacles hyaline, or partly tinged with purple, acute, with row of 1-3 cells apically, margins with scattered teeth and papillae. Archegoniophores dorsal, stalk 1-2 mm long, with slightly thickened, dark purplish epidermal cells. Involucres 2-3, purplish. Spores 82-94 µm diameter, brown, with complete areoles on both faces, ridges tuberculate. Elaters with 2(-3) helical bands. Gametophytic chromosome number unknown. Distribution and ecology. This species is known from a single locality in Mexico (Chiapas, 17° N), growing in the town of San Cristóbal (2100 m) on a wall of calcareous mortar, in a relatively dry and warm environment.

Bischler et al. 155 Specimens examined. MEXICO. CHIAPAS: Type. Plagiochasma münchianum is most closely related to P. wrightii and P. jamaicense. In P. wrightii, the thallus is narrower, the epidermis finely granulose, and the pores smaller. In P. jamaicense, the appendages of the scales are smaller and narrower, without teeth, and the thallus produces numerous apical adventitious branches.

8. Plagiochasma wrightii Sull., in A.Gray, Manual, ed. 2, 688. 1856. Aitonia wrightii (Sull.) Underw., Bull. Illinois State Lab. Nat. Hist. 2: 43. 1884. Type (Evans, 1915). U. S. A. Texas: San Marco, 1848, Wright 9 (lectotype, FH; isolectotype, NY).

Figs. 131: H-O, 132, 136: G

Plagiochasma crenulatum Gottsche var. ß minus Gottsche, Mexik. Leverm. 266. 1863 (Kongel. Danske Vidensk. Selsk. Naturvidensk. Math. Afh., ser. 5, 6: 362. 1868). Type. Mexico. Oaxaca: Chinantla, Liebmann 519 (holotype, W; isotype, NY). Thallus 3.3-4.2 mm wide, whitish, yellowish or brownish, with purplish margins; apical adventitious branches frequent. Epidermis finely granulose; epidermal cells 26-31 x 21-24 µm, with large, often nodulose trigones. Epidermal pores 27-66 per mm2, hardly raised above epidermis, 17-24 µm diameter, bounded by 2(-3) concentric rings of 5-8 cells, radial walls hardly thickened. Ventral scales purplish or orange, with border of 1-2 rows of light red or hyaline, sinuose cells; appendage 1(2), triangular, apex acute, with row of (1-)2-3 cells apically, hyaline, rarely light red or purplish, widest basally, 12-17 cells wide, slightly constricted and horizontally plicate, length : width ratio 1.51.9 : 1; margins not differentiated or partly bordered by 1-2 rows of smaller cells, without papillae. Male receptacle at base of apical adventitious branch, rarely dorsal, usually reniform, with 20-30 antheridia. Scales of male and female receptacles hyaline or purplish, or partly tinged with purple, acute, with 1-2 cells apically, margins with scattered papillae. Archegoniophores dorsal, stalk 0.2-4 mm long, with slightly thickened, yellow, brown or dark purplish epidermal cells. Involucres 1-4, yellow or purplish. Spores 70-82 µm diameter, light or dark brown, with complete areoles on both faces, ridges tuberculate. Elaters with 2-3 helical bands. Gametophytic chromosome number unknown. Distribution and ecology. A New World species recorded from 38° N to 17° N. It is quite common in southern U.S.A (Arizona, Colorado, Kansas, Missouri, Nevada, New Mexico, North Carolina, Oklahoma, Texas) and Mexico (Chiapas (Bourell, 1992), Chihuahua, Coahuila, D. F., Durango, Guerrero, Hidalgo, Mexico, Michoacán, Nuevo León, Oaxaca, Puebla, San Luis Potosí, Sonora, Tlaxcala, Veracruz, Zacatecas). Plagiochasma wrightii has been collected on damp sandy soil in rock crevices (limestone, sandstone, granite), from 800 m to 3000 m, in dry but warm climatic areas, in exposed habitats or rather open vegetation.

Bischler et al. 156 Selected neotropical specimens examined. MEXICO. CHIHUAHUA: 55 mi. N of Chihuahua, at km 87 on road to El Paso, Wittemore et al. 2905 (MEXU). COHAHUILA: 6 mi. SE of Saltillo, Palmer 423 (FH, W, YU). D. F.: Taltlan, Amable 1545 (US); El Desierto, Amable 1650 (F, US). DURANGO: Durango, near Santiago, Palmer 58 (FH, NY, US, YU). GUERRERO: Taxco, 1947, Ruiz Oronoz s.n. (MEXU). HIDALGO: Jacala, Chase 7405 (US); N of Atotonilco, Schwab SN 120 (JE, NY, PC). MEXICO: Barranca de Malitenango, Pócs 9552A (MEXU). MICHOACÁN: Morelia, Los Tetecos, Arsène s.n. (F, NY, US); NUEVO LEÓN: 19 mi. W Linares, Mc Gregor & Del Rosario 202 (NY, US); 21 mi. W Linares, Hwy 60, Mc Gregor & Del Rosario 182 (NY). OAXACA: Type of P. crenulatum var. ß minus ; E Tamazulapan, Schwab SN 132 (JE, PC); Yavesca, Liebmann 256 (W). QUERÉTARO: Peña de Bernal, Long & Delgadillo 29698 (E); below La Florida, W of Conca, Long & Delgadillo 29730 (E). PUEBLA: Acatlan-Oaxaca, 100 km before Oaxaca, Düll A 63 (F, JE); Santa Barbara, Nicolas 5612 (FH, G, PC). SAN LUIS POTOSÍ: 4 mi. E Ciudad del Maíz, McGregor & Del Rosario 787 (JE, NY, PC); San Luis Potosí, 30 mi. E, McGregor & Del Rosario 624 (NY, PC). SONORA: 18 mi. SW of Cumpas, Richards et al. 726 (F, JE); S of Jecori, Richards et al. 715 (F, PC). TLAXCALA: Rio Zahuapan near Amaxoc, Sharp 398 (MEXU). VERACRUZ: Cordoba, Sallé s.n. (NY, PC); Mts. Songolica, Berendt s.n. (F, FH, G); Haciena Jalapilla, 1948, Ruiz Oronoz s.n. (MEXU); Piedra de Agua, between Xalapa and Naolinca , Long et al. 29636 (E). ZACATECAS: Concepción del Oro, Palmer 307 (FH, NY). Plagiochasma wrightii is characterized by its single, hyaline, acute appendage of the scales. Plagiochasma münchianum and P. jamaicense have similar appendages but are separated by the smooth epidermis (granulose in P. wrightii) and by the wider thalli.

14b. PLAGIOCHASMA subgen. MICROPYLUM Bischl., Rev. Bryol. Lichénol. 43: 103. 1977. Aytonia J. R. & G. Forst., Char. gen. pl. ed. 1, 75. 1775 (nom. rej.). Ruppinia L. f., Suppl. pl. 69. 1782. Aitonia J. R. Forst., Comment. Soc. Regiae Sci. Gött. 9: 73. 1787 (orth. var.). Type. Aytonia rupestris J. R. & G. Forst. (= Ruppinia lichenoides L. f. = Plagiochasma rupestre (J. R. & G. Forst.) Steph.). From micro=small, pylum=opening, referring to the small epidermal pores. Thallus with infrequent constrictions or apical adventitious branches. Epidermis with irregular, hydrophobic, granular deposit. Epidermal pores 130-280 per mm2, not raised above epidermis, 5-6 µm in diameter, bounded by a single ring of 4-6 cells, often irregular in size, without hyaline inner ring. Scales of thallus and receptacles with undifferentiated margins, without papillae. Male and female receptacles dorsal. The subgenus is monospecific.

Bischler et al. 157

9. Plagiochasma rupestre (J.R. & G.Forst.) Steph., Sp. hepat. 1: 80. 1898. Aytonia rupestris J. R. et G. Forst., Char. gen. pl. ed. 1, 74. 1775. Ruppinia rupestris (J.R. & G.Forst.) Sw., Meth. musc. 122. 1781. Type (Bischler & Sergio, 1984). Madeira, Forster s.n. (neotype: Madeira. Funchal, Quinta do Bom Sucesso, Sérgio & Nobrega 3873, neotype, LISU; isoneotypes, BM, G, MADJ, PC). Thallus 2.3-5 mm wide, usually bluish green, sometimes whitish or partly tinged with purple, with narrow purplish margins; apical adventitious branches rare. Epidermis covered with a hydrophobic deposit; epidermal cells 31-40 x 20-24 µm, with medium sized, rarely small or large, usually not nodulose trigones. Epidermal pores (90-)130-280 per mm2, not raised above epidermis, 5-6 µm diameter, bounded by a single ring of 4-6 cells, often irregular in shape, radial walls more or less thickened. Ventral scales purplish, sometimes borders lighter in colour, of cells of the same size and shape, without papillae; appendages 1-2(-3), triangular, apex acute, with 1-2 cells, or acuminate, with row of 2-5 long cells with thickened walls apically, hyaline or light red, rarely purplish, widest basally, 10-14 cells wide, not constricted but sometimes horizontally plicate, length : width ratio 1.7-5 : 1; margins not differentiated, without papillae. Male receptacle dorsal, rounded or reniform, with 850 antheridia. Scales of male and female receptacles hyaline or partly tinged with purple, acute, with 1-5 cells apically, margins without papillae. Archegoniophores dorsal, stalk usually 2-4 mm long, with 1-3 rows of thickened, yellow, red or purplish epidermal cells. Involucres 1-3(-4), yellow or brownish, sometimes tinged with purple. Spores 70-87 µm diameter, light or dark brown, with complete areoles on both faces, ridges tuberculate. Elaters with 2-3(-4) helical bands. Gametophytic chromosome number n = 9 or 18. Distribution and ecology. Plagiochasma rupestre has a pantropical distribution, from 46° N to 40° S. The species is widespread in warm regions of the world and has been recorded from Europe, the Mediterranean area and Macaronesia, SW Asia, India, tropical and S Africa, the Mascarenes, Australia, New Zealand, and Oceania. However, it has not been collected in eastern Asia and on the tropical Asiatic islands. In the New World, it is common in the southern United States, Mexico, and warm-temperate South America. The species varies considerably, but in a similar manner in the Old World and New World populations.

KEY TO THE VARIETIES OF PLAGIOCHASMA RUPESTRE 1. Epidermal pores bounded by 4-5 cells with thin radial walls. Length : width ratio of appendages =

Bischler et al. 158 1.7-2.7 : 1; uniseriate apex consisting of row of 1-2 (-4) thin-walled, short cells..... 9a. var. rupestre 1. Epidermal pores bounded by 5-6 cells with thickened radial walls. Length : width ratio of appendages = 4.5-5 : 1; uniseriate apex consisting of row of 3-5 thick-walled, elongate cells....... .... ...................................................................................................................................9b. var. volkii 9a. Plagiochasma rupestre var. rupestre

Figs. 1: A, I, 3: A, I, 133: A-E, 135

Reboulia chlorocarpa Nees & Mont., Ann. Sci. Nat., Bot. sér. 2, 5: 70. 1836. Plagiochasma chlorocarpum (Nees & Mont.) Mont., in A.D.Orbigny, Voy. Amér. mérid. 7, Bot. 2: 59. 1839. Ruppinia chlorocarpa (Nees & Mont.) Trevis., Mem. Reale Ist. Lombardo Sci. Ser. 3, Cl. Sci. Mat. 4: 437. 1877. Aitonia chlorocarpa (Nees & Mont.) Kuntze, Revis. gen. pl. 1: 143. 1891. Type. Chile. Without locality, Bertero 1129 (holotype, PC). Plagiochasma peruvianum Nees & Mont, in Montagne, Ann. Sci. Nat., Bot. sér. 2, 9: 44. 1838. Plagiochasma boliviense Mont., Cent. 1° pl. cell. 45. 1838. Ruppinia peruviana (Nees & Mont.) Trevis., Mem. Reale Ist. Lombardo Sci. Ser. 3, Cl. Sci. Mat. 4: 437. 1877. Aitonia peruviana (Nees & Mont.) Kuntze, Revis. gen. pl. 1: 143. 1891. Type. Bolivia. La Paz: Yungas, between Chupé and Yanacaché, d'Orbigny 218 (holotype, PC; isotypes G, STR). Plagiochasma validum Bisch., Handb. bot. Termin. 2: 59, tab. 56, fig. 2753 explicatio. 1838. Ruppinia valida (Bisch.) Trevis., Mem. Reale Ist. Lombardo Sci. Ser. 3, Cl. Sci. Mat. 4: 437. 1877. Aitonia valida (Bisch.) Kuntze, Revis. gen. pl. 1: 143. 1981. Type (Bischler, 1979). Chile. O'Higgins: Rancagua, Bertero 409 (lectotype, W; isolectotype, PC). Plagiochasma elongatum Lindenb. & Gottsche, in Gottsche, Lindenb. & Nees, Syn. hepat. 519. 1846. Ruppinia elongata (Lindenb. & Gottsche) Trevis., Mem. Reale Ist. Lombardo Sci. Ser. 3, Cl. Sci. Mat. 4: 437. 1877. Aitonia elongata (Lindenb. & Gottsche) Kuntze, Revis. gen. pl. 1: 143. 1891. Type (Evans, 1915). Mexico.Veracruz: Comaltepec, Liebmann 290 (lectotype, YU; isolectotype, G). Plagiochasma mexicanum Lindenb. & Gottsche, in Gottsche, Lindenb. & Nees, Syn. hepat. 519. 1846. Ruppinia mexicana (Lindenb. & Gottsche) Trevis., Mem. Reale Ist. Lombardo Sci. Ser. 3, Cl. Sci. Mat. 4: 437. 1877. Aitonia mexicana (Lindenb. & Gottsche) Kuntze, Revis. gen. pl. 1: 143. 1891. Lectotype (Evans, 1915). Mexico. Puebla: San Lorenzo Tehuacan, Liebmann 71 (lectotype, YU; isolectotypes, BM, G, NY). Plagiochasma validum Bisch. var. ß minus Gottsche, Lindenb. & Nees, Syn. hepat. 520. 1846. Type. Chile. Valparaiso: Quillota, Bertero 1130, 1132, 1133 (syntypes, W). Aitonia lanigera Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 568. 1885. Plagiochasma lanigerum

Bischler et al. 159 (Spruce) Steph., Sp. hepat. 1: 85. 1898. Type. Ecuador. Tunguragua: Ambato; Los Baños, river Pastaza, Spruce s.n. (syntypes, MANCH). Aitonia subplana Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 567. 1885. Plagiochasma subplanum (Spruce) Steph., Sp. hepat. 1: 86.1898. Type. Ecuador. Tunguragua: Antombos, near river Pastaza, 1800 m, Spruce s.n. (holotype, MANCH). Plagiochasma brasiliense Steph., Sp. hepat. 1: 83. 1898. Type. Brazil. Paraná: Apiahy, Puiggari 851 (holotype, G; isotypes, FH, PC). Aytonia evansii Haynes, Bull. Torrey Bot. Club 34: 57. 1907. Plagiochasma evansii (Haynes) Steph., Sp. hepat. 6: 7. 1917. Type. Jamaica. Portland Gap, Blue Mountains, 21 Jul 1903, Evans 213 (holotype, YU; isotypes BM, FH, G, NY). Plagiochasma bornmülleri Steph., Sp. hepat. 6: 6. 1917. Type. Brazil. Rio Grande do Sul: Without locality, Bornmüller s.n. (holotype, FH; isotypes, FI, G, JE). Plagiochasma hosseusii Herzog, Feddes Repert. Spec. Nov. Regni Veg. 55: 4. 1952. Type. Argentina. Misiones: Dep. San Martín (Corpus), San Ignacio, Hosseus 581 (holotype, JE). Plagiochasma rupestre (J.R. et G.Forst.) Steph. fo. glaucum Schier, Nova Hedwigia 25: 561. 1974. Type. Peru. Lima: Rimac-Tal, May 1958, Volk 4384 (isotypes, PC, Hb. Volk). Thallus usually bluish green with purplish margins, 2.3-5 mm wide. Epidermal pores bounded by a ring of 4-5(-6) cells, often irregular in shape, radial walls usually not thickened. Appendages of scales usually purplish, acute apically and often hyaline, length : width ratio = 1.7-2.7 : 1, with row of 1-2(4) thin-walled, short cells apically. Gametophytic chromosome number n = 9 or 18. Distribution and ecology. The variety is widespread in warm regions of the world and has been recorded from Europe, the Mediterranean area and Macaronesia, SW Asia, India, tropical and S Africa, the Mascarenes, Australia, New Zealand, and Oceania. However, it has not been collected in eastern Asia and on the tropical Asiatic islands. Plagiochasma rupestre var. rupestre is distributed from 35° N to 40° S in the New World and is common in the southern part of the United States (Arizona, Colorado, Kansas, New Mexico, Oklahoma, Texas), in Mexico (Baja California, Chiapas (Düll, 1999), Chihuahua, D. F., Durango, Guerrero, Hidalgo, Jalisco, Mexico, Michoacan, Morelos, Nuevo León, Oaxaca, Puebla, Querétaro, San Luis Potosí, Sonora, Tlaxcala, Veracruz, Zacatecas) and in the subtropical and warm-temperate parts of southern South America: Chile (Atacama, Biobio, Los Lagos, O'Higgins, Santiago, Valparaíso), Juan Fernandez, Argentina (Buenos Aires, Catamarca, Córdoba, Jujuy, La Pampa (Hässel, 1963), Mendoza (Hässel, 1963), Misiones, Rio Negro (Hässel, 1963), Salta (Hässel, 1963), Tucuman), and Uruguay. Records from the inner Tropics are more scattered and are from Guatemala, Costa Rica, Panama, Jamaica, Haiti, Colombia (Cundinamarca,

Bischler et al. 160 Magdalena (Winkler, 1976), Santander, Tolima), Venezuela (Mérida), Ecuador (Azuay, Los Rios, Pastaza, Tunguragua) and the Galápagos Islands (Fernandina (Evans, 1902), Floreana, Isabela, Pinta), Peru (Amazonas, Ancash, Arequipa, Cajamarca, Cuzco, Junín, La Libertad, Lima, Piura), Brazil (Bahía (Yano, 1981), Espírito Santo (Schäfer-Verwimp, 1991), Minas Gerais, Paraná, Pernambuco (Yano & Andrade-Lima, 1987), Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo), and Bolivia (Cochabamba, Chuquisaca, La Paz, Potosí, Santa Cruz, Tarija). The variety prefers dry, warm areas and was collected from 150 m to 3800 m, on different soils, in rock crevices of various kinds of rocks (calcareous as well as acidic), and in stonework. Its habitats are either exposed or shaded by open vegetation. Selected neotropical specimens examined. MEXICO. BAJA CALIFORNIA: without locality, Brandegee 25 (NY, YU). CHIHUAHUA: 55 mi. N Chihuahua, km 87 on road to El Paso, Whittemore 2904 (MEXU). D. F.: Botanical Garden, 1979, Jovet-Ast s.n. (PC); Pedregal de San Angel, Delgadillo 1650 (MEXU); Desierto de los Leones, walls of El Convento, 1944, Ruiz Oronoz s.n. (MEXU). DURANGO: Tejamen, Palmer 473 (FH, NY, YU). GUERRERO: Taxco, 1947, Ruiz Oronoz s.n. (MEXU). HIDALGO: Hacienda Palmar near Pachuca, Rose et al. 8743 (NY, US, YU); San Miguel Regla, 1948, Ruiz Oronoz s.n. (MEXU). JALISCO: Guadalajara, barranco de Oblatos, Barnes & Land 122 (F, NY, YU). MEXICO: Sierra Nevada, W slopes of Popocatepetl, Hermann 20825 (PC); Barranca Malitenango Pócs 9551N (MEXU). MICHOACÁN: Morelia, Arsène 72 (F); SE Morelia, Rio Los Filhos, 2 km E of junction with Agua Sarca, Cerro Azul, Clausen s.n. (NY). MORELOS: S. Antonio Falls, Barnes & Land 480 (F, YU); Mun. Tepoztlan, Sierra Tepoztlan, 1955, Clausen s.n. (MEXU). NUEVO LEÓN: 5 mi. E Iturbide, Mc Gregor & Del Rosario 7 (NY); 19 mi. W Linares, Hwy 60, Mc Gregor & Del Rosario 230 (NY). OAXACA: San Felipe, E. & P. Hegewald 6471 (Hb. Hegewald). PUEBLA: Type of P. mexicanum; San Baltazar, Nicolas 2 (YU). QUERÉTARO: 1 km NW of Jurica, NW of Querétaro, Long & Delgadillo 29755 (E); above Santa Rosa Xajay, NE of San Juan del Rio, Long & Delgadillo 29757 (E). SAN LUÍS POTOSÍ: without locality, Parry & Plamer 1016 (NY). SONORA: Cerro Agua Caliente, Richards & Drouet 758 (BM, F). TLAXCALA: Near Amaxoc, Sharp 398 (US). VERACRUZ: Type of P. elongatum; Orizaba, Heller 1846 (W); Rio Jamapa between Huatusco and Coscomatepec, Long & Garcia 29674 (E). ZACATECAS: Concepción de Oro, Palmer 307 (F, NY, YU). GUATEMALA. Sacatepequez, Las Lajas, Standley 58128 (F, NY). COSTA RICA. Cartago, Taras, Melendez-Howell s.n. (PC). PANAMA. Chiriquí, Boquete, Svihla 40-447 (Hb. Fulford). JAMAICA. Type of Aytonia evansii; Blue Mts.., Content Gap, Evans 613 (NY, YU).

Bischler et al. 161 HAITI. Vicinity of Furcy, Leonard 4571 (YU). COLOMBIA. CUNDINAMARCA: Páramo de Guasca, Murillo & Fayad 157 (NY). SANTANDER: Vicinity of Vetas, Killip & Smith 17390 (JE). TOLIMA: without locality, Felix s.n. (E). VENEZUELA. MÉRIDA: Parque de los Escritores, Onraedt 78 V 5749 (JE). ECUADOR. AZUAY: Gualaquiza, Allioni 6601 (G). Loja: road Celica-Zapatillo, between Celica and Pozul, Anderson 971 (U). LOS RIOS: Guaranda-Bodegas, Rémy s.n. (PC). PASTAZA: Canelos, Spruce s.n. (BM, E, G, MANCH). TUNGURAGUA: Type of Aitonia lanigera; type of Aitonia subplana; Ambato, Spruce H 72 (MANCH). GALÁPAGOS: Floreana, Wittmer Farm, Gradstein H 134 (PC, U), Stewart 940 (YU); Isabela, Volcano Cerro Azúl, Gradstein H 444 (U); Pinta, Stewart 8676 (YU). PERU. AMAZONAS: Chachapoyas, Cauchille (Balsas-Leimebamba), P. & E. Hegewald 6759 (Hb. Hegewald); Chachapoyas, above Leimebamba, Frahm 2347 (U). ANCASH: Huaraz, Monterrey, P. & E. Hegewald 7471 (JE, Hb. Hegewald); Chavin Huantar, Menzel P-14 (U). AREQUIPA: Way to Puno, P. & E. Hegewald 5463 (PC). CAJAMARCA: Contumaza, Tambo de Lima (Contumaza-Cascas), P. & E. Hegewald 7376 (PC). CUZCO: Collpapampa, Bües 1523 (YU). JUNÍN: Yauli, Chacapalpa (La Croya-Janja), P. & E. Hegewald 5818 (PC). LA LIBERTAD: Otuzco, Casmiche, P. & E. Hegewald 5073 (PC). LIMA: Type of P. rupestre fo. glaucum; Huarochiri, Canchahuara, Cerrate & al. 4762 (US). PIURA: Huancabamba, W of Porculla-Pass, P. & E. Hegewald 7149 (PC). BRAZIL. MINAS GERAIS: Coxoeira, Claussen 94 (PC). PARANÁ: Type of Plagiochasma brasiliense. RIO DE JANEIRO: without locality, Glaziou 3402 (NY). RIO GRANDE DO SUL: Type of P. bornmülleri; Cruz Alta, Bornmüller 4994 (FH, FI, G, JE). STA CATARINA: Lauro Müller, Vital 2001 (PC). SÃO PAULO: Barra do Turvo, Vital 1855 (PC). BOLIVIA. COCHABAMBA: Loma San Pedro, Hermann 24598 (G, NY, US). CHUQUISACA: Yamparaez, 1 km S of Tarabuco, Lewis 83-1623 (F). LA PAZ: Type of P. peruvianum; Ingavi, Asplund 16 (JE); La Paz, Rusby 11 (NY); Sorata, 4030m, Rusby 3000 (NY). POTOSI: Chayanta, 1 km NE of Macha, Lewis 84-1650 (F). SANTA CRUZ: Florida, SE slopes of Cerro Porto Bermejo, Lewis 85-1428 (F). TARIJA: Aviles, 20 km NW of Padcaya, Lewis 84-2932 (F). CHILE. Type of Reboulia chlorocarpa. ATACAMA: without locality, Morong s.n. (NY, YU). BIOBIO: Concepción, Schwabe 50 (JE). LOS LAGOS: Valdivia, Hollermayer s.n. (JE). O'HIGGINS: Syntype of P. validum; Osorno, Pahyehue, Rio Cleauleufen, Sparre 2213 (G). SANTIAGO: Cuesta Barriga, Mahu 5025 (US). VALPARAÍSO: Syntype of P. validum and type of P. validum var. ß minus; La Leona, Bertero 408 (W). JUAN FERNANDEZ: Masafuera, Quebrada Casa, Hatcher & Engel 123 (US); Masatierra, Pangal, Skottsberg 1 (S, YU).

Bischler et al. 162 ARGENTINA. BUENOS AIRES: Sierra de la Ventana, Hanstal 61 (NY). CATAMARCA: Andalgala, Sleumer s.n. (S). CÓRDOBA: Los Gigantes, Capurro 9990 (PC, S). JUJUY: Sta Catalina, La Puna, Claren 11379 (S). MISIONES: Type of P. hosseusii; S. Martin (Corpus), San Igancio, Hosseus 576 (JE). TUCUMÁN: Siambon, Lorentz s.n. (BM, EGR, FH, G). URUGUAY. Cerro Montevideo, Herter 82030 (PC). Plagiochasma rupestre var. rupestre comprises haploid and polyploid, morphologically indistinct populations. In Peru (Lima) and Argentina (Córdoba), the two cytotypes coexist. The haploid cytotype was additionally recorded from Mexico (San Luis Potosí), Brazil (São Paulo), Guatemala, and Argentina (Buenos Aires, Catamarca).

9b. Plagiochasma rupestre var. volkii Bischl., Rev. Bryol. Lichénol. 44: 289. 1978. Type. Africa. Namibia. Nevdamm near Windhoek, Volk 948 (holotype, JE).

Figs. 133: F-L, 134

Thallus usually bright green with narrow purplish margins, 3-3.9 mm wide. Epidermal pores bounded by a ring of (4-)5-6 cells, usually regular in shape, with thickened radial walls. Appendages of scales usually hyaline, length : width ratio = 4.5-5 : 1, acuminate and with row of 3-5 elongate cells with strongly thickened walls apically. Gametophytic chromosome number n = 9. Distribution and ecology. The variety, much less common than var. rupestre, is known from Zimbabwe, Lesotho, Namibia, South Africa, and the Neotropics. In the New World, collections are from Peru (Lima), Bolivia (Tarija), and Argentina (Catamarca, Tucumán), south of the equator (1229° S), from dry, warm areas, between 1200 and 2600 m, in rock crevices shaded by shrubs. Neotropical specimens examined. PERU. LIMA: Huarochiri, Sta Eulalia, Goodspeed 33011 (US). BOLIVIA. TARIJA: Prov. Cercado, road Tarija-Entre Rios km 25, Cerro Mesón, Gradstein 7650 (U). ARGENTINA. CATAMARCA: Ciudad Belén, Sleumer 2145 (S). TUCUMÁN: Tafi, La Cienaga, Sleumer s.n. (S). The neotropical collections are sterile and their identification remains tentative. Sexual structures are needed to confirm their identity.

15. REBOULIA Raddi, Opusc. Sci. 2: 357. 1818, nom. et orth. cons. Type. Reboulia hemisphaerica

Bischler et al. 163 (L.) Raddi (Marchantia hemisphaerica L.). Genus dedicated to Eugen de Reboul, citizen of Florence in the early 19th century. Thallus 3-9 mm wide, dull green, often tinged with purple. Epidermal cells with conspicuous trigones. Epidermal pores with 3-6 rings of 6-9 cells, radial walls thickened. Air chambers in several layers. Basal tissue without mucilage cavities. Ventral scales with oil-cells and marginal papillae, and 2-4 filiform appendages, not constricted basally. Monoecious. Antheridia and archegonia as in the family, archegonia 1-2 per archegonial cavity. Archegoniophore stalk without assimilatory strip, with 1 rhizoid furrow, with scales at top and bottom; receptacle 4-6-lobed. Calyptrae 3-4-layered after fertilization. Involucres bilabiate, Involucres opening obliquely downwards. Sporophyte one per involucre. Capsules opening to 1/3 of capsule length by a fragmenting lid. Spores 2500-3000 per capsule, 60-90 µm diameter, proximal and distal faces with tuberculate areoles and tuberculate ridges, trilete scar distinct. Elaters with 2-3 helical bands. Gametophytic chromosome number n = 9 or 18. A genus comprising 1-3 drought-tolerant species inhabiting rock crevices and rock outcrops, recorded from relatively warm areas in both hemispheres, 62°N to 50°S. It is frequent in the Mediterranean area, but is absent from areas with long, cold winters and from tropical, humid zones.

1. Reboulia hemisphaerica (L.) Raddi, Opusc. Sci. 2: 357. 1818. Marchantia hemisphaerica L., Sp. pl. ed. 1, 1138. 1753. Grimaldia hemisphaerica (L.) Lindenb., Nova Acta Acad. Caes. Leop.Carol. Nat. Cur. 14, Suppl. 106. 1829. Type. Europa (syntype, OXF, n.v.). Figs. 1: L, 3: K, 137, 138, 160: B Thallus 3-8 mm wide, light green with narrow, purplish borders which are often lobulate-sinuose and ascending; branching dichotomous, ventral and by apical adventitious branches. Epidermal cells with thin to slightly thickened walls and conspicuous trigones. Epidermal pores 190-250 µm diameter, bounded by 4-6 concentric rings of 5-9 cells, radial walls usually strongly thickened. Air chambers in 5-6 layers. Scales purplish, imbricate, with oil cells and marginal papillae, with 2-3 filiform appendages, 2-3 cells wide basally, with row of 2-4 cells apically. Monoecious. Antheridia in irregular groups or in cushions bounded by more or less numerous, purplish scales, in neotropical specimens usually behind archegoniophore. Archegoniophore stalk 6-25 mm long; receptacle 4-7lobed, hemispherical, with compound epidermal pores on dorsal side. Spores yellowish-brown, 60-90 µm diameter, areolate on both faces, with 4-5 large areoles across diameter and broad wing, ridges of areoles and wing tuberculate. Elaters with 2-3 helical bands. Gametophytic chromosome number n = 9 or 18. Distribution and ecology. Reboulia hemisphaerica is widespread in warm-temperate areas all over the world. It is recorded from Europe up to 61° N, the Mediterranean area, Macaronesia, tropical eastern and South Africa, and SW Asia. The eastern Asiatic and Australasiatic records (eastern Asia, India, Sri Lanka, Japan, Taiwan, tropical Asiatic Islands, New Caledonia, New Zealand, Australia,

Bischler et al. 164 Tasmania), are genetically distinct and might belong to other species (Boisselier-Dubayle et al., 1998). In the U. S. A. the species has been recorded from 42 (out of 49) states. In the Neotropics, the species is quite common in south-eastern Brazil, but rare elsewhere. It is known from Mexico (Chihuahua (Whittemore, pers. comm.), Chiapas, Colima, D. F., Durango, Hidalgo, Mexico, Morelos, Nuevo León, Oaxaca, Puebla, Querétaro, Socorro Is., Tlaxcala, Veracruz), Bermuda, Cuba, Haiti, Jamaica, Puerto Rico, Brazil (Rio Grande do Sul), Bolivia (Rusby, 1895), Chile (Los Lagos (Reimers 1926), Magellanes, Santiago, Juan Fernandez), Argentina (Buenos Aires (Hässel, 1963), Chubut (Hässel, 1963), Misiones, Neuquén, Rio Negro (Hässel, 1963), Tierra del Fuego (Engel 1976)), and Uruguay. The species grows usually in dense colonies on humid or wet clay, sand, or sandy clay, overlying limestone, dolomite or other calcareous rocks, in rock crevices or stone walls, on exposed river banks, in grassland, on roadsides, under scattered shrubs, or in open forests, from sea level to 4000 m. Selected specimens examined. MEXICO. CHIAPAS: Palenque ruins, Maslin L-41415 (MEXU). COLIMA: Nevado de Colima, S of Ciudad Guzman, Eiten 1137 (NY). D. F.: Botanical Garden, 1979, Jovet-Ast s.n. (PC); La Cima, Barnes & Land 408 (NY), Whittemore & Leidig 1850 (MEXU); Salazar, Stahl s.n. (G); Cañada, Pringle 15348 (FH, MEXU), Barnes & Land 424 (NY); Parque Ejidal San Nicolas, Totolapan, Long 29560 (E). DURANGO: Tejamen, Palmer 472 (NY). HIDALGO: Camino de Pachuca to Real del Monte, 1948, Ruiz Oronoz & Herrera s.n. (MEXU); above Real del Monte, Sharp 507 (TENN). MEXICO: Popocatepetl, Sierra Nevada, Hermann 20825 (PC, US); Road to Popocatepetl, 15 km from Amecameca, Düll 2/388 (PC); NW portion of Popocatepetl, Delgadillo 2195 (MEXU); La Jagua de Alicalican, S slope of Ixtaccihuatl, Whittemore 4107 (MEXU); La Marquesa-Tenango, 1975, Perez de Lamothe s.n. (PC); Volcán Ixtaccihuatl, Purpus 550 (PC), Eggers & Frahm 792559 (PC), Delgadillo 1726 (MEXU); Rio Frio, Sharp 312 (MEXU); MORELOS: near Cuernavaca, along Tapeite River, W. A. & E. L. Merrill 13 (NY). NUEVO LEÓN: Cerro Potosi, McGregor & Del Rosario 384 (NY). OAXACA: Laguna Santiago Tilapa, Rzedowski 19421 (TENN). PUEBLA: Near Huejotzingo, Sharp 1199 b (MEXU); Tezuitlan, Sharp 9804 (MEXU, TENN). QUERÉTARO: Sierra Gorda, Chuveje Cascades, W of Jalpan, Long & Delgadillo 29722 (E); below La Florida, W of Conca, Long & Delgadillo 29734 (E). SOCORRO IS.: 1932, Howell 176 (NY). TLAXCALA: Amaxoc, Sharp 382 (TENN). VERACRUZ: Córdoba, 2 km W on road to Orizaba, Frye 2240 (NY, TENN); Orizaba, Barnes & Land 677(NY); La Pitahaya near Xalapa, Long 29686 (E). BERMUDA. S. loc., Moseley s.n. (BM), Dodge s.n. (NY); Paget Parish, 1900, Howe s.n. (NY); North Side, Britton 380 (NY); near Harrington House, Brown 442 (NY); Painters Vale, Britton 1863 (NY), Marble 807 (NY); Hamilton, 1900, Howe s.n. (NY); Tucker's Farm, Brown & Britton 819 (NY).

Bischler et al. 165 CUBA. Without locality, Wright s.n. (NY). JAMAICA. S. loc., Wilson 595 (NY), Palisot de Beauois s.n. (G), Wilds s.n. (NY); Chestervale, Underwood 1173 (FH); Cinchona, Cummings 30 (NY); trail from Morce's Gap to Vinegar Hill, Maxon & Killip 686 (NY). HAITI. Curcy, 1980, Tixier s.n. (PC). PUERTO RICO. Mt. Britton peak, Sierra, Steere 7083 (U). BRAZIL. RIO GRANDE DO SUL: S. loc., Rick 169 (G), Lindman 89, 102 (G), Gaudichaud 90 (PC); São Leopoldo, Rick s.n. (PC). CHILE. MAGELLANES: Punta Arenas, 1906, Thaxter s.n. (NY). SANTIAGO: near the town, Costes 93 (PC), s.n. (NY). JUAN FERNANDEZ: Masafuera, Skottsberg 159 (G); Masatierra, Skottsberg 124 (G); Masatierra, near base of Piramide, Hatcher & Engel 509 (NY). ARGENTINA. CHUBUT: Lago Futalaufquen, Hässel & Donterberg 1710 (BA). MISIONES: Loreto, Perez Moreau 8381 (NY). NEUQUÉN: Nahuel Huapi, Pto Lavallol, Isla Victoria, Hässel 9638 (PC). URUGUAY. Tacuarembo, Herter 1243 (G, NY, UC, U); Lavalleja, Penitente, Herter 2147 (G). Reboulia hemisphaerica is easily distinguished by its ventral scales having several filiform appendages. The species comprises at least three genetically distinct but morphologically hardly differentiated, allopatric components that may deserve specific status: two haploids, one from Europe and Africa, found also in southern U. S. A., a second from Japan, and a polyploid from Australasia (BoisselierDubayle et al., 1998). The neotropical specimens have not been examined genetically. Whether they belong to one of these three components or to additional ones, remains to be investigated. None of the seven subspecies described by Schuster (1992b) has been recorded or seen from the Neotropics.

9. LUNULARIACEAE H.Klinggr., Die höheren Crypt. Preuss. 9. 1858. Type. Lunularia Adans. A monogeneric family.

16. LUNULARIA Adans., Fam. 2: 15. 1763. Type. Lunularia cruciata (L.) Dumort. (Marchantia cruciata L.). Named for the lunate gemma cups. Lunularia is monospecific.

Bischler et al. 166 1. Lunularia cruciata (L.) Dumort. ex Lindb., Notiser Sällsk. Fauna Fl. Fenn. Förh. 9: 298. 1868. Marchantia cruciata L., Sp. pl. ed. 1, 1137. 1753. Type. Europe (syntype OXF, n.v.). Figs. 1: F, K, M, 2 :G, M, 3 :C, 139, 140, 160: C Preissia cucullata Nees & Mont., in Montagne, Ann. Sci. Nat., Bot. ser. 2, 9: 44. 1838. Type. Chile. Bertero s.n. (isotypes, PC, STR, W). Lunularia thaxteri A.Evans & Herzog, in Herzog, Arch. Esc. Farm. Fac. Ci. Med. Cordoba, Sec. Ci. 7: 5. 1938. L. cruciata fo. thaxteri (A.Evans & Herzog) Hässel, Opera Lilloana 7: 129. 1963. L. cruciata subsp. thaxteri (A.Evans & Herzog) R.M.Schust., Hep. N. America 6: 91. 1992. Type. Chile. Biobio: Dep. Coronel, Concepción, 1905/1906, Thaxter s.n. (JE). Thallus 4-13 mm wide, light green, sometimes tinged with purple. Branching dichotomous and apical. Epidermal cells thin- or thick-walled. Epidermal pores simple, with 3-5 concentric rings of 317 cells, with hyaline inner ring, radial walls thin or thickened. Air chambers in one layer, with chlorophyllose filaments. Basal tissue without mucilage cavities. Ventral scales hyaline or purplish, in two rows, with oil-cells and marginal papillae and one large, basally constricted reniform appendage. Asexual reproduction by discoid gemmae in crescent-shaped gemma cups with entire margins. Dioecious. Antheridia in terminal, elliptical receptacles, bounded by a membrane, without scales. Archegonia 2-8 per archegonial cavity, in terminal, 4-lobed cushions bounded by white, involute scales. Archegoniophore stalk elongating at time of spore maturity, without assimilatory strip and without rhizoid furrow, with scales; receptacle 4-lobed, very reduced, without epidermal pores and assimilatory layer. Calyptrae 2-4-layered after fertilization. Involucres 4, tubular. Pseudoperianth absent. Sporophytes 1-3 per involucre, with massive, shortly elongating seta (2-3 times capsule length). Capsules opening by a lid and 4 regular valves to capsule bottom, walls without annular thickenings. Spore : elater ratio more than 4 : 1. Spores 5000-6000 per capsule, 14-22 µm diameter, proximal and distal faces faintly tuberculate, trilete scar indistinct. Elaters with 2-3 helical bands. Gametophytic chromosome number n = 9. Distribution and ecology. Lunularia cruciata is nearly cosmopolitan, distributed worldwide from 62°N to 45°S, very common in rather warm areas but rare in the tropical belt and in zones with long, cold winters where it was most likely introduced recently. It has been recorded from Europe north to southern Scandinavia, the Mediterranean area, Macaronesia in eastern and South Africa, the Mascarenes, in Asia from SW Asia through Afghanistan to India, Sikkim, Sri Lanka, and Japan, from Australia, and New Zealand. In North America the species has been recorded from Canada and the U. S. A (at least from 24 states), but over most of the area, only from greenhouses. In the Neotropics, it is known from Mexico (D. F.), Bermuda (Evans, 1906), Costa Rica (Morales, 1991), Jamaica, Colombia (Cundinamarca), Bolivia (Chuquisaca, Cochabamba (Stephani, 1916), La Paz, Santa Cruz, Tarija), Peru (Cuzco (Hässel, 1963), Huanuco (Bryan, 1929), La Libertad (Hegewald & Hegewald, 1977)), Brazil (Minas Gerais (Gradstein & Pinheiro da Costa, 2003), Rio Grande do Sul, Rio de Janeiro

Bischler et al. 167 (Yano, 1995), São Paulo (Yano, 1995)), Chile (Biobio, Los Lagos (Herzog, 1938), O'Higgins (Montagne, 1850), Santiago, Valparaíso, Juan Fernandez), Argentina (Buenos Aires, Catamarca, Chubut , Entre Ríos, Mendoza (Hässel, 1963)), Misiones, Neuquén, Rio Negro, Salta (Hässel, 1963), Tucumán), and Uruguay. The species grows in dense colonies or scattered among other thallose liverworts, on humid or wet sand, sandy clay, bare or rocky soil, overlying limestone, dolomite, tuff or other calcareous rocks, in rock crevices, at base of cliffs, in stone walls, on river banks, in caves, in grassland, exposed or shaded by open vegetation, from 1000 to 4200 m in the tropical zone but more common below 2000 m, reaching sea level in the temperate zones. It is a colonist of man-made habitats (greenhouse weed), and might have been introduced in many areas with garden soil and plants, but is also known from natural habitats. The species reproduces sexually very rarely and only in warmer areas. Selected specimens examined. MEXICO. D. F.: Desierto de los Leones, Delgadillo 1345 (MEXU); Parque Ejidal San Nicolas, Totolapan, Long 29553 (E); Parque Nacional Desierto de los Leones, near Exconventa, Long 29594 (E). JAMAICA. Morce's Gap, Underwood 500 (Hb. Fulford). COLOMBIA. CUNDINAMARCA: Bogotá, Schultes 11323 (U). BRAZIL. RIO GRANDE DO SUL: Dos Irmãos, Vianna 5693 (U); Bagé, Bueno 39804 (U). BOLIVIA. CHUQUISACA: Hernando Siles, 8 km W Monteagudo, Lewis 84-912 (F). LA PAZ: Inquisivi, Río Chichipata, Lewis 86-2146 (F); Larecaja, 4 km NE Soratá, Lewis 82-224A (F). SANTA CRUZ: Vallegrande, 12 km SW Vallegrande, Lewis 85-752 (F). TARIJA: Aviles, 20 km NW of Padcaya, Lewis 84-2932 (F); O'Connor, 1 km E Canaletas, Lewis 84-2565 (F); O'Connor, 5 km NW Narvaez, Lewis 84-2649 (F); O'Connor, Abra Castellón, 7 km NW of Entre Ríos, Gradstein 5677 (U); O'Connor, Narvaez-San Josecito, Gradstein 7662 (U); O'Connor, road Tarija-Entre Ríos, Gradstein 7620 (U). CHILE. Central Provinces, Gay s.n. (PC). BIOBIO: Type of L. thaxteri ; Concepción, Lota, Parque Isidora Cousiño, Mahu 20198 (PC). SANTIAGO. Baños de Apoquindo, 1905, Thaxter s.n. (JE). VALPARAÍSO: Corral, s.col., s.n. (JE). JUAN FERNANDEZ: Masatierra, s.col. 3 (G); Masatierra, Cerros de Bahia Cumberland, Pisano & Montaldo 1424 (UC). ARGENTINA. BUENOS AIRES: Capital, 1981, Hässel s.n. (PC); Punta Lara, Hässel 8341 (PC), Castellanos 729 (BA); Sierra de la Ventana, Kühnemann 4147 (BA); near the Capital, Villa Devoto, Sanchez 200 (BA). CATAMARCA: La Cuesta del Singuil, km 1421, Vianna 14265 (BA); Las Chacritas, Vianna 13974 (BA). CHUBUT: Lago Futalaufquén, Hässel 22947 (BA); Esquel, Kühnemann 5015 (BA). ENTRE RÍOS: Gualeguay, Il Distrito, Estancia El Retiro, Wright 9264 (BA). MISIONES: Yabebiri, Perez Moreau 8279 (BA). NEUQUÉN: Lago Nahuel Huapi, Perez Moreau 9265 (BA, PC); Nahuel Huapi-cena, Pen. Quebrihue, 1991, Hässel s.n. (PC); Lago Traful, Kühnemann

Bischler et al. 168 2363 (BA). RIO NEGRO: Lago Guillermo, Kühnemann 2272 b (BA); Huapi, C° Lopez, Huapi, Perez Moreau 9267 (BA). TUCUMÁN: way to Salta, Castellanos 7341 (BA); Sierra de Medina, Vianna 13945 (BA). URUGUAY. Tacuarembo, Herter 1243 (G, UC, U); Atahualpa, Herter & Strahl 1472 (G, UC, U); Florida, La Palma, Herter 1243a (G, NY); Lavalleja, Aigua, Herter 94518 (G). Lunularia cruciata is easily distinguished from all other Marchantiales by its crescent-shaped gemma cups which are almost always present. The type of L. thaxteri from Chile has strongly thickened epidermal cells, purplish ventral side of thallus and purplish ventral scales, and epidermal pores bounded by fewer rings but with more cells in each ring. A plant with these characters from Madeira did not show any genetic difference with Lunularia cruciata (Boisselier-Dubayle et al., 1995). Lunularia thaxteri is therefore treated here as a synonym. However, whether the neotropical specimens are genetically identical with the European and African representatives needs to be investigated.

10. MARCHANTIACEAE (Bisch.) Lindl., Nat. Syst. Bot., ed. 2, 412. 1836. Type. Marchantia L. Thallus branching dichotomous and apical. Epidermal pores compound with 4-7 rings of cells, with hyaline inner ring, or vestigial, bounded by a single ring of 4 cells, or absent. Air chambers in one layer, with chlorophyllose filaments, or with papillae, or empty, or vestigial, with chlorophyllose filaments only, not roofed by epidermis, or absent. Ventral scales in 2-10 rows. Asexual reproduction by discoid gemmae in cup-shaped gemma-cups, or absent. Monoecious or dioecious. Antheridia and archegonia developing on stalked, terminal gametangiophores. Archegoniophore receptacle with compound or vestigial epidermal pores on dorsal side. Calyptrae 2-4-layered after fertilization. Involucres tubular, cup-shaped or bivalved. Pseudoperianths campanulate, or absent. Sporophytes with hardly or slightly elongating seta. Capsules splitting into irregular valves, wall with annular thickenings. Spore : elater ratio more than 4 : 1. Elaters with helical bands. Gametophytic chromosome number x = 9. A family comprising five genera; two, Marchantia and Dumortiera, are represented in the Neotropics. A third genus, Preissia, has been recorded from Mexico and Bolivia. As shown by Grolle (1980) and Bischler (1984), the neotropical specimens of Preissia were misidentified and belong to Marchantia chenopoda L. Three subfamilies have been described, two of which are represented in the Neotropics: Marchantioideae (genus Marchantia) and Dumortieroideae (R.M.Schust., Phytologia 56: 71. 1984) (genus Dumortiera).

Bischler et al. 169 KEY TO THE NEOTROPICAL GENERA OF THE MARCHANTIACEAE 1. Thallus without purplish pigmentation. Epidermis, pores and air chambers absent or vestigial. Ventral scales in 2 rows, small, without marginal papillae. Bristles on ventral side of thallus and gametangiophores. Gemma-cups never produced. Archegoniophore stalk without assimilatory strips. Involucres tubular. Pseudoperianth lacking. Distal face of spores tuberculate............ 17. Dumortiera 1. Thallus often with purplish pigmentation. Epidermis, pores and air chambers well developed. Ventral scales large, in 4 -10 rows, each pair of rows of different shape, the median ones with a large appendage, with marginal papillae and oil-cells. Bristles lacking. Gemma-cups (cup-shaped) usually present. Archegoniophore stalk with assimilatory strip(s). Involucres bilabiate. Pseudoperianth present, campanulate. Distal face of spores not tuberculate ........................................ 18. Marchantia

17. DUMORTIERA Nees, in Reinwardt, Blume & Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.Carol. Nat. Cur. 12: 410. 1824. Type. Dumortiera hirsuta (Sw.) Nees (Marchantia hirsuta Sw.). Genus dedicated to Count B. C. J. Dumortier (1797-1878), Belgian politician and botanist. Hygrophila Taylor, in J.Mackay, Fl. Hibern. 2: 53. 1836, nom. illeg. Type. H. irrigua (Wilson) Taylor (Marchantia irrigua Wilson). Dumortiera contains several cytotypes, which are often considered as separate species or subspecies. Since morphological differences among the cytotypes are few and rather weak, the genus is treated here as monospecific. It is a perennial stayer of forest floors, able to survive under weak light intensities, and has a worldwide, oceanic distribution between 40°N and 40°S, but reaching up to 50°N in atlantic Europe.

1. Dumortiera hirsuta (Sw.) Nees, in Reinwardt, Blume & Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12: 410. 1824. Marchantia hirsuta Sw., Prodr. 145. 1788. Type (Grolle, 1976). Jamaica. Swartz s.n. (holotype, S; isotype MW, UPS) (n.v.). Figs. 1: M, 141, 142, 160: D Marchantia irrigua Wilson, in Hooker, Brit. Fl. 2: 106. 1833. Hygrophila irrigua (Wilson) Taylor, in J. Mackay, Fl. Hibern. 2: 54, 1836. Dumortiera irrigua (Wilson) Nees, Naturgesch. eur. Leberm. 4: 159. 1838. Dumortiera hirsuta var. irrigua (Wilson) Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 566. 1885. Type. Ireland (n.v.). Hygrophila nepalensis Taylor, Trans. Linn. Soc. 17: 392. 1835. Dumortiera nepalensis (Taylor) Nees, Naturgesch. Eur. Leberm. 4: 169. 1838. Type. Nepal, Wallich s.n. (n.v.). Dumortiera hirsuta var. angustior Gottsche, and fo. depauperata Gottsche, Lindenb. & Nees, Syn.

Bischler et al. 170 hepat. 544. 1846. Type. Mexico. Veracruz: Mirador, Liebmann s.n.; Peru. Poeppig s.n. (n.v.). Dumortiera hirsuta var. intermedia Gottsche, Lindenb. & Nees, Syn. hepat. 544. 1846. Syntypes. South Africa, Brazil, Beyrich s.n. (n.v.). Dumortiera hirsuta var. latior Gottsche, Lindenb. & Nees, Syn. hepat. 544. 1846. Neotropical syntypes. Venezuela. Caracas, Moritz 46b; Brazil. Minas Gerais: Martius s.n.; Mexico. Puebla: Chinantla, Liebmann s.n. (n.v.). Dumortiera hirsuta var. trichopus Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 567. 1885. Type. Ecuador. Chimborazo: Spruce s.n. (n.v.). Dumortiera hirsuta var. brasiliensis Schiffn., in Schiffner & Arnell, Oesterr. Akad. Wiss., MathNaturwiss. Kl. Denkschr. 111: 8. 1964. Type. São Paulo. Near Lapa, Schiffner 2244 (n.v.). Thallus 8-30 mm wide, flat, deep green, not tinged with purple. Branching dichotomous and apical. Epidermal pores absent, or at thallus apex only, bounded by a single ring of 4 cells, radial walls thin. Air chambers absent, or in one layer and vestigial, with chlorophyllose filaments not roofed by epidermis. The floor of the air chambers becomes the expose surface of the mature thallus. Basal tissue with few oil-cells, without mucilage cavities. Bristles present on ventral thallus side and on gametangiophores. Ventral scales in two rows, small, hyaline, without oil-cells, papillae or appendage. Asexual reproduction by specialized propagules lacking. Monoecious or dioecious. Antheridiophores with disc-shaped receptacle, with small ventral scales underneath and bristles on margins and dorsal side, stalk short, without assimilatory strip, with 2 rhizoid furrows; rarely, antheridial branches develop in the female receptacles. Archegonia in stalked, terminal archegoniophores, up to 7 per archegonial cavity. Archegoniophore stalk long, without assimilatory strip, with 2 rhizoid furrows and small scales at top; receptacle shallowly 6-16-lobed, without or with few epidermal pores, sometimes with open cavities bounded by a ring of 8-13 cells, with bristles on margins and dorsal side. Calyptrae 2-3-layered after fertilization. Involucres tubular, opening at top by a ventral-terminal slit, with bristles. Pseudoperianths lacking. Sporophytes 1-2 or more per involucre with slightly elongating seta. Capsules splitting into 4-8 irregular valves to 3/4 of capsule length. Spore : elater ratio more than 4 : 1. Spores to 20 000 per capsule, 20-35 µm diameter, distal and proximal faces tuberculate, trilete scar distinct. Elaters with 2-4 helical bands. Gametophytic chromosome number n = 9, 18, or 27. Distribution and ecology. Dumortiera hirsuta has been recorded from tropical and warmtemperate areas : from western Europe up to Great Britain and Ireland, the Mediterranean area, Macaronesia, Africa (Ethiopia, tropical Africa, South Africa), the Mascarenes, Asia (from India and Sri Lanka, to eastern Asia, Japan and Taiwan), the tropical Asiatic islands, Oceania, and New Zealand. The species is known from 19 states in the U. S. A. and is common in the Neotropics. Its range extends from Mexico (Chiapas, D. F. (Svilha, 1941), Hidalgo, Jalisco, Mexico (Reed & Robinson, 1967), Michoacán (Svilha, 1942), Morelos, Oaxaca, Puebla, Querétaro, San Luis Potosí,

Bischler et al. 171 Tamaulipas, Veracruz, Guadalupe Is. (Frye & Clark, 1937)), Guatemala (Evans, 1919a), Honduras (Evans, 1919a), Nicaragua, El Salvador (Standley & Calderon, 1925), Costa Rica, Panama (Stotler et al., 1998), Bermuda (Evans, 1906), Cuba (Arnell, 1956), Jamaica, Haiti, Dominican Republic, Puerto Rico, Leeward Islands (Monserrat (Evans, 1919a)), Windward Islands (Barbados, Dominica, Guadeloupe, Martinique, St. Vincent (Fulford, 1984), Grenada (Evans, 1919a)), Colombia (Caldas, Chocó, Cundinamarca, Magdalena, Meta, Riseralda (Gradstein, 1995), Santander (Herzog, 1934), Tolima (Robinson, 1967), Valle (Gradstein & Hekking, 1979)), Venezuela (Aragua (Evans, 1919a), Caracas, Zulia (Griffin, 1977)), Trinidad (Underwood, 1895), Ecuador (Chimborazo (Spruce, 1885), Cotopaxi, Los Ríos, Pichincha, Tunguragua, Galápagos (Isabela)), Peru (Amazonas, Cajamarca, Cuzco (Hässel, 1963), Huanuco, Loreto (Spruce, 1885), Puno (Soukoup, 1955), San Martín), Brazil (Acre (Yano & Costa, 1992), Amazonas (Spruce, 1885), Espírito Santo (Schäfer-Verwimp, 1991), Mato Grosso (Stephani, 1897), Mato Grosso do Sul (Hell, 1969), Minas Gerais (Nees, 1833), Paraná, Rio de Janeiro, Rio Grande do Sul (Vianna, 1970), Santa Catarina, São Paulo), Bolivia (Beni, Cochabamba (Stephani, 1916), La Paz, Santa Cruz, Tarija), Paraguay, Argentina (Buenos Aires (Hässel, 1963), Córdoba (Hässel, 1963), Jujuy (Hässel, 1963), Misiones (Hässel, 1963), Salta (Hässel, 1963), Tucumán), to Uruguay. The species is very common and has been collected in the Neotropics on moist soil, dripping or mossy rocks, usually in shaded places, on streamsides, near waterfalls, in caves, in disturbed and undisturbed montane rain forests, from 150 to 3500 m, more commonly below 2000 m. Selected specimens examined. MEXICO. CHIAPAS: Mun. Union Juarez, volcán Tacana, way between Talquian and Chiquihuite, Martinez & Reyes 20442 (MEXU); Mun. Barrizabal, above Finca El Suspiro, s.col., s.n. (MEXU); Mun. Ocosingo, near Lacantus river, Gradstein & Equihua 8170 (MEXU); trail to village of San Manuel, S of Palenque, Mattews 1234 B (U). HIDALGO: Mun. Mineral del Chico, Lot & Novelo 1066 (MEXU). JALISCO: Mun. Tecolotlan, 10 km S of Quila to Tecolotlan, Novelo & Philbrick 1144 (MEXU). MORELOS: near Cuernavaca, Pringle 16671 (MEXU). OAXACA: 48 km NE Llano de las Flores, towards Valle Nacional, Delgadillo 848 (MEXU); road from Oaxaca to Tuxtepec, Sharp 9845 (MEXU); Sierra Juarez, 1962, Robinson s.n. (MEXU); Mun. Juquila, near Santa Rosa, Rzedowski 19585 (MEXU). PUEBLA: Mun. Zaragoza, Cerro Totoltepec, 10 km N of Zaragoza, on road to Zacapoaxtla, Grimes 2746 (MEXU); Mun. Xochitlan, Arroyo Atena, 15 km SE Zapotitlan, Grimes 2762 (MEXU). QUERÉTARO: Sierra Gorda, Rio Escañela, W of Jalpan, Long & Delgadillo 29717 (E). SAN LUIS POTOSÍ: near Zacatipa, Sharp 9918 a (MEXU). TAMAULIPAS: Rancho del Cielo above Gomez Ferias, Delgadillo 2702 (MEXU). VERACRUZ: Mirador, Liebmann 17 (PC); Córdoba, Motzorongo, Maury 5508, 5509 (PC); valley of Córdoba, E. B. 2143 (PC); Córdoba, Bourgeau 2143 (PC); Ojo de Agua, 1949, Ruiz Oronoz & Herrera s.n. (MEXU); Hacienda Tenajapa, 1948, Ruiz Oronoz s.n. (MEXU); way to Rio Sardo, Tlapacoyan, Ruiz Oronoz & Herrera s.n (MEXU); Xalapa, natural forest beside Jardin Botanico,

Bischler et al. 172 Long 29663 (E); Rio Jamapa between Huatusco and Coscomatepec, Long & Garcia 29672 (E); Piedra de Agua, between Xalapa and Naolinca, Long et al. 29627 (E). NICARAGUA. Chinandega, Volcán El Viejo, Baker 2179 (W); Carillo, Pittier 6016 (G). COSTA RICA. S. loc., Endres 65 (W); Carillo, Pittier 6016 (PC); Prov. Heredia, Braulio Carrillo, area Zursui, Gomez 20105 (U); Prov. Puntarenas, San José, Cerro Echandi, Gomez et al. 21847 (U). JAMAICA. Trail from Morce’s Gap to Vinegar Hill, Maxon & Killip 1314 (PC). HAITI. Massif Hotte, Dept. Grande Anse, 41 km S Roseaux, Zanoni et al. 24158 K (U). DOMINICAN REPUBLIC. S. loc., Balbis s.n. (PC); Prov. Puerto Plata-Santiago, Loma Diego O'Campo, 15 km NE Santiago, Zanoni et al. 24158K (U). PUERTO RICO. B. Cialitos, Jayuya, Pagán 333 (MEXU); Upper Rio de la Mina, Luquillo Mts., Pagán 4094 (U). WINDWARD ISLANDS. Barbados: mis. Hooker (W). DOMINICA: S. loc., Bory s.n. (PC). GUADELOUPE: S. loc., L'Herminier s.n. (PC), Richard s.n. (PC), Le Normand s.n. (PC). MARTINIQUE: S. loc., 1824, Duperrey s.n. (PC). COLOMBIA. CALDAS: Mun. Cinchina, opposite Cenicafé, Van Zanten & Gradstein 581 (U). CHOCÓ: Cerro del Torrà, NE slope, Silverstone-Sopkin 1367 (U). CUNDINAMARCA: Bogotá, Cipacón, 1861, Lindig s.n. (PC); La Vega, 1906, Apollinaire s.n. (PC). MAGDALENA: Sierra Nevada, Los Arroyitos, Kirkebride 2378 (U). META: Cerro Nevado de Sumapaz, Cleef 7816 (U); Villavicencio, N of town near Rio Guaitiquia, Van Zanten & Gradstein 699 (U). VENEZUELA. CARACAS: Galipan, Linden 338 (G, PC); Galipan, Funck & Schlim 338 (PC). ECUADOR. COTOPAXI: around Pilalo, Holm-Nielsen & Jeppesen 1182 (U); Lacatunga, Tenafuerte, Rio Lilao, km 52-53, Dodson & Gentry 12820 (U). LOS RIOS: Rio Palenque forest res., Gradstein et al. 6944 (U). PICHINCHA: 15 km E of Santo Domingo de los Colorados, Arts 18/056 (MEXU); road from Lloa (10 km W of Quito) to Rio Cristal, Gradstein et al. 6780, 6799 (U); road from Quito to Santo Domingo, W slope of Cordillera, Gradstein et al. 6743 (U). TUNGURAGUA: Baños, Spruce s.n. (G). GALÁPAGOS: Isabela, Cerro Azul, Gradstein & Sipman H 408 (PC, U), H 437 (U). PERU. Mt. Campana, Spruce s.n. (G); AMAZONAS: Chachapoyas, between Chachapoyas and Cajamarca, above Leimebamba, Frahm et al. 877 (PC). CAJAMARCA: Contumaza, Tambo de Lima, between Contumaza and Cascas, P. & E. Hegewald 7382 (PC). HUANUCO: Leoncido Prado, Tingo Maria, P. & E. Hegewald 7817, 7841 (PC); Leoncido Prado, Tingo Maria, Cuevas de las Pavas, P. & E. Hegewald 7789, 8339 (PC). SAN MARTÍN: Tarapoto, Cumba-Kiwi, Spruce s.n. (G); Rioja, between Chachapoyas and Moyobamba, Frahm et al. 87, 101, 689, 1592 (PC), 429, 1591 (U); Lamas, between Yurimaguas and Tarapoto, Frahm et al. 1696 (PC, U).

Bischler et al. 173 BRAZIL. S. loc., Weddell s.n. (PC). PARANÁ: Cataractas de Iguazu, 20 km of Foz de Iguazu, Vitt 21322 (U). RIO JANEIRO: Glaziou 7239 (PC); 1823, Gaudichaud s.n. (PC); Petropolis, Volk 83-623 (PC). SANTA CATARINA: S de Espigão, 22 km NW Dr. Pedrinho, on road BR 477, Frahm 1701 (U). SÃO PAULO: Salto dos Treis Ranjos waterfall, near Cerqueira-Cesar, Schiffner 4275 (G, PC). BOLIVIA. BENI: Prov. Ballívian, Serranía del Pilón Lajas, Gradstein 7155 (U). LA PAZ: Inquisivi, 1.5 km S of Comunidad Phullchiri, Lewis 83-3551 (F); Inquisivi, 2 km NW of Licoma Pampa, Lewis 83-3642 (F); Inquisivi, 7 km SE Miguillas, Lewis 86-1472 ( F); Rio Miguillas, Lewis 86-611 (F); Inquisivi, 3 km E Licoma Pampa, Lewis 86-1002 (F); Larecaja, 8 km NE of Consata, Lewis 83-390 (F); N Yungas, 11 km NW Coroico, Lewis 86-1753 (F); N Yungas, 20 km N Caranavi, Lewis 86-1975 (F); N Yungas, Cerro Uchumani, Lewis 83-2708 (F). SANTA CRUZ: Vallegrande, Cerro El Centinela, Lewis 85-616 (F). TARIJA: Acre, 2-5 km NW of Comunidad La Mamora, Lewis 84-2232, 2240 (F); Acre, Comunidad Sidras, Lewis 84-2316, 2334 (F); Acre, Loma Las Lagunas, Lewis 84-2335 (F); O'Connor, Abra Castellón, 7 m NW of Entre Ríos, Gradstein 1671 (U). PARAGUAY. Cerro León, Balansa 3711 (G, PC); Dep. Guaira, S Colonia Independencia, Cordillera del Ybytyruzu, Tacuara, Geissler 15104 (G); Dep. Central, S Colonia Independencia, Cordillera Ybytyruzu, N Cerro Guayaqui, arroyo Tacuaro, Geissler 15037 p.p. (G); Mbatobi near Paraguari, Balansa 3620 (G); Dep. Paraguari, Salto Cristál, Spichiger & Loiseau 1606 (G); Dep. Alto Paraná, 35 km N Hernandarias, Reserva Biologica Itabo, Salto Syi, Geissler 14584 (G); Dep. Alto Paraná, 80 km N Hernandaria, Reserva Biologica Limoy, near R. Limoy, Geissler 14678 (G); Dep. Alto Paraná, Arboretum, Geissler 14711 (G); Dep. Paraguari, Cordillera de los Altos, Jololo, Geissler 15179 (G); Dep. Amaby, W Pedro Juan Caballero, arroyo Naranja-hai, Parque Nacional Cerro Cora, Geissler 14838 (G). ARGENTINA. TUCUMÁN: Tafi, San Javier, Kühnemann 3107 (PC); Siambon, 1873, Lorentz s.n. (G); trail to Tafi Viejo, Nuñez 10525 (U); Sierra de Medina, Vianna s.n. (BA). URUGUAY. Montevideo, San Gabriel, 1833, Carbon s.n. (PC). Dumortiera hirsuta probably is a complex species, comprising several genetically distinct siblings. Three cytotypes are known, but chromosome counts are not available for neotropical material. Important genetic differentiation has been found within the haploid cytotype, but the large distribution of the species and its rarity in Europe and the Mediterranean precluded a thorough investigation of its genetic structure. Awaiting further research, all neotropical specimens have been attributed to Dumortiera hirsuta sensu lato, thus including D. nepalensis (Tayl.) Nees (Evans, 1919a). This species, treated as a subspecies by Schuster (1992b), is characterized by crowded, papilliform cells on the dorsal surface of the thallus that correspond to remnants of air chamber partitions. Dumortiera hirsuta is easily distinguished from most neotropical Marchantiales by its large, dark green thallus without epidermal pores and with "bristles" on the ventral side. However, in sterile condition the species may be difficult to distinguish from Monoclea gottschei (see under the latter

Bischler et al. 174 species).

18. MARCHANTIA L., Spec. Pl. ed. 1, 1137. 1753; emend. Raddi, Opusc. Sci. (Bologna) 2: 358. 1818. Type species (Léman, 1823). Marchantia polymorpha L. (lectotype). Genus dedicated to N. Marchant (...-1678), Director of the Garden of Duc Gaston d' Orléans. Thallus 2.7-20 mm wide, often tinged with purple. Branching dichotomous. Epidermal pores compound, with 4-7 rings of cells, radial walls thin. Air chambers in one layer, with chlorophyllose filaments. Basal tissue with oil-cells, often with sclerotic cells and mucilage cavities. Rhizoids smooth and tuberculate. Ventral scales in 4-10 rows, with oil-cells and marginal papillae, each pair of rows of different shape, the two median with large appendage. Asexual reproduction by discoid gemmae with 2 growing points. Gemma cups cup-shaped. Dioecious, rarely with male branches developing in the female receptacles and then monoecious. Antheridiophore terminal; stalk with 0-2 assimilatory strips, 2-4 rhizoid furrows and scales, sometimes with modified ventral thallus scales sheathing the base; receptacle peltate or palmate, 2-12-lobed, with compound pores on dorsal side and scales underneath. Archegoniophore terminal; stalk with 1-2 assimilatory strips, 2-4 rhizoid furrows, and scales, sometimes with modified ventral thallus scales sheathing the base; receptacle 2-13-lobed, with compound pores on dorsal side. Involucres bilabiate, alternating with the receptacular lobes in neotropical species. Archegonia 2-14 per archegonial cavity, each fertilised archegonium enclosed within a campanulate pseudoperianth. Calyptrae 2-3-layered after fertilization. Sporophytes several per involucre, maturing sequentially; mature sporophyte with slightly elongating seta; capsule splitting into irregular valves to 1/2 of capsule length, walls with annular thickenings; spore : elater ratio more than 4 : 1. Spores 5000 to more than 10 000 per capsule, 8-36 µm diameter, distal face either with tuberculate areoles, or with ridges and depressions, or irregularly pitted or grooved, or vermiculate, proximal face usually irregularly pitted or vermiculate, trilete scar more or less distinct. Elaters with 2-3 helical bands. Gametophytic chromosome number n = 9 or 18. Marchantia is a genus of worldwide distribution, comprising 36 species. They occur predominantly in humid-temperate, subtropical, and tropical areas without pronounced dry seasons. Some species reach the Arctic and the Antarctic. All are terrestrial and grow on bare or rocky soil, exposed or shaded by open vegetation. They avoid the shaded understory of forests but are common at forest edges and along roads inside the forests. They are indifferent to edaphic conditions (type of rocks, or soil, or pH), but are not drought-tolerant; the substratum has to be permanently moist. Newly opened habitats where space is available, especially open, ruderal and disturbed, man-made sites are frequently colonized. The distributions of some taxa seems to be limited by high or low temperatures, but the availability of water and free space are the main conditions determining their geographical ranges. They grow from sea level up to 4500 m but are rare in equatorial lowland areas. Marchantia species are among the most common thallose liverworts in the Neotropics. The genus

Bischler et al. 175 is represented by two (out of three) subgenera and eight species. Four of them have large ranges, including more than one continent, the four others are known from the New World alone. Marchantia foliacea Mitt. from southern Chile, Juan Fernandez, Australia, and New Zealand does not occur in the Neotropics and has been excluded from this treatment. Invalid names are not cited among the synonyms of the species. They are listed in Bischler (1984).

KEY TO THE NEOTROPICAL SUBGENERA AND SPECIES OF MARCHANTIA 1. Thallus margins crisped-plicate, lobulate, rarely entire. Ventral scales in 4-8(-10) rows, extending over 75-100% of thallus width. Laminal scales wider than long, without apical papillae. Receptacle of archegoniophore deeply dissected into 9-11 terete rays. Spores (9-)10-16 µm diameter ....................................................................................................................... 18a. subgen. Marchantia..2 2. Thallus margins lobulate. Ventral scales in 4-6 rows; marginal scales, if present, not extending beyond thallus margins. Appendages of median scales bordered by very small cells, margins entire to crenulate .................................................................................................................1. M. berteroana 2. Thallus margins entire or crenulate. Ventral scales in 6-8(-10) rows; marginal scales extending beyond thallus margins. Appendages of median scales usually sharply toothed, rarely entire, marginal cells slightly smaller than inner cells.................................................................................... 3 3. Thallus and gametangiophores tinged with black or dark red. Ventral scales in 6-8(-10) rows. Laminal and marginal scales dark red, with strongly arched insertion, curved away from thallus. Margins of appendages of median scales toothed, at least some teeth 2-celled. Female receptacle without papillae on rays................................................................................................2. M. plicata 3. Thallus and gametangiophores green, sometimes tinged with purple. Ventral scales in 6 rows. Laminal and marginal scales hyaline or light red, with hardly arched insertion, appressed to thallus. Margins of appendages of median scales with unicellular teeth, rarely almost entire. Female receptacle with papillae on rays................................................... ...........3. M. polymorpha 1. Thallus margins entire. Ventral scales in 4 rows, extending over 25-60% of thallus width. Laminal scales as long as or longer than wide, with apical papillae. Receptacle of archegoniophore shallowly to deeply divided into 5-11 apically flattened lobes. Spores 19-36 µm diameter ................................ .....................................................................................................................18b. subgen. Chlamidium 4. Inner opening of epidermal pores with straight or convex inner walls, not cruciate (inner pore opening is not completely closed). Appendage of median scales without, rarely with 1-2 oil-cells. Gemma cups ciliate or nearly entire, without papillae on outer side. Male receptacle palmate. Female involucres entire or ciliate......................................................................................... .............5

Bischler et al. 176 5. Margin of appendage of median scales irregularly angular. Cilia of gemma cups 2-5(-6) cells long. Female receptacle shallowly divided into 5 or (7-) 9-11 lobes. Cilia of involucres 3-8 cells long ................................................................................................................................................ 6 6. Thallus margin with thin-walled cells. Inner opening of epidermal pores with strongly convex inner walls. Appendage of median scales orbicular to ovate (length : width ratio = 11.2), not acuminate, rounded or bluntly acute apically, without oil-cells. Male receptacle without papillae on dorsal side. Archegoniophore stalk with single band of air chambers; female receptacle nearly symmetric, (7-)9-11-lobed ...................... .................4. M. breviloba 6. Thallus margins with thick-walled cells. Inner opening of epidermal pores with straight or hardly convex inner walls. Appendages of median scales broadly triangular (length : width ratio = 1.3-2.3 : 1), acute and acuminate apically, often with 1-2 oil-cells. Male receptacle with numerous papillae on dorsal side. Archegoniophore stalk with 2 bands of air chambers; female receptacle strongly asymmetric, 5-lobed ........................................... . 5. M. chenopoda 5. Margin of appendage of median scales sharply toothed. Gemma cups with entire margin, or with cilia 1-2, rarely 3-4 cells long. Female receptacle deeply divided into 5-9(-11) lobes. Involucres crenulate, or with short cilia, 1-2 cells long................................................................. 7 7. Thallus without distinct median band on dorsal side, margins hyaline. Gemma cups ciliate, with 1-3(-4)-celled cilia; lobes of female receptacle flat basally, hardly broadened apically. Margins of involucre with 1-2-celled cilia..............................................................6. M. inflexa 7. Thallus with narrow, dark median band on dorsal side, margins usually purplish. Gemma cups crenulate, or with scattered, 1-2-celled cilia; lobes of female receptacle convex basally, broadened apically. Margins of involucre crenulate ........................................... 8. M. papillata 4. Inner opening of epidermal pores cruciate, with inner pore opening completely closed. Appendage of median scales with numerous oil-cells. Gemma cups with ciliate lobes and papillae on outer side. Male receptacle peltate. Female involucres with ciliate lobes. ....... .......7. M. paleacea

18a. MARCHANTIA subgen. MARCHANTIA. Marchantia sect. Astromarchantia Nees, Naturgesch. Eur. Leberm. 4: 60, 61. 1838 (nom. illeg.). Marchantia Stellatae Gottsche, Lindenb. & Nees, Syn. hepat 522. 1846 (without rank). Marchantia Marchantiotypus Dumort., Bull. Soc. Roy. Bot. Belgique 13: 150. 1874 (without rank). Type species. Marchantia polymorpha L. Thallus margins crisped-plicate or lobulate, often crenulate. Epidermal pores with cruciate inner opening. Basal tissue without sclerotic cells. Ventral scales in 4-8(-10) rows, extending over 75-100 % of thallus width; appendage of median scales ovate, orbicular or reniform, with several oil-cells; margins entire, crenulate or toothed. Laminal scales wider than long, rounded apically, without apical papillae; cell walls towards apex usually with thickened angles. Marginal scales, when present, ovate,

Bischler et al. 177 rounded apically, cell walls towards apex with thickened angles. Gemma cups with ciliate lobes and papillae on outer side. Stalks of gametangiophores sheathed basally by conspicuous scales, these scales rounded apically, without appendages. Male receptacles peltate, without papillae on dorsal side, shallowly divided into wide, rounded lobes. Female receptacles deeply dissected into 9-11 terete rays. Involucres with ciliate lobes. Spores 10-16 µm diameter; spore-wall vermiculate. The subgenus comprises three species, among the largest of thallose liverworts. It is distributed from Svalbard and Franz Josef Land (81° N) in the northern hemisphere, to the Antarctic islands and the Antarctic Peninsula in the southern, and is common in the temperate zones with extensions into the subtropical and tropical areas on islands and along mountain ranges at higher elevations. In the Neotropics, the three species occur in permanently moist or wet habitats in high mountains, mostly above 2000 m.

1. Marchantia berteroana Lehm. & Lindenb., in Lehmann, Nov. Stirp. Pug. 6: 21. 1834. Type (Bischler, 1984). Chile. Juan Fernandez: Bertero s.n. (lectotype, W; isotypes, FH, G, NY, PC, STR, W).

Figs. 143, 144, 160: E

Marchantia berteroana var. biflora Nees, Lindenb. & Gottsche, in Meyen, Nov. Actorum Acad. Caes. Leop.-Carol. Nat. Cur. 19, suppl. 1: 476. 1843. Type. Chile. Valparaiso: Quebrada S. Augusto and Sierra de S. Francisco de Valparaiso, Meyen s.n. (syntypes, STR, W). Marchantia berteroana var. γ anactis Gottsche, Lindenb. & Nees, Syn. hepat 525. 1846. Type. Chile. Juan Fernandez: Bertero s.n. (n.v.); St. Helena, Ludwig s.n. (syntypes, FH, W). Marchantia cephaloscypha Steph. var. breviradiata A.Massal., Atti Reale Ist. Veneto Sci. 87: 242. 1928. Type. Argentina. Staten Isl.: Port Cook and Penguin-Rockery, Spegazzini s.n. (syntype, G). Marchantia berteroana var. polylepida Herzog, Hedwigia 74: 79. 1934. Type. Colombia. Cundinamarca: Bogotá, Troll 2087 (holotype, JE). Marchantia discoidea Steph., in Bonner, Candollea 14: 105. 1953. Type. Colombia. Mayor 57 (holotype, G). Thallus bluish or yellowish green, tough and leathery, often pigmented with purple or brown, usually 7.1-11 mm wide, without distinct median band on dorsal side; margins hyaline, purplish or brown, crisped-plicate and irregularly lobulate. Epidermal pores usually 46-73 µm diameter, inner opening cruciate. Basal tissue with few or without mucilage cavities. Ventral scales in 4-6 rows, laminal scales appressed to thallus, with hardly arched insertion, marginal scales, if present, not extending beyond thallus margins; appendages of median scales hyaline or purplish, ovate to orbicular, usually 21-28 cells wide, rounded apically; margins entire or slightly crenulate, bordered by 1-3 rows of very small, subquadrate cells, usually 10-22 times smaller than the inner cells; laminal and marginal scales hyaline or light red; marginal scales numerous, few, or absent. Gemma cup lobes

Bischler et al. 178 with row of 3-10 thick-walled cells apically; papillae on outer side 1-3-celled. Antheridiophore stalk without band of air chambers, with two rhizoid furrows; receptacle almost symmetric, with 6-8 (-9) lobes. Archegoniophore stalk with single, broad band of air chambers and 2 rhizoid furrows; receptacle almost symmetric, dorsal side with rounded projection; rays sometimes clavate apically, without papillae. Involucres hyaline, seldom light red. Spores yellow to brownish, (9-)11-12 µm diameter.Gametophytic chromosome number n = 9. Distribution and ecology. The species has a circumantarctic distribution, with extensions in high mountain areas in Oceania (New Guinea), and in the Neotropics in the Andes, to the Galápagos Is. and Costa Rica. It has not been recorded from continental Ecuador, Peru and Bolivia. Its distribution includes Costa Rica, Colombia (Antioquia, Boyacá, Cauca, Cundinamarca, Magdalena (Winkler, 1976), Meta, Santander), Venezuela (Mérida), Ecuador (Galápagos Is.: Sta Cruz), Brazil (Minas Gerais (Gradstein & Pinheiro da Costa, 2003), Rio de Janeiro, Rio Grande do Sul (Vianna, 1970), Chile (Aisén, Araucania, Los Lagos, Magellanes, Valparaíso), Juan Fernandez Is., Argentina (Buenos Aires, Chubut, Córdoba, Neuquén, Rio Negro, San Luís (Hässel, 1963), Tierra del Fuego, Tucumán (Hässel, 1963)), Uruguay (Hässel, 1964), and the Antarctic (Staten Is., Falkland Is., South Georgia, South Sandwich Is. (Grolle, 1972b), Antarctic Peninsula (Corte, 1962)). The species grows on humid or wet soil, or soil over rocks (lava, schist), in seepage areas, or on road banks, sometimes in marshes, often in burnt areas, exposed or shaded by open vegetation. It has been recorded from 2000 to 4500 m but occurs at lower elevations on islands and in the southern part of its range. It is less common than the two other species of the subgenus. Selected neotropical specimens examined. COSTA RICA. S. José, Irazu, Pittier 12 (G); Alto del Poas, Pittier 6021 (G); Panamerican road, 1976, Melendez s.n. (PC); Cerro Muerte, Schulthes 12032 (FH). COLOMBIA. ANTIOQUIA: Type of M. discoidea; Aguas Termales, 1874, Wallis s.n. (G). BOYACÁ: Páramo de la Rusia, Cleef 6885 (G, U). CAUCA: Mt. Puracé, Killip 6793 (FH, JE); Silva, Luteyn et al. 10125 (NY). CUNDINAMARCA: Type of M. berteroana var. polylepida; Chisacá, Cleef 5272 (PC, U); San Cayetano, Laguna de la Bessia, Cleef 6558 (U). META: Páramo de Sumapáz, Cleef 7567 (U). SANTANDER: Páramo Rico, Killip & Smith 17869 (FH); near Las Vegas, Killip & Smith 15914 (JE). VENEZUELA. MÉRIDA: without locality, Moritz 134 (G). ECUADOR. Galápagos Is.: Santa Cruz, Mt. Crocker-Puntudo area, Gradstein & Weber H 126 (G, U). BRAZIL. RIO DE JANEIRO: Parque Nacional Itatiaia, Frahm 1622 (PC). CHILE. VALPARAISO: Type of M. berteroana var. biflora. JUAN FERNANDEZ: Type; syntype of M. berteroana var. γ anactis. MAGELLANES: I. Riesgo, Seno Skyring, Fulford 360 (Hb. Fulford).

Bischler et al. 179 ARGENTINA. BUENOS AIRES: Qilmes, Hässel 9351 (JE). CÓRDOBA: Copina, Castellanos 586 (PC). NEUQUEN: Lago F. Frias, Perez Moreau 4602 (JE, PC). STATEN ISLAND: Type of M. cephaloscypha var. breviradiata. The presence or absence of marginal scales is subject to variation in all parts of the geographical range of the species. The lobulate thallus margin, the entire and bordered (by very small cells) appendage of the median scales, and the absence of any distinct median band on the dorsal side of the thallus separate M. berteroana from the two other species of the subgenus.

2. Marchantia plicata Nees & Mont., in Montagne, Ann. Sci. Nat., Bot. sér. 2, 9: 43. 1838. Type (Bischler, 1984). Bolivia. La Paz: Yungas, between Chupé and Ianacaché, d'Orbigny s.n. (lectotype, PC; isotypes, G, JE).

Figs. 145, 146

Marchantia lamellosa Hampe & Gottsche, in Gottsche, Lindenb. & Nees, Syn. hepat. 527. 1846. Type (Bischler, 1984). Venezuela. Mérida: Páramo de Mucuchies, Moritz s.n. (lectotype, BM; isotypes, G, W). Thallus green or yellowish-green, pigmented with black or dark red on margins and ventral side, stalks, margins of male receptacles and margins of involucres, usually 7.7-18.3 mm wide and up to 30 cm long, without distinct median band on dorsal side; margins dark red or black, entire or crenulate. Epidermal pores usually 50-84 µm diameter, inner opening cruciate, or cells with strongly convex inner walls, rarely almost straight. Basal tissue without mucilage cavities. Ventral scales in 6-8(-10) rows, laminal and marginal scales dark red , curved away from thallus, with strongly arched insertion, marginal scales extending beyond thallus margins; appendage of median scales purplish, rarely hyaline, ovate to orbicular, usually 15-23 cells wide, usually rounded, sometimes acute or shortly apiculate apically, margins toothed, with 1-2-celled teeth, marginal cells usually 3.9-5.9 times smaller than inner cells; laminal and marginal scales purplish, sometimes hyaline; laminal scales in several rows, alternating irregularly with median scales, irregular in shape; marginal scales numerous. Gemma cup lobes with a row of 3-7 thin-walled cells apically, papillae on outer side 1-2-celled. Antheridiophore stalk without band of air chambers, with 2(-4) rhizoid furrows; receptacle almost symmetric or slightly asymmetric, (6-)8-10-lobed. Archegoniophore stalk with single, broad band of air chambers and 2(-4) rhizoid furrows; receptacle symmetric or slightly asymmetric, dorsal side with rounded projection, rays often clavate apically, without papillae. Involucres hyaline, purplish or tinged with black. Spores yellow or brownish, usually 14-16 µm diameter. Gametophytic chromosome number n = 9. Distribution and ecology. The species is known only from high elevations in the Andes (20° N to 34°30' S). Its range extends from Mexico (Mexico, Puebla (Hässel, 1963)), Guatemala, Costa Rica,

Bischler et al. 180 Panama (Stotler et al., 1998), Colombia (Boyacá, Caldas, Caquetá, Cauca, Cundinamarca, Chocó, Magdalena, Meta, Santander, Tolima), Venezuela (Mérida), Ecuador (Azuay, Chimborazo, Cotopaxi, Imbabura, Loja, Napo, Pastaza, Pichincha, Tungurahua), Peru (Amazonas, Ancash, Apurimac (Hässel, 1963), Ayacucho, Cajamarca, Cuzco, Huanuco, Huaráz, Junín, La Libertad, Lima, Pasco, San Martín), Bolivia (Cochabamba, La Paz), to Argentina (Catamarca (Hässel, 1963), San Luís (Hässel, 1963), Tucumán). The species has been collected on soil and soil over rocks (volcanic and calcareous), in stonework, in seepage areas, on road banks, in marshes and páramos, at water borders, exposed or shaded by open vegetation, between 600 and 4500 m, but is most common between 2000 and 4000 m. Marchantia plicata has been recorded as the world’s largest terrestrial liverwort from the surroundings of Quito, Ecuador, where it grows abundantly on volcanic soil on steep, cut road banks at about 3000 m, forming huge rosettes of up to 75 cm in diameter (Gradstein, 1989). Selected specimens examined. MEXICO. MEXICO: Lagunas de Zempoala, 14 km W Tres Cumbres, Eggers & Frahm s.n. (PC). GUATEMALA. San Marcos, NW slopes of volcán Tajumulco, Steyermark 36894 (Hb. Fulford); 6 km N of Quiche, Volk 6/715 (JE). COSTA RICA. Volcán Barba, Valerio 24 (JE). COLOMBIA. BOYACÁ: Nevado del Cocuy, Ritacuba, Bischler 2741 (PC). CALDAS: Nevado del Ruiz, Bischler 294 (PC); between Manizales and Termales, Bischler 378, 379 (PC). CAQUETÁ: between Garzon and Florencia, Mason 13965 (UC). CAUCA: S.loc., Luteyn 10072 (NY). CUNDINAMARCA: Bogotá, Boquerón, 1851-1857, Triana s.n. (G, W). CHOCÓ: between Quibdo and Bolivár, Bischler 205 (PC). MAGDALENA: Santa Marta, Schlim 1046 (PC). META: Páramo de Sumapáz, Cleef 8260 (PC, U); Chisacá, Río S. Rosa, Cleef 5273 (U). SANTANDER: Río de la Baja, Killip & Smith 18311 (FH, JE). TOLIMA: between Cajamarca and Armenia, Bischler 253 (PC); Manizales-El Fresno, Páramo de las Letras, Pinto 438 (PC, U). VENEZUELA. MÉRIDA: Type of M. lamellosa; s.loc., Luteyn et al. 9795 (NY); El Gavilán, páramo Timotes, Onraedt 78.V.5245 (PC). ECUADOR. San Juan, Benoist 2501 (PC). AZUAY: Cuenca, Harling 2167 (JE). CHIMBORAZO: SW Chimborazo, 1903, Meyer s.n. (JE); 15 km N of Alausi, Gradstein et al. 3368 (U). COTOPAXI: Near Sindipampa, Arts 23/021 (PC). IMBABURA: Lago San Marcos, Cayambe Mt., Cazalet & Pennington 71 (NY). LOJA: San Lucas, Lehmann 5 (W). NAPO: near Cosanga, Baeza-Tena, El Paso del Guacamayo, Davis 510 (NY). PASTAZA: River Pastaza, Spruce s.n. (MANCH). PICHINCHA: Volcán Pululahua, Fantz B301 (NY); Quito, Jameson s.n. (FH); 3 km W of Lloa, south slope of Volcán Pichincha, Steere & Balslev 26058 (NY). TUNGURAHUA: Mt. Tunguragua, Spruce s.n. (G, MANCH).

Bischler et al. 181 PERU. AMAZONAS: Vicinity of Leimebamba, Frahm et al. 1033 (PC). ANCASH: Huaráz, P. & E. Hegewald 7475 (Herb. Hegewald); NE Huaráz, laguna Llaca, Geissler 8857 (G). AYACUCHO: Cangallo, between Chumbis and Ocros, P. & E. Hegewald 8959 (PC). CAJAMARCA: Rio Chucsen, between Matara and Namura, P. & E. Hegewald 6149 (PC). CUZCO: Urubamba, Machu Pichu, P. & E. Hegewald 5607 (PC). HUANUCO: W summit of Carpish, Stork & Horton 9901 (UC). HUARÁZ: Laguna Llaca, NE Huaraz, Geissler 8878 (PC). JUNÍN: Huancapistana, Killip & Smith 24276 (FH). LA LIBERTAD: Otuzco, Casmiche, P. & E. Hegewald 6007 (Herb. Hegewald). LIMA: Huarochiri, Rio Rimac, Zarate, Hutchison & Wright 7089 (UC). PASCO: Cerro Pasco, Hooker s.n. (FH). SAN MARTÍN: Rioja, Moyobamba-Chachapoyas, Frahm et al. 549 (PC); Rioja, Olmos-Moyobamba km 403, Geissler 8678 (G). BOLIVIA. COCHABAMBA: Arani, 14 km S Tiraque, Lewis 85-100 (F); Carrasco, 7 km W of Rancho Pairumani, Lewis 85-325 (F). LA PAZ: Type; between Chupé and Yanacaché, d'Orbigny 209 (G, JE, P); Zongo, Bang 910 (FH); Cañon La Paz, Shepard s.n. (F); Cerro La Paz, Shepard 155 (F); Inquisivi, 3 km W of Quime, Lewis 83-3406 (F); Larecaja, 16-17 km N of Sorata, Lewis 83-227 (F); S Yungas, 8 km NE of summit of Nevado Illamani, Lewis 83-2457A, 2412 (F). ARGENTINA. TUCUMÁN: Anfama, La Cienaga, Castellanos 10343 PC). The shape of the inner opening of the epidermal pores is usually cruciate in Marchantia plicata, but sometimes the inner walls are nearly straight. The colour of the ventral scales varies from purplish to hyaline tinged with black purple. Marchantia polymorpha, a closely related species, occurs in similar habitats and can grow intermixed with M. plicata. It is not tinged with black purple, the number of rows of scales is of six, and the laminal and marginal scales are appressed to the thallus. In addition, the appendages of the median scales in M. plicata often have two-celled marginal teeth and the rays of the female receptacle are devoid of papillae.

3. Marchantia polymorpha L., Spec. Pl. ed. 1, 1137. 1753. Lectotype (Bischler & BoisselierDubayle, 1991). Europe, Dillenius, Hist. musc. tab. 76 f. 6E-6F, 1742 (typotype, Dillenius, fol. 166 f. 6, OXF).

Figs. 2: H-I, 147, 148

Thallus light or yellowish green, sometimes tinged with purple, usually 6.6-10.9 mm wide, often with wide, blackish median band composed of elongate air chambers with black borders, or lacking air chambers; margins purplish or hyaline. Epidermal pores usually 43-78 µm diameter, inner opening usually cruciate. Basal tissue sometimes with mucilage cavities. Ventral scales in 6 rows, laminal and marginal scales hyaline or light red, appressed to thallus, with hardly arched insertion, marginal scales extending beyond thallus margins; appendage of median scales hyaline, or hyaline with purplish borders, rarely purplish, orbicular to reniform, usually 14-23 cells wide, usually rounded apically, margins usually with sharp, unicellular teeth, sometimes nearly entire, the marginal cells usually 2.17.7 times smaller than inner cells; laminal and marginal scales hyaline or light red. Gemma cup lobes

Bischler et al. 182 with row of 3-7 thin-walled cells apically, papillae on outer side 1-3-celled. Antheridiophore stalk without band of air chambers, with two rhizoid furrows; receptacle almost symmetric, with 6-8(-10) lobes. Archegoniophore stalk with one band of air chambers and 2(-4) rhizoid furrows; receptacle nearly symmetric, dorsal side with rounded median projection, rays sometimes slightly clavate, with numerous, unicellular papillae. Involucres hyaline or purplish. Spores yellow, 11-13 µm diameter. Gametophytic chromosome numbern = 9. Distribution and ecology. The species is nearly cosmopolitan, distributed worldwide in temperate and cold areas of the northern and southern hemisphere (80°40' N to 63° S), on continents as well as on islands. However, it has not been recorded below 1000 m in continental tropical areas. In the north, the species has been recorded from all states of U. S. A., from Canada and Greenland. Its neotropical distribution includes Mexico (D. F., Chiapas, Hidalgo, Mexico, Michoacán, Oaxaca (Evans, 1917), Puebla, San Luis Potosí, Veracruz), Guatemala, El Salvador, Costa Rica (Morales, 1991), Panama, Bermuda, Cuba, Jamaica, Haiti, Colombia (Boyacá, Cauca, Cundinamarca, Magdalena (Winkler, 1976), Nariño, Norte de Santander, Santander, Tolima), Venezuela (Caracas, Mérida), Ecuador (Chimborazo, Pichincha, Tunguragua, Galápagos Is. (Isabela, Gradstein & Weber, 1982)), Peru (Cajamarca, Cuzco, Junín, Lima (Carillo & Chanco, 1971), Puno (Soukoup, 1955)), Brazil (Rio Janeiro, Rio Grande do Sul (Vianna, 1970)), Bolivia (Cochabamba, La Paz), Chile (Aisén (Hässel, 1963), Araucania (Hässel, 1963), Los Lagos (Hässel, 1963), Magellanes, Santiago, and Juan Fernandez), Argentina (Buenos Aires, Córdoba, Chubut, Mendoza (Hässel, 1963), Neuquén, Rio Negro, Santa Cruz (Hässel, 1963), Santa Fé (Hässel, 1963), Tierra del Fuego, Tucumán (Hässel, 1963)), Uruguay and the Antarctic Islands. It has not been recorded from the Leeward and Windward Islands. The species colonizes moist or wet, sandy or clayey soil, soil over rocks (limestone, sandstone, granite) and burnt soil, seepage areas, montane marshy grasslands including intermittently flooded zones, road banks, ruderal sites or other disturbed zones, exposed or sheltered by open vegetation. It is a common intruder into gardens, cultures and greenhouses, introduced with garden soil and plants. Its neotropical occurrence in natural habitats is most frequent between 2000 and 4000 m, but it can be found in man-disturbed sites, on islands and in its southern range below 2000 m. Selected neotropical specimens examined. MEXICO. CHIAPAS: Mun. Siltepec, above Colonia El Rosario, Breedlove 6885 z (MEXU); Velaso, Omitlan, Sanchez Mejorado 113 (MEXU). D. F. : Aqueducto de los Leones, Maury 3121 (PC); Popocatepetl, Sanchez Mejorada 1 (MEXU); Desierto de los Leones, 1944, Ruiz Oronoz s.n. (MEXU); Parque Ejidal San Nicolas, Totolapan, Long 29564 (E). HIDALGO: Chapulhuacan, Frye & Frye 2848 (FH); Zacaulipan, Poza Obscura, Sanchez Mejorado 285 (MEXU). MEXICO: Popocatepetl, Düll 364 (PC). MICHOACAN: Morelia, Frye & Frye 3092 (G). PUEBLA: Sta Barbara, Nicolas 5102 (PC). SAN LUIS POTOSÍ: Tamazunchale, Frye & Frye 2997 (G). VERACRUZ: Orizaba, Smith s.n. (FH); Cofre de Perote, Balls B 4637 (UC). GUATEMALA. Finca Carmona, SE of Antigua, Sacatepequez, Standley 63692 (FH); Fuentes

Bischler et al. 183 Georginas, volc. Zunil, Quezaltenango, Standley 67420 (FH); Quezaltenango, near Zunil, Standley 67420 (FH); SE Antigua, Sacatepequez, Standley 63692 (FH). EL SALVADOR. Chalatenango, E slope of Los Esesmiles, Tucker 1019 (G, PC, UC). PANAMA. Alto Pineda, Salazar Allen 1108 (PMA); El Aguacatal, Volcán, Salazar Allen 1178 (NY, PMA); Finca de la Fortuna, Alto Quiel, Boquete, Salazar Allen 775 (PMA). BERMUDA. Hamilton, Haynes s.n. (FH). CUBA. Guantanamo, La Prenda, Hioram & Maurel 5118 (NY); Sierra Maestra, Cumbre Bayamesa, Lopez Figueiras 2121 (JE). JAMAICA. Hardware Gap, Evans 176 (FH). HAITI. La Selle, Croix-des-Bouquets, Ekman 7700 (Herb. Fulford). COLOMBIA. BOYACA: near Páramo Chita, Aguirre & Gradstein s.n. (PC); Nevado del Cocuy, Bischler 2833 (PC); Páramo de Pisba, Cleef 4337 (MEXU). CAUCA: Volcán Puracé, Cleef & Fernandez 633 (U); Páramo de las Papas, Bischler 715 (P). CUNDINAMARCA: Sabana de Bogotá, Schulthes 11107 (FH); Chisacá, Bischler 2336 (PC); Bogotá, Triana s.n. (G). NARIÑO: Pasto, Espinosa 4020 (Herb. Fulford). NORTE DE SANTANDER: Páramo Hatico, Toledo-Pamplona, Killip & Smith 20710 (FH, JE). SANTANDER: Piedecuesta-Las Vegas, Killip & Smith 15548 (FH, JE). TOLIMA: Manizales-Fresno, Pinto 439 (PC). VENEZUELA. CARACAS: Caracas, Linden 572 (NY). MÉRIDA: Páramo de los Conejos, Bernardi 1263 (NY); Trujillo, Linden 579 (PC); San Rafael de Mucuchies, Pittier 12894 (NY). ECUADOR. CHIMBORAZO: Pungala, 20 km S of Riobamba, Arts 28/008 (PC). PICHINCHA: without locality, Spruce s.n. (NY). TUNGURAGUA: Baños, Spruce K 1843 (MANCH). PERU. CAJAMARCA: Enseñada-Celendin, P. & E. Hegewald 6588 (PC); Contumaza, P. & E. Hegewald 7373 (PC). CUZCO: Paucartambo valley, Herrera 2997 (JE). JUNÍN: Tarma-Yauli, Vitt 21678 (PC). BRAZIL. RIO JANEIRO: Serra do Itatiaia, Dusen 52 (JE). BOLIVIA. COCHABAMBA: Ayopaya, 22 km W Quilacollo, Lewis 83-4405 (F); Carrasco, Zapato Rancho near Thola Pampa, Lewis 83-4437A (F); Carrasco, 17 km E Totora, Lewis 85-1577 (F); Parangani-Ayopoya, Cardenas 4807 (JE). LA PAZ: S Yungas, 1 km NE of summit of Nevado Illamani, Lewis 83-2413A (F); S Yungas, 1 km W of Estancia Totoral, Lewis 83-2389A (F). CHILE. SANTIAGO: Cerro S. Cristóbal, Mahu 6356 (JE). MAGELLANES: Port Jamine, Marivault s.n. (PC). JUAN FERNANDEZ: Masafuera, Quebrada Casas, Hatcher & Engel 639 (Herb. Fulford). ARGENTINA. BUENOS AIRES: Alrededores de Tandil, Castex & Jussieu 1211 (PC). CÓRDOBA: Cerro Uritorco, Hosseus 559 (JE). TIERRA DEL FUEGO: Road to Lapataia, Castellanos 1314 (JE,

Bischler et al. 184 PC). URUGUAY. Without locality and collector (PC). Marchantia polymorpha shows considerable morphological variation, as indicated by its specific epithet. Recent electrophoretic studies (Boisselier-Dubayle & Bischler, 1989; Bischler & BoisselierDubayle, 1991; Boisselier-Dubayle et al., 1995) have identified three genetically different races (described as subspecies), hardly distinct morphologically but ecologically differentiated. M. polymorpha subsp. polymorpha is known only from Europe, subsp. montivagans Bischl. et Boisselier from Europe, Central Asia, Canada, and the Kerguelen Is., and subsp. ruderalis Bischl. et Boisselier from Europe, Georgian SSR, Nepal, China, Japan, and Canada. Neotropical specimens have not been analysed electrophoretically. Whether these subspecies exist in the Neotropics or whether other genetically distinct elements are present in this area, remains unknown.

18b. MARCHANTIA subgen. CHLAMIDIUM (Corda) Bischl., Cryptogamie, Bryol. Lichénol. 3: 362. 1982. Chlamidium Corda, in Opiz, Beitr. Naturgesch. 647. 1829. Marchantia sect. Chlamidium (Corda) Nees, Naturgesch. Eur. Leberm. 4: 60, 101. 1838. Type. Chlamidium indicum Corda (= Marchantia chenopoda L.). Thallus margins entire, sometimes crisped. Epidermal pores with inner opening bordered by cells with straight or convex inner walls, sometimes cruciate. Basal tissue usually with sclerotic cells. Ventral scales in 4 rows, extending over 25-60 % of thallus width; appendage of median scales diverse in shape and structure, with or without oil-cells. Laminal scales as long as or longer than wide, with numerous apical papillae; cell walls towards apex not thickened. Marginal scales absent. Gemma cups with ciliate to nearly entire margins, or with ciliate lobes; outer side with or without papillae. Stalks of gametangiophores rarely sheathed basally with conspicuous scales. Male receptacles palmate, seldom peltate, with or without papillae on dorsal side. Female receptacles shallowly or deeply divided into 5-11 lobes, flat and rounded, truncate or shortly emarginate apically. Involucres entire, crenulate, ciliate, or with ciliate lobes. Spores 19-36 µm diameter; spore-wall not vermiculate. Marchantia subgen. Chlamidium comprises 20 species, five of which occur in the Neotropics where their range extends from 36° N to 35° S. In contrast to the species of subgen. Marchantia, they are more common at lower elevations (0-2500 m), even though isolated specimens were collected up to 4000 m. They often grow in drier places and endure short dry seasons.

4. Marchantia breviloba A.Evans, Trans. Connecticut Acad. Art 21: 265. 1917. Type. Jamaica. Jul 1903, Evans 175 (holotype, YU).

Figs. 149, 150

Bischler et al. 185 Thallus light to yellowish green, usually 4.6-6.6 mm wide, without distinct median band on dorsal side; margins light red or purplish, seldom hyaline, marginal cells hardly thickened. Epidermal pores usually 74-106 µm diameter, inner opening bordered by cells with strongly convex inner walls. Basal tissue with red sclerotic cells and numerous mucilage cavities. Ventral scales in 4 rows; appendage of median scales orange or purplish, seldom hyaline, ovate or orbicular (length : width ratio usually 11.2: 1), 11-18 cells wide, with lower edges often folded over, apex rounded, obtuse, bluntly acute and shortly apiculate, margins irregularly angular, or crenulate, marginal cells sinuose, usually 1.7-4.3 times smaller than inner cells, oil-cells absent; laminal scales purplish or hyaline. Cupules regularly ciliate, cilia 2-4(-6) cells long, outer side without papillae. Stalks of gametangiophores without conspicuous scales basally. Antheridiophore stalk with single, narrow, often interrupted band of air chambers or without band, with 2-4 rhizoid furrows; receptacle palmate, usually 5.9-9.3 mm diameter, deeply dissected into (2-)4-8 irregular rays, without papillae on dorsal side. Archegoniophore stalk with single, broad band of air chambers and 4(-2 ) rhizoid furrows; receptacle usually 4.5-6.9 mm diameter, concave or with very shallow median projection, nearly symmetric, shallowly dissected into (7-)9-11, apically not broadened lobes; scales of receptacle without sinuose cells apically. Involucres with 3-8 cells long cilia at the margins. Spores brown, seldom yellow, usually 30-36 µm diameter, with tuberculate areoles. Gametophytic chromosome number n = 18. Distribution and ecology. Marchantia breviloba is a species restricted to the Neotropics, where it occurs scattered between 20° N and 23° S. It has been recorded from Cuba, Jamaica, Haiti, Colombia (Boyacá, Cundinamarca, Norte de Santander), Venezuela (Mérida), Ecuador (Pichincha), Peru (Amazonas, Cajamarca, Cuzco, Junín (Bryan, 1929), San Martín), Brazil (Rio de Janeiro), and Bolivia (Cochabamba). It seems to be absent from Central America and the Leeward and Windward Islands. The species has been collected on soil and rocks, at water edges, along wet road banks, or along tracks, in tropical rainforest. It grows from 500 m to 4000 m, but mostly above 1000 m. Selected specimens examined. CUBA. Santiago de Cuba, Bisse & Lippold 15136 (JE); Loma San Juan, Hioram 12363 (HAC). JAMAICA. Hardwar Gap, Evans 175 (YU), Underwood 2234, 2251, 2262 (U, Herb. Fulford); St. Catherine's Peak, Evans 441 (FH, YU); Silver Gap to Hardwar Gap, Maxon & Killip 1224, 1231 (FH, YU). HAITI. Between Acuel and Port Salut, Ekman 284 (Herb. Fulford); Vicinity of Furcy, Leonard 4430 (YU). COLOMBIA. BOYACÁ: Sogamoso-Pajarito, Bischler 1713 (PC). CUNDINAMARCA: Fomeque, E of Bogotá, Florschütz 4508 (PC). NORTE DE SANTANDER: E slope of Páramo de Santurban, Killip & Smith 19625 (JE). VENEZUELA. MÉRIDA: without locality, Moritz 44 b, 70 (BM, W).

Bischler et al. 186 ECUADOR. PICHINCHA: without locality, Benoist 3125 (PC). PERU. AMAZONAS: Chachapoyas-Cajamarca, near Leimebamba, Frahm et al. 1033 (G, PC). CAJAMARCA: Puente Bajo , between Enseñada and Celendin, P. & E. Hegewald 6568 (PC). CUZCO: Sassaihuaman, Vargas 2205 (Herb. Fulford). SAN MARTÍN: Rioja, along road to Chachapoyas, Frahm et al. 538 (PC). BRAZIL. RIO DE JANEIRO: Serra do Itatiaia, Dusén 52 (JE); Serra do Mar, road Paratí to Cunha, Frahm 1621 (PC). BOLIVIA. COCHABAMBA: Chaparé, near Chimaré, Eyerdam 24681 (UC). The appendages of the scales of M. breviloba are similar in shape to those of M. paleacea. However, the epidermal pores are not cruciate in M. breviloba and the cupules are ciliate, not provided with ciliate lobes. Male and female receptacles yield additional distinctive characters. Marchantia breviloba is distinguished from M. chenopoda and M. inflexa by the appendages of the median scales, which are much longer than wide in M. chenopoda and coarsely toothed in M. inflexa. Marchantia breviloba is the sole neotropical Marchantia species with a duplicated gametophytic chromosome set. Sometimes, male branches develop between the lobes in the female receptacles.

5. Marchantia chenopoda L. Spec. Pl. ed. 1, 1137. 1753. Marchantia anapodocarpos Neck., Meth. Musc. 117. 1771. Type (Bischler, 1984). Martinique, s. col. (Plum. fil. 143. T. 142). Figs. 2: L, 3: B, G, 151, 152, 160: F Chlamidium indicum Corda, in Opiz, Beitr. Naturgesch. 647. 1829. Type (Bischler, 1984). Martinique, Sieber 378 p.p. (neotype, W; isoneotypes, STR, W). Marchantia swartzii Lehm. & Lindenb., in Lehmann, Nov. Stirp. Pug. 4: 9. 1832. Grimaldia swartzii (Lehm. & Lindenb.) Bisch., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 17, 2: 1034. 1835. Marchantia dillenii Lindb., Moss. Dillen. Hist. Musc. 47. 1884. Type. Jamaica (isotype, HSOL, n.v.). Marchantia cartilaginea Lehm. & Lindenb., in Lehmann, Nov. Stirp. Pug. 4: 31. 1832. Marchantia chenopoda var. cartilaginea (Lehm. & Lindenb.) Schiffn., Nova Acta Acad. Caes. Leop.-Carol. German. Nat. Cur. 60: 288. 1893. Type (Bischler, 1984). St. Vincent (lectotype, W; isotypes BM, MANCH, PC, STR). Marchantia brasiliensis Lehm & Lindenb., in Lehmann, Nov. Stirp. Pug. 4: 32. 1832. Type (Bischler, 1984). Brazil (lectotype, W; isotypes, G, STR). Marchantia martinicensis Sieb. ex Bisch., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 17, 2: 972-973. 1835. Type (Bischler, 1984). Martinique, Sieber s.n. (lectotype and isotype, W).

Bischler et al. 187 Grimaldia peruviana Mont. & Nees, in A.D.Orbigny, Voy. Amér. Mérid. 63. 1839. Marchantia peruviana (Mont. & Nees) Mont., in Gottsche, Lindenb. & Nees, Syn. hepat. 358. 1846. Type (Bischler, 1984). Bolivia. La Paz: Near Irupana, Rio de Chica, d'Orbigny 226 (lectotype, PC; isotype, G). Marchantia flabellata Hampe, Linnaea 20: 335. 1847. Type (Bischler, 1984). Venezuela. Caracas: Galipan, Moritz 47b (lectotype, BM; isotype, W). Marchantia notarisii Lehm., Nov. Stirp. Pug. 10: 22. 1857. Type. Chile. Valparaiso: Guilielmus s.n. (isotypes, G, S). Preissia mexicana Steph., Hedwigia 22: 49. 1883. Cyatophora mexicana (Steph.) Underw., Bot. Gaz. 20: 68. 1895. Type. Mexico (Bischler, 1984). Puebla: Sierra S. Pedro Nolasco, Juergensen 654 (lectotype, G; isotypes, BM, NY). Marchantia bescherellei Steph., Rev. Bryol. 15: 86. 1888. Type (Bischler, 1984). Brazil. Rio de Janeiro: Glaziou 6348 (lectotype, G; isotypes, PC, NY). Plagiochasma bolivianum Steph. ex Herzog, Biblioth. Bot. 87, 2: 173. 1916. Type. Bolivia. Cochabamba: Incacorral, Tablas, Herzog 4619 (syntype, G). Preissia barbata Herzog, Biblioth. Bot. 88: 27. 1921. Type: Bolivia. Cochabamba: Upper valley of Tocorani, 2800 m, Herzog s.n. (holotype, JE). Marchantia columbica Steph., in Bonner, Candollea 14: 104. 1953. Type. Colombia. Antioquia: Near Angelopolis, 1912, Mayor 48 (hototype, G). Marchantia pittieri Steph., in Bonner, Candollea 14: 109. 1953. Type. Costa Rica. San Francisco dos Rios, May 1891, Pittier 15563 (holotype, G). Marchantia ecuadorensis S.W.Arnell, Svensk Bot. Tidskr. 56: 342. 1962. Type. Ecuador. NapoPastaza: El Topo, Harling 3427 (holotype, S). Thallus green to light or yellowish green, usually 2.9-5.3 mm wide, without distinct median band on dorsal side; margins usually purplish, marginal cells thick-walled. Epidermal pores usually 71111µm diameter, inner opening bordered by cells with straight to slightly convex inner walls. Basal tissue with yellow or purplish sclerotic cells, with or without mucilage cavities. Ventral scales in 4 rows; appendage of median scales orange or purplish, seldom yellow, triangular (length : width ratio usually 1.3-2.3: 1); usually 7-13 cells wide, apex acuminate, acute or apiculate, margins entire, irregularly crenulate or angular, sometimes irregularly toothed with 1-celled teeth, marginal cells usually 1.3-2.9 times smaller than inner cells, oil-cells rare; laminal scales purplish or light red. Gemma cups with cilia usually 3-5 cells long, outer side without papillae. Stalks of gametangiophores without conspicuous scales basally. Antheridiophore stalk with two narrow, often interrupted bands of air chambers or without bands, and 2 rhizoid furrows; receptacle palmate, usually 4.1-7.1 mm diameter, deeply dissected into (2-)3-4(-9) rays, with numerous papillae on dorsal side.

Bischler et al. 188 Archegoniophore stalk with two narrow bands of air chambers and 2 rhizoid furrows; receptacle usually 3.7-5.5 mm diameter, strongly asymmetric, hemispherical, without median projection dorsally, shallowly dissected into 5 rounded lobes not broadened apically; scales of receptacle without sinuose marginal cells apically. Involucres with cilia usually 3-5 cells long at the margins. Spores yellowish brown, usually 25-27 µm diameter, with tuberculate areoles. Gametophytic chromosome number n = 9. Distribution and ecology. Marchantia chenopoda has a wide distribution, from 23° N to 35° S in the New World. It has been recorded from Mexico (Chiapas, D. F., Hidalgo, Jalisco, Mexico, Michoacan, Oaxaca, Puebla, Veracruz), Guatemala, Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Cuba, Jamaica, Haiti, Dominican Republic, Puerto Rico, Leeward Is. (St. Kitts), Windward Is. (Guadeloupe, Dominica, Martinique, St. Vincent, Grenada (Evans, 1917)), Colombia (Antioquia, Arauca, Boyacá, Caldas, Cauca, Chocó, Cundinamarca, Huila (Van Zanten & Gradstein, 1988), Magdalena, Meta, Nariño, Risaralda (Gradstein, 1995), Santander, Santander del Norte, Tolima, Valle), Venezuela (Aragua, Carabobo, Caracas, D. F. of Caracas, Mérida, Trujillo), Trinidad, Suriname, French Guiana (?; only one old record (Cayenne, 1821, leg. Sieber, W), might be based on erroneous label), Ecuador (Carchi, Chimborazo, Morona Santiago, Napo-Pastaza, Pastaza, Pichincha, Tunguragua, Galápagos Is. (Isabela, Sta Cruz)), Peru (Ayacucho, Cuzco, Huanuco, Junín, Loreto (Stephani, 1905), Pasco, San Martín), Brazil (Amazonas (Nees, 1933), Espíritu Santo, Mato Grosso (Vianna 1976), Minas Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul (Vianna, 1970; Oliveira e Silva & Yano 2000), Santa Catarina, São Paulo), Bolivia (Cochabamba, La Paz, Santa Cruz (Gradstein et al., 2003)), Chile (Los Lagos (Stephani 1911), Valparaíso), Paraguay, Argentina (Salta, Tucumán (Hässel, 1963)), Uruguay. Marchantia chenopoda is a common species in the Neotropics, growing on clayey and sandy soil, and on soil over rocks (sandstone, limestone, granite, schist), on water edges (streams, waterfalls, drainages), often ruderal, on roadsides, banks, in meadows, exposed or under open vegetation. It is most common below 2000 m, but has been collected up to 3300 m. Selected neotropical specimens examined. MEXICO. CHIAPAS: Tapachuela, Cerro Tres Picos, Eggers & Frahm 792743 (PC); Francisco Geomania, 1943, Villalobos s.n. (MEXU); 5 1/2 mi. SW Motozinla, on road to Huixtla, Whittemore 2199 (MEXU); Mun. Angel Albino Corzo, Xuxtepeque and Finca Cabañas, Breedlove & Daniel 71141 (MEXU). D. F.: Ciudad Universitaria, 1979, Perez de la Mothe s.n. (PC); Parque Ejidal San Nicolas, Totolapan, Long 29565 (E). HIDALGO: Chapulhuacan, Frye & Frye 50 (TENN), 2846 (NY), 2994 (G, PC, STR). JALISCO: Old La Cuesta road, 2.5 mi. W of jct. with new road, Whittemore 2145 (MEXU); along hwy 15, E of La Venta del Astillero, 8 mi. W of Guadalajara, Whittemore 2094 (MEXU); Guadalajara, Kiener 18312 (MEXU). MEXICO: Temascaltepec, near river Ixtapan, Hinton 3913 (NY). MICHOACÁN: Uruapan, vicinity of Zararacua Falls, Frye & Frye 3028 (G). OAXACA: Near La Esperanza, Düll 2/157 (JE); Barricón, Totolapan, Berendt s.n. (FH); Comaltepec, road Tuxtepec to Oaxaca, Sierra de Juarez, Relampago, 1966,

Bischler et al. 189 Calderon s.n. (MEXU); Sierra Juarez above Valle Nacional, 1962, Robinson s.n. (MEXU); Cuicatlan, Tentila, Conzatti 3835, 3837 (MEXU). PUEBLA: Type of Preissia mexicana; Tulancingo, near Huanchinango, Sharp 855 b (MEXU); Necaxa, 1948, Ruiz Oronoz s.n. (MEXU). VERACRUZ: Fortin de las Flores, Düll A 127 (JE); Orizaba, Botteri s.n. (FH); Jalapa, Pringle 15326 (MEXU, STR); Mirador, Liebmann s.n. (STR); Cordoba, along road to Orizaba, Frye 2241 (NY, TENN); near Santecompan, Delgadillo 2619 (MEXU); Mun. Banderilla, Banderilla, Ventura 637 (MEXU); Xalapa, natural forest beside Jardin Botanico, Long 29660 (E); Rio Jamapa between Huatusco and Coscomatepec, Long & Garcia 29666 (E). GUATEMALA. Alta Verapaz, E of Tactic, Steyermark 44011 (Herb. Fulford); Suchitepequez, volcán Sta Clara, Steyermark 46791 (Herb. Fulford); Solola, Santiago Atitlan, Svihla 45-689 (Herb. Fulford) HONDURAS. Comayagua, near Siguatepeque, Yuncker et al. 6410 (Herb. Fulford); Morazán, vicinity of El Zamorano, Standley 2335 (Herb. Fulford). EL SALVADOR. Chalatenango, E slope of Los Esesmiles, Tucker 1010 (G, UC); San Salvador, Standley 22684 (FH). NICARAGUA. S of Managua, Maxon et al. 7427 (JE); front of volcán Ometepe, 1893, Shimeck s.n. (Herb. Fulford) COSTA RICA. Type of M. pittieri; S. José, Pittier 6035 (G); Cartago, Turrialba, DeWolf 182 (BM); Prov. Limón, Bananito, Stork 1211 (UC); Volcán Poás, Jovet-Ast s.n. (PC); Santa Cruz, 9 km NW of Turrialba, Maas 778 (U). PANAMA. Bocas del Toro, Division Central, Valdespino 868, 870a (PMA); Boquete, Bajo Chorro, Salazar Allen 1184, 1185, 1189 (NY, PMA); Cerro Azul, Valdespino & Spörle 752 (PMA). CUBA. Oriente, S. Maestra, loma Siberia, Bisse & Lippold 13695 (JE); Santiago de Cuba, Alto de Villalón, Reyes 7457 (HAC); Santiago de Cuba, Cima Gran Piedra, 1962, Mustelier & Vicario s.n. (MEXU). JAMAICA. St. Andrews, Richards & Massey 3126 (FH); Cinchona, Evans 248 (BM, FH, PC). HAITI. Petit Bargue to Mt. Casse, Nash 523 (Herb. Fulford). DOMINICAN REPUBLIC. Prov. Vega, Zanoni et al. 19936A (NY); Mt. Barrero, Eggers 2056 (G). PUERTO RICO. Mt. Morales, Utuado, Britton & Marble 1098 (BM); Ponce to Adjuntas, Underwood & Griggs 732 (BM, FH). LEWARD IS. ST. KITTS: Mt. Misery (BM, W). WINDWARD IS. GUADELOUPE: Cambrefort, without collector 14 (PC), Baudoin s.n. (PC). DOMINICA: Roseau Valley, Armitage s.n. (MANCH); without locality, Proctor Cooper 17 (FH).

Bischler et al. 190 MARTINIQUE: Types of Chlamidium indicum and M. martinicensis; Mt. Pelée, 1977, Baudoin s.n. (PC); St. Pierre, Sources Chaudes, Sastre 7427 (PC). ST. VINCENT: Type of M. cartilaginea; path to Souffrière, Elliott 37 B (BM), Guilding s.n. (STR). COLOMBIA. ANTIOQUIA: Type of M. columbica; El Boquerón (near Medellín), Onraedt 83.A.10067 (PC); Guarne, Piedras Blancas, Onraedt 83.A.10242 (PC). ARAUCA: Sarare, Alto del Mirador, Bischler 2164 (PC). BOYACÁ: Páramo Chita-Sácama, 1982, Aguirre & Gradstein s.n. (PC). CALDAS: near Manizales, Bischler 457 (PC); between Manizales and Termales, Bischler 365 (PC). CAUCA: Monchique, Luteyn et al. 10280 (FH). CHOCÓ: Tutunendo-El Carmen, Aguirre 9042, 9043 (NY); Tutunendo, Gradstein 9038 (PC). CUNDINAMARCA: Salto del Tequendama, Smith & Idrobo 1088 (UC). MAGDALENA: Sierra Nevada de Sta Marta, Curcunnate, Martin 3565 (Herb. Fulford). META: Villavicencio, Bischler 79 (PC). NARIÑO: Ricaurte, between Pasto and Tumaco, Espinosa 2927 (Herb. Fulford). SANTANDER: between Piedecuesta and Las Vegas, Killip & Smith 15549 (JE). SANTANDER DEL NORTE: Catatumbo, Campo Oru, Bischler 2406 (PC). TOLIMA: near Ibagué, Bischler 252 (PC). VALLE: near Cali, Pichinde, Onraedt 72.A.6531, 6531 (PC); near Palmira, La Montanita, Maas & Escobar 605 (U). VENEZUELA. ARAGUA: Maracay, Parque Pittier, Onradt 78.V.4516, 4518, 4519, 4576 (PC); Parque Primera Toma, Cucube de Choroni, Onraedt 78.V.4578 (PC); Maracay, Rancho Grande near Portachuelo and Ocumare, Mägdefrau 445 (FH). CARABOBO: Campanero, Funck 785 (G). CARACAS: Type of M. flabellata; Colonia Továr, Onraedt 78.V.4778, 4781 (JE); Galipan, Funck & Schlim 337 (BM, G, PC). DISTRITO FEDERAL: El Junco, Agostini et al. 81 (JE, MEXU, U). MÉRIDA: S.loc., Luteyn et al. 9971 (NY); Mucuy, 1978, Onraedt 78.V.5606 (PC); Sierra Nevada, Onraedt 78.V.5884 (PC). TRUJILLO: Bocono, Luteyn & Cotton 9718 (NY). TRINIDAD. Thaxter s.n. (FH). SURINAME. Without locality, Wullschlägel s.n. (BM). ECUADOR. CARCHI: 0°47' N, 78°09' W, Steere 26598 (PC). CHIMBORAZO: Mt. Chimborazo, Spruce H 91e (MANCH). MORONA SANTIAGO: River Gualaquiza, Allioni in Levier 6408 (G).NAPOPASTAZA: Type of M. ecuadorensis; 0°30' S, 77°54' W, Brako 5409 (NY); Near Mera, Mexia 6962 (NY, UC); Puyo, convent of Dominicans, Mexia 6940 (FH, NY, UC). PASTAZA: Canelos, Spruce 94c (MANCH). PICHINCHA: Finlandia, 18 km E Sto Domingo de los Colorados, Arts 18/113 (FH, MEXU); Toachi river, Quito-Sto Domingo, Arts 17/012, 17/020 (PC). TUNGURAGUA: Baños, Spruce 1673 (MANCH). GALÁPAGOS IS.: Isabela, volcán Santo Tomás, Eliasson 1519 (U); Sta Cruz, Gradstein & Sipman H 366 (U). PERU. AYACUCHO: La Mar, near Ayna, P. & E. Hegewald 9063 (PC). CUZCO: Paucartambo, Tambomayo, Vargas 5112 (Herb. Fulford). HUANUCO: Leoncio Prado, P. & E. Hegewald 7889 (PC). JUNÍN: La Merced, Killip & Smith 24065 (JE). PASCO: Oxapampa, P. & E. Hegewald 8511 (PC). SAN MARTÍN: Tarapoto, Spruce 1676 (MANCH); Rioja, road Olmos-Moyobamba km 403, Geissler

Bischler et al. 191 8679 (G). BRAZIL. Type of M. brasiliensis ; Morro Velho, Gardner 191 (FH). ESPÍRITU SANTO: Sta Barbara do Caparo, Chase 1007 (JE). MINAS GERAIS: Gongo Soco, Hoehne in Verd. exs. 377 (BM, G, JE, MANCH). PARANÁ: Serra da Esperança, Vitt 21394 (PC), Vitt 21396 (MEXU); km 119 on road Br 277, Frahm 1619 (MEXU). RIO DE JANEIRO: Type of M. bescherellei; near Nova Friburgo, Glaziou 17992 (G, PC). SANTA CATARINA: Faragua, Carl 59 (JE). SÃO PAULO: Alto da Serra, Mosén 38 (G, PC, W). BOLIVIA. COCHABAMBA: Types of Plagiochasma bolivianum and Preissia barbata; Comarcacha, Cardenas & Cutler 1373 (Herb. Fulford). LA PAZ: Type of Grimaldia peruviana; N Yungas, 18 km S of Santa Ana de Alto Beni, Lewis 86-1989 (NY); S Yungas, 5-6 km W of Puenta Villa, Lewis 83-2867, 2864 (F); Mapiri, Bang 1545 (F, NY). CHILE. VALPARAISO: Type of M. notarisii. ARGENTINA. SALTA: Orán, Aguas Blancas, Capurro 327 (PC). PARAGUAY. Amabay, W of Pedro Juan, Caballero, arroyo Naranja-hai, Parque Nacional Cerro Corá, Geissler 14837 (G). URUGUAY. Montevideo, without collector (W). Marchantia chenopoda is a polymorphic species, able to adapt to a variety of habitats. Female plants are easily distinguished from the other neotropical species by the archegoniophore stalks with two narrow bands of air chambers. The hemispherical receptacles are strongly asymmetric, always shallowly 5-lobed. In M. breviloba, the number of lobes is greater than 5 and the receptacle nearly symmetric. Sterile samples can be distinguished by the shape of the appendages of the median scales, which are acuminate and acute in M. chenopoda but wide and usually rounded apically in M. breviloba. The thallus margins separate M. chenopoda from M. inflexa: usually purplish and with thickened marginal cells in M. chenopoda and hyaline and with thin-walled cells in M. inflexa. The cilia of the cupules are 3-5 cells long in M. chenopoda, 1-2 cells long in M. inflexa.

6. Marchantia inflexa Nees & Mont., in Montagne, Ann. Sci. Nat., Bot. sér. 2, 9: 43. Jan 1838. Type (Bischler, 1984). Martinique (lectotype, PC; isotypes, G, PC, W).

Figs. 153, 154, 160: K

Marchantia quinqueloba Nees, Naturgesch. Eur. Leberm. 4: 98. Sep 1838. Type (Bischler, 1984). Without locality (lectotype, STR; isotype, G). Marchantia disjuncta Sull., in Gray, Amer. J. Sci. Arts ser. 2, 1: 74. 1846. Type (Bischler, 1984). Borders of river Alabama, Claiborne, Sullivant 128, 286 (lectotype, FH; isotypes FH, G, NY, syntypes, FH, NY). Marchantia elliottii Steph., Spec. hepat. 1: 172. 1899. Type. Dominica. Elliott 1292 (holotype, G).

Bischler et al. 192 Marchantia caracensis Steph., Spec. hepat. 1: 188, 1899. Type (Bischler, 1984). Mexico. Wawra s.n. (lectotype, G). Thallus green to light green or glaucous, without or with weak purplish pigmentation, usually 2.84.2 mm wide, usually without distinct median band on dorsal side; margins hyaline, marginal cells thin-walled. Epidermal cells without papillae. Epidermal pores usually 60-92 µm diameter, inner opening bordered by cells with straight to slightly convex inner walls. Basal tissue with scattered oilcells and purplish sclerotic cells, with or without mucilage cavities. Ventral scales in 4 rows; appendage of median scales purplish, light red or orange, broadly triangular (length : width ratio usually 1-1.6 : 1), usually 10-14 cells wide, acute or shortly apiculate apically, margins nearly regularly toothed, with 1-2-celled teeth, marginal cells usually 1.5-3.7 times smaller than inner cells, oil-cells rare; laminal scales purplish or light red, rarely hyaline. Gemma cups with irregular cilia 13(-4) cells long, outer side without papillae. Stalks of gametangiophores without conspicuous scales basally. Antheridiophore stalk with single, narrow, often interrupted band of air chambers, and 2(-4) rhizoid furrows; receptacle palmate, usually 4.7-7.1 mm diameter, deeply dissected into 4-6(-7) rays, with few, scattered papillae on dorsal side, or papillae absent. Archegoniophore stalk with single, broad band of air chambers and 2(-4) rhizoid furrows; receptacle usually 3.3-5.3 mm diameter, symmetric or slightly asymmetric, with hardly distinct median projection dorsally, deeply dissected into 5-9(-11) lobes, not or slightly broadened apically; scales of receptacle without sinuose marginal cells apically. Involucres at margins crenulate or with irregular cilia 1-2 cells long. Spores yellowish or brown, usually 25-29 µm diam, with tuberculate areoles. Gametophytic chromosome number n = 9. Distribution and ecology. Marchantia inflexa is a New World species, known from 36°N to 7°30' N in the U. S. A. (Alabama, Arizona, Arkansas, Florida, Georgia, Louisiana, Oklahoma, Tennessee, Texas), and in the Neotropics from Mexico (Chiapas, Guerrero, Hidalgo, Nuevo León, Oaxaca (Düll, 1999), Puebla, Querétaro, Tamaulipas, Veracruz), Guatemala, El Salvador, Nicaragua, Costa Rica, Panama, Bahamas (Britton & Millspaugh, 1920), Cuba, Jamaica, Haiti, Dominican Republic, Puerto Rico, Leeward Islands (St. Kitts, Monserrat Is.), Windward Islands (Guadeloupe, Dominica, Martinique, St. Vincent, Grenada), Colombia (S. Andres Is., Santander), Venezuela (Caracas), Trinidad. It is a common species in the Caribbean area, growing on soil and soil over rocks (limestone, volcanic), in humid and usually shaded places in tropical rain forest under open vegetation, or at water edges, or in seepage areas where it may be intermittently flooded, more rarely on road banks, preferentially at low elevations (below 1000 m) but reaching 3100 m in Central America. Selected neotropical specimens examined. MEXICO. Type of M. caracensis. CHIAPAS: Sumadero San Cristóbal, Las Casas, Alexander 1186 (NY). GUERRERO: Petlacala, Mexia 9093a (BM, FH, UC). HIDALGO: Chapulhuacan, near Jacala, Frye & Frye 2848 (FH); Tula, Pringle 10675 (MEXU). NUEVO LEÓN: El Salto, 3 km S of Zaragoza, Whittemore 2457 (MEXU). PUEBLA: without locality, Nicolas 5101 (PC). QUERÉTARO: Sierra Gorda, Chuveje Cascades, W of Jalpan, Long &

Bischler et al. 193 Delgadillo 29718 (E); S bank of Rio Santa Maria near Ayutla, Long & Delgadillo 29725 (E). TAMAULIPAS: near Tamaulipas, Palmer 446 (FH, NY). VERACRUZ: Orizaba, Botteri s.n. (FH); near Mirador, Sartorius s.n. (FH); Córdoba, Underwood s.n. (STR); Los Chorros, Steyermark 32919 (Herb. Fulford); Orizaba, Schweinitz 114 (Herb. Fulford); Barranca Metlac, 1948, Ruiz Oronoz & Herrera s.n. (MEXU); Tlapacoyan, Camino a Rio Sardo, Ruiz Oronoz & Herrera s.n. (MEXU); Piedra de Agua, between Xalapa and Naolinca, Long et al. 29628 (E); Rio Jamapa between Huatusco and Coscomatepec, Long & Garcia 29673 (E). GUATEMALA. Alta Verapaz, Cobán, Türckheim in Levier 4960 (FH); Black River, E Verapaz, Watson 295b (FH, NY) EL SALVADOR. San Vicente, Standley 21753 (FH); Ahuachapan, Standely & Padilla 2539 (Herb. Fulford) NICARAGUA. Near Grenade, Levy 1310 (PC). COSTA RICA. S. José, Schultes 12062 (FH); Coradonga, 1901, Tonduz s.n. (G); Guanacaste, Finca Pacifica, 7 km NE of Cañas, Delgadillo 2778 (MEXU). PANAMA. Cocle, Welch 19949 (Herb. Fulford). CUBA. Loma del Gato, Leon et al. 10004 (FH, NY); Oriente, S. Maestra, Loma de la Mesa, Lippold 16694 (JE). JAMAICA. Hope Grounds, Harris s.n. (FH); St. Elisabeth, Maxon & Killip 1453b (FH); Portland, Nannipot Is., Morley & Whiteford 742 (FH); Portland, Maxon & Killip 9 (FH); Portland, Blue Mts., Stony River Base Camp, Morley & Whiteford 578 (FH); St. Andrews, Newcastle road, McMorris s.n. (NY), Farr 269 (Herb. Fulford). HAITI. Marmelade, Leonard 8158 (FH, JE); Massif de la Hotte, Zanoni et al. 24332B, 25670 (NY); St. Louis du Nord, Leonard & Leonard 14445 (FH, JE); SW St. Louis, Leonard & Leonard 14234 (FH). DOMINICAN REPUBLIC. Loma Diego de O'Campo, Zanoni et al. 25610L (NY); Pedernales, Buck s.n. (NY); River Mameyes, Eggers 1734 (G); Puerto Plata, Ekman 14509 (Herb. Fulford). PUERTO RICO. Between Arecibo and Utuado, Britton 2074 (NY); near Cayey, Sintenis H 42 (G); Rio Abajo, Pagán 2150 (MEXU). LEEWARD ISLANDS. ST. KITTS: Bethseda, Breutel s.n. (G, PC, STR, W). MONSERRAT IS.: Tar river, Shafer 871 (Herb. Fulford). WINDWARD ISLANDS. GUADELOUPE: La Madeleine, Baudoin s.n. (PC); Basse Terre, Duss s.n. (FH); Basse Terre, cascade des Ecrevisses, De Sloover 23552 (JE); Regretté, Questel 4414 (PC). DOMINICA: Type of M. elliottii; Laguna Roseau, Eggers H 3 (G); Roseau, Elliott 982 (BM, G). MARTINIQUE: Type; St. Pierre, 1820, Plée s.n. (FH, G); Morne de la Calabasse, Bélanger 487 (PC);

Bischler et al. 194 Morne Rouge, Le Gallo 882 (PC). ST. VINCENT: Kingston, Elliott 232 (BM); Morne Soufrière, Guilding s.n. (BM, FH). GRENADA: Thaxter s.n. (FH). COLOMBIA. S. ANDRES IS.: without collector (BM). SANTANDER: near Soratá, Killip & Smith 16515 (JE); near California, Killip & Smith 17072 (JE). VENEZUELA. CARACAS: Río Carude, 1856, Gollmer s.n. (G). TRINIDAD. Santa Cruz, Broadway 6770 (NY), Britton et al. 1505 (FH). Marchantia inflexa is most closely allied to M. papillata and has been considered a synonym of the latter by many authors (e.g., Hässel, 1963). However, the two species differ in the shape of the female receptacle, which is more deeply divided and with lobes convex and costate basally, strongly broadened apically and with a conspicuous median projection dorsally in M. papillata. The margins of the cupules and the involucres are shortly ciliate in M. inflexa, whereas they are crenulate in M. papillata. Other characteristics separating M. inflexa from M. papillata are the distinct median band on dorsal side of thallus, with purplish pigmentation of margins, the absence of mucilage cavities, and the spore wall ornamentation in M. papillata. Moreover, the scales of the female receptacle are sinuose apically in the latter species, whereas they are straight in M. inflexa. Their geographical ranges do not overlap. Sterile Marchantia chenopoda specimens can be distinguished from M. inflexa by their purplish thallus margins, bordered by thick-walled cells.

7. Marchantia paleacea Bertol., Opusc. Sci. (Bologna) 1: 242. 1817. Type (Grolle, 1976). Italy. Oriental Liguria, Borgonuovo, 1810, D. Turio s.n. (lectotype, BOLO). subsp. paleacea

Figs. 1: G, 155, 157, 160: G, L

Marchantia squamosa Raddi ex Lehm. & Lindenb., in Lehmann, Nov. Stirp. Pug. 4: 12. 1832. Type (Bischler, 1984). Brazil. Raddi s.n. (lectotype, W; isotypes, G, NY, S). Marchantia domingensis Lehm. & Lindenb., in Lehmann, Nov. Stirp. Pug. 6: 22. 1834. Type. Dominican Republic (holotype, W). Marchantia tholophora Bisch., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 17, 2: 989. 1835. Type (Bischler, 1984). Mexico. Oaxaca: Sommerschuh s.n. (lectotype, W; isotype, G). Thallus light or bright green, usually tinged with light red or purplish, usually 4.9-7.6 mm wide, without distinct median band on dorsal side; margins purplish or light red, rarely hyaline, cells thinwalled. Epidermal pores usually 67-92 µm diameter, inner opening cruciate. Basal tissue usually with purplish or yellow sclerotic cells, with few mucilage cavities. Ventral scales in 4 rows; appendage of median scales purplish or orange, ovate (length : width ratio usually 1-1.4 : 1), usually 16-22 cells

Bischler et al. 195 wide, rounded, acute or apiculate apically, margins entire, irregularly angular, crenulate, or irregularly toothed, sometimes with 1-2 conspicuous teeth towards base, marginal cells usually 2-3.8 times smaller than inner cells, oil-cells usually numerous; laminal scales purplish or light red. Gemma cups with ciliate lobes, outer side with 1-2(-3)-celled papillae. Stalks of gametangiophores with conspicuous scales basally, rounded apically, without appendage. Antheridiophore stalk without band of air chambers, with 2(-4) rhizoid furrows; receptacle peltate, usually 4.3-6.5 mm diameter, nearly symmetric, usually with 6-8 shallow, rounded lobes, dorsal side without papillae. Archegoniophore stalk with one band of air chambers and 2(-3) rhizoid furrows; receptacle usually 4.5-6.8 mm diameter, nearly symmetric to asymmetric, with rounded median projection on dorsal side, deeply divided into (5-)7-11 lobes, convex and costate basally, hardly broadened apically; scales of receptacle without sinuose cells apically. Involucres hyaline or purplish at margin, with ciliate lobes. Spores brown, 19-24(-28) µm diameter; spore-wall pitted. Gametophytic chromosome number n = 9 Distribution and ecology. The species has a circumtethyan range. In Europe, it is common in the Mediterranean area to 45° N. The species has also been found in northern Africa, SW Asia, the southern border of the Himalayas, China, Japan, and Malaysia. Outside this main range it has been collected in Yemen, Ethiopia, South Africa, and on Réunion. In the New World, it has been recorded from 36° N in U. S. A. (Alabama, Arizona, Florida, Oklahoma, Tennessee, Texas) to 5°N. Neotropical records include Mexico (Chiapas, Chihuahua, D. F., Hidalgo, Jalisco, Mexico, Michoacan, Morelos, Nuevo León, Oaxaca, Puebla, Querétaro, San Luis Potosí, Veracruz), Guatemala, Honduras, Costa Rica, Bermuda (Evans, 1906), Cuba, Jamaica, Haiti, the Dominican Republic, Colombia (Cundinamarca), Venezuela (Mérida). Records from farther south (Brazil, Bolivia and Chile) need confirmation. Marchantia paleacea is quite common in the Caribbean area, growing on clayey and sandy soil, on soil over rocks (limestone, sandstone, travertine), in rock crevices, on road banks, at water edges, in exposed sites or sheltered by open vegetation. It endures a short dry season and has been found most frequently in the altitudinal belt between 1000 and 2000 m, but was recorded from 500 to 3300 m. Selected neotropical specimens examined. MEXICO. CHIAPAS: 2 km N of San Cristóbal, Den Held & Van Rijn 41 (MEXU, U); 6 km E Bochil, Clayton B-420 (BM); Mun. Tenejapa, Paraeje Achilum, Breedlove 68398 (MEXU). CHIHUAHUA: Mun. Batopilas, W of La Buta, Buga 5928 (MEXU). D. F.: Beyond Contreras, Parker 704 (Herb. Fulford); Desierto de Los Leones, 1945, Ruiz Oronoz s.n. (MEXU). HIDALGO: Chapulhuacan, 5 km S Jacala, Frye & Frye 2848 (FH); Zacualtipan, Sanche Mejorado 287 (MEXU); San Miguel Regla, 1948, Herrera s.n. (MEXU). JALISCO: San Sebastian, Sierra Madre, Mexia 1518 a (G, UC). MEXICO: Paso Puerto de las Cruces, Düll 2/339 (FH, JE); 1 km W of Cahuacan, Cruz Cisneros 190 (MEXU); Mun. Oculian, Barranca de Mexicapa on road from Sta Monica de Ocuilan to Cuernavaca, Pócs 9547F (MEXU). MICHOACAN: Morelia, along Hwy to Hidalgo, Frye & Frye 3029 (G); Urupan, Frye & Frye 3031 (G); Barranca San

Bischler et al. 196 Lorenzo, Rees & Baltazar 676 (MEXU). MORELOS: Cuernavaca, Frye & Frye 3129 (G). NUEVO LEÓN: Ejido Purisima, 21.4 mi. S of hwy. 60 at Iturbide, Whittemore 2432 (MEXU); El Salto, 3 km S of Zaragoza, Whittemore 2455 (MEXU). OAXACA: Type of M. tholophora; Huautla de Jimenez, 1956, Heim s.n. (PC); 2 km N San Pedro y San Pablo Teposcoluta, Gracia Mendoza 483 (MEXU). PUEBLA: Sta Barbara, Nicolas 5101 (PC, STR); Río Alsecca, Arsène s.n. (PC); Mun. Xochitlan, Arroyo Atena, 15 km SE Zapotitlan, Grimes et al. 2764 (MEXU); Mun. Jopala, 24 km SE de San Antonio de Tilapa, Castrechon & Corichi 432 (MEXU); Matamoros, Ruiz Oronoz s.n. (MEXU). QUERÉTARO: Barranca de Amecalco, Argüelles 1361 (MEXU). SAN LUÍS POTOSÍ: near San Luís Potosí, 7670 (PC). VERACRUZ: Orizaba, Quelle s.n. (JE); Jalapa, Pringle 15326 (FH); Fortin de las Flores, Düll 2/114 (PC); Tlapacoyan, Planta de Luz, 1949, Ruiz Oronoz & Herrera s.n. (MEXU); Encino Gacha near Las Vigas de Ramirez, NW of Xalapa, Long et al.29621 (E); Rio Jamapa between Huatusco and Coscomatepec, Long & Garcia 29683 (E). GUATEMALA. 55 km N Antigua, Volk 6/0714 (JE); Alta Verapaz, Cobán, Türckheim in Levier 4960 (FH); Guatemala-San Raimundo, Standley 63025 (FH); Nahuala, Champagne 2/03, 07 (PC); NW Antigua, Standley 64644, 64635 (FH). HONDURAS. Comayagua, El Banco, Rodriguez 2574 (BM, UC). COSTA RICA. S. José, Schulthes 12062 (FH), Standley 41248 (FH); Zarcero-Laguna de Alfaro Ruíz, Rodriguez 289 (UC); Cord. de Talamanca, 1979, Jovet-Ast s.n. (PC). CUBA. Without locality, Wright s.n. (FH, G, NY); Oriente, Santiago de Cuba, Gran Piedra, Bisse & Lippold 15136/b (JE); Guantanamo, Hioram & Maurel 4660 (NY); Santiago de Cuba, Mt. Liban, Linden 2196 (G). JAMAICA. Silver Gap to Hardwar Gap, Maxon & Killip 1223 (FH); St. Andrews, Farm Hill, Blue Mts., Barry 63 A (Herb. Fulford); Morce's Gap, Blue Mts., Papenfuss s.n. (UC). HAITI. Kenskoff, Sastre 7257 (PC); La Selle, Croix-des-Bouquets, Ekman 7628 (Herb. Fulford); Dep. S, Massif de la Hotte, just outside Park Nat. Pic Macaya, N of Villa Formon, Judd et al. 5762 (U). DOMINICAN REPUBLIC. Type of M. domingensis; S Boaruco, Zanoni et al. 20467H (NY); S de Neibes, Zanoni et al. 24889 g (NY). COLOMBIA. CUNDINAMARCA: Usaquén near Bogotá, 1907, Apollinaire s.n. (G); San Francisco near Bogotá, 1907, Apollinaire s.n. (G). VENEZUELA. MÉRIDA: Vicinity of Timotes, Pittier 12673 (G). Marchantia paleacea is easily distinguished from the other species of subgen. Chlamidium by the cruciate epidermal pores and the cupules with ciliate lobes. The ventral side of the thallus with four rows of scales restricted to the median portion separates it from the species of subgen. Marchantia.

Bischler et al. 197 Two subspecies have been described: the widespread subsp. paleacea and subsp. diptera (Nees & Mont.) Inoue from Asia. All neotropical specimens belong to subsp. paleacea.

8. Marchantia papillata Raddi, Critt. Brasil. 20. 1822 (Mem. Mat. Fis. Soc. Ital. Sci. Modena, Pt. Mem. Fis. 19: 44. 1823). Type (Bischler, 1984). Brazil (lectotype, PI; isotypes, G, NY, STR, W). subsp. papillata

Figs. 156, 157, 160: H

Marchantia pusilla Nees, in Meyen, Nov. Actorum Acad. Caes. Leop.-Carol. Nat. Cur. 19, Suppl. 1: 476. 1843. Type. Chile. Without locality, Meyen s.n. (holotype, STR). Marchantia subandina Spruce, Trans. & Proc. Bot. Soc. Edinburgh 15: 561. 1885. Type (Bischler, 1984). Peru. San Martín: Near the falls of river Huallaga, Spruce s.n. (lectotype, MANCH; isotype, G). Marchantia faxinensis Schiffn., in Schiffner & Arnell, Oesterr. Akad. Wiss., Math. Naturwiss. Kl. Denkschr. 111: 9. 1964. Type. Brazil. São Paulo: Near Faxina, 650 m, Schiffner 2349 (holotype, W). Thallus yellowish or dark green, usually 2.4-3.4 mm wide, usually with narrow, black median band on dorsal side. Margins usually purplish, cell walls thin. Epidermal pores usually 50-84 µm diameter, inner opening bounded by cells with straight or slightly convex inner walls. Basal tissue with numerous, purplish sclerotic cells, without mucilage cavities. Ventral scales in 4 rows; appendage of median scales purplish, orange or hyaline, triangular (ratio length : width usually 1-1.7: 1), (6-)8-14 cells wide, acute or apiculate, with row of (1-)2-3 cells apically, margins toothed, teeth mostly unicellular, usually 1.8-3.4 times smaller than inner cells, without oil-cells; laminal scales purplish or hyaline. Gemma cups nearly entire, crenulate or with some irregular cilia, 1-2 cells long, without papillae on outer side. Stalks of gametangiophores not sheathed basally by large scales. Antheridiophore stalk with 1 narrow, often interrupted band of air chambers or without band, and 2 rhizoid furrows; receptacle palmate, usually 3.5-5.5 mm diameter, deeply dissected into 4-7 rays, without papillae on dorsal side. Archegoniophore stalk with single, broad band of air chambers, sometimes splitting into two bands, and 2 rhizoid furrows; receptacle nearly symmetric, usually 3.24.4 mm diameter, with rounded median projection dorsally, deeply dissected into 5-9 lobes, lobes strongly convex basally, broadened apically; scales of receptacle with sinuose cells apically, ending in row of 1-2 cells. Involucres hyaline at margin, entire or irregularly crenulate. Spores yellow or brown, 24-26 µm diameter, with smooth ridges. Gametophytic chromosome number n = 9. Distribution and ecology. Marchantia papillata subsp. papillata is known in the Neotropics from 2°S to 35°S, from Ecuador (without locality), Peru (Huanuco, Lima (Carrillo & Chanco, 1971), San Martin), Brazil (Amazonas, Mato Grosso, Minas Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul (Vianna, 1976), Santa Catharina, São Paulo), Bolivia (Santa Cruz, Tarija), Chile (without locality),

Bischler et al. 198 Paraguay, Argentina (Buenos Aires, Córdoba, Jujuy, Misiones, Tucumán (Hässel, 1963)), to Uruguay (Hässel, 1964). The species is common only in south-eastern Brazil, rare elsewhere, and grows on soil and soil over rocks, often near water edges, mainly at low elevations (0-500 m), but has been collected up to 1200 m in Argentina. Selected specimens examined. ECUADOR. Without locality, Eggers 15009 (G). PERU. HUANUCO: Leoncio Prado, Tingo Maria, P. & E. Hegewald 7833 (Herb. Hegewald). SAN MARTÍN: Type of M. subandina. BRAZIL. Type. AMAZONAS: Jurua Miry, Ule 527 (BM, G, JE); Manaos, Rio Negro, Spruce s.n. (MANCH). MATO GROSSO: Caceres, Facenda Jacobina, Höhne 243 (JE). MINAS GERAIS: Widgren s.n. (G); Ituiutaba, Macedo 3409 (W); Dante, Cachoeira do Oculo, Mexia 4999a (UC). PARANÁ: Rio Negro, d'Orbigny 104 (G). RIO DE JANEIRO: Corcovado, Bandeira 28 (JE), Glaziou 1995 (PC); Mosén 285 (FH, G, W). SANTA CATARINA: Rio Itajahi, Pabst 1646 (229) (FH). SÃO PAULO: Type of M. faxinensis; near Iguape, Loefgren 33094 (JE); Raiz de Serra, Schiffner 152 (W); Brotas, Vital s.n. (PC) BOLIVIA. SANTA CRUZ: Florida, 5 km E of Samaipata, Lewis 83-5321 (F). TARIJA: Arce, 1 km W of Agricultural Experimental Station, Communidad Sidras, Lewis 84-2334 (F). CHILE. Type of M. pusilla. PARAGUAY. Paraguari, Balansa 4006 (G, PC); Dep. Paraguari, SE Piribebuy, Salto Pirareta, Geissler 15164 (G); Dep. Central, S Colonia Independencia, Cordillera del Ybytyruzu, near Cerro Guayaqui, arroyo Tacuara, Geissler 15057 (G). ARGENTINA. BUENOS AIRES: Punta Lara, Hässel 9350 (JE, U). CÓRDOBA: La Paz, Loma Bola, Hässel 22964 (G, JE, U). JUJUY: S.loc., Fries 35 (FH). MISIONES: Teyucuaré, Perez Moreau 1980 (JE, PC). Marchantia papillata comprises two subspecies, the neotropical subsp. papillata and the Asiatic subsp. grossibarba (Steph.) Bischl., distinguished mainly by the shape of the appendage of the median scales. Additionally, subsp. grossibarba is more robust, nearly all parts of the plants being larger, and the lobes and rays of the female and male receptacles are more numerous. The shape of the female receptacle, with lobes costate basally and broadened apically, separate M. papillata subsp. papillata from the other species of subgen. Chlamidium. The scales of the female receptacle have sinuose cells apically. Spore-wall ornamentation in M. papillata is also distinctive. Marchantia papillata subsp. papillata is most closely related to M. inflexa and sometimes the two have been considered conspecific (see under the latter species).

Bischler et al. 199

MONOCLEALES

by S. R. Gradstein

2. MONOCLEALES R.M.Schust., J. Hattori Bot. Lab. 26: 296. 1963. Type. Monocleaceae (Nees) A.B.Frank The order comprises a single family.

11. MONOCLEACEAE (Nees) A.B.Frank, in Leunis, Syn. Pflanzenk. (ed. 2) 2, 3: 1556, 1576. 1877. Tribe Monocleae Nees, in Gottsche et al., Syn. hepat.: 507. 1846. Type. Monoclea Hook. The family is monogeneric.

19. MONOCLEA Hook., Musci Exot. 2: 27, t. 174. 1820. Type. Monoclea forsteri Hook. Name referring to the capsule opening with a single slit. Thallus pure green, growing prostrate, apical cell cuneate, with four cutting faces; thallus variable in size, small to vary large, 0.5-30 mm wide, without purplish pigmentation, the upper surface densely spotted by whitish or darkish dots in fresh material, margins usually crisped-undulate, midrib usually lacking. Branching dichotomous. Epidermal pores and differentiated assimilatory layer lacking. Basal tissue with oil cells containing some chloroplasts, without mucilage cavities. Ventral scales lacking. Mucilage papillae 1- to several-celled, on ventral side at thallus apex. Rhizoids smooth, sometimes with isolated papillae but not distinctly tuberculate, often branched. Asexual reproduction by specialized propagules lacking. Dioecious. Antheridia cylindrical-ellipsoidal, jacket unistratose except distally where 2 layers may be present, stalk short, embedded in dorsal receptacles, sometimes bounded by cilia; ostioles formed by protruding thallus tissue. Archegonia with 14-20 neck canal cells, in cross section with 6 neck cell tiers, terminal on main thallus, several per involucre, accompagnied by mucilage hairs; involucre single, tubular, laterally fused to thallus margins; scales associated with gynoecia lacking. Calyptrae dividing prior to fertilization (2-3 layers), becoming 6-7layered after fertilization. Pseudoperianths lacking. Sporophytes 1-3 per involucre, with conspicuous, obtuse-conical foot and long, thick seta arising near thallus apex, from beneath an elongate swelling of thallus. Capsules ellipsoidal, opening by a longitudinal slit; wall unistratose, with internal and external anastomosed thickenings; spore/elater ratio more than 4 : 1. Spores hardly polar, 10.000 or more per capsule, 12-23 µm diameter, proximal and distal faces with reticulate ornamentation,

Bischler et al. 200 triradiate scar indistinct. Elaters long, often branched, with 2-3 helical bands. Gametophytic chromosome number n = 9. A genus of two species growing on wet forest floor or on shady, damp banks and rocky riversides. One species is endemic in New Zealand (M. forsteri Hook.), the other (M. gottschei Lindb.) is common in the Neotropics, from Central America to tropical and temperate South America. Sporophytes are not uncommon in locations where male and female populations occur side by side. Sporophytes have been little collected, however, and are scarce in herbarium materials. The presence of sporophytes is indicated in the specimens examined section by “c. spor.”. Collections with numerous sporophytes are the ones from Jamaica gathered by Howe.

1. Monoclea gottschei Lindb., Rev. Bryol. 13: 102. 1886. Type (Gradstein et al., 1992). Chile. Los Lagos: Valdivia, Corral, 5 June 1896, Dusen 125, with male receptacles and sporophytes (neotype, G; isoneotypes, NY, S). Antheridial receptacles variable, orbicular to longer than wide, margins irregularly and obtusely crenate-crispate, rarely entire, bristle-like hairs lacking. Sporophyte seta 0.6-0.9 mm diameter, made up of 20-25 rows of cells; capsule 5-7 times longer than wide; capsule wall thickening bands approx. 2.5 µm wide (longitudinal section), subvertical in position and not bulging outwards, the radial walls in cross section smooth. Gametophytic chromosome number n =9. Distribution and ecology. Widely distributed in the mountains of the Neotropics (subsp. elongata) and at low elevations in warm-temperate regions of Chile and Argentina (subsp. gottschei). Monoclea gottschei grows in moist, shaded places in disturbed and undisturbed montane rain forests, on moist earth, leaf-litter, rotten wood, trunk bases and thin, wet soil over rock, usually near running water, from sea level in warm-temperate regions to nearly 4000 m in the tropical Andes. Monoclea gottschei often grows associated with Dumortiera hirsuta. Fertile material of the latter species is easily recognized by the short-stalked, rounded male and female receptacles with hairy margins but sterile thalli of the two species may be confusingly alike. The thallus of Dumortiera is usually more glossy and evenly green (becoming olive-green when dry), the margins are plane or somewhat irregularly sinuate, never crispate, and the thallus surface does not have any dots. However, the characteristic dots of Monoclea are usually not well visible in old herbarium material. A very good difference is the presence of a whitish to yellowish "midrib" on the underside of the thallus of Dumortiera, made up of very tightly associated, tuberculate rhizoids forming a narrow strand along the midline of the thallus. The rhizoidal strand is often surrounded by more loosely associated rhizoids. Long, pale rhizoids may curve away from the midline to the thallus margins in a parallel fashion. In thin thalli, the rhizoidal strand may be clearly visible in dorsal view, resembling a midrib. Such a false midrib is lacking in Monoclea; its presence or absence is a reliable character to separate

Bischler et al. 201 sterile, dried thalli of the two genera. Moreover, rhizoids in Monoclea are smooth or only slightly tuberculate, never densely tuberculate as in Dumortiera.

KEY TO THE SUBSPECIES OF MONOCLEA GOTTSCHEI 1. Antheridial receptacles (3-)4-5 mm wide, always orbicular............................. 1a. subsp. gottschei 1. Antheridial receptacles narrower, up to 2.5 mm wide, variable in shape, orbicular to elongate ...... ...............................................................................................................................1b. subsp. elongata

1a. Monoclea gottschei subsp. gottschei Type. See under the species.

Antheridial receptacles relatively large, (3-)4-5 mm wide, always orbicular. Distribution and ecology. Monoclea gottschei subsp. gottschei occurs in the warm-temperate regions of Chile (Aisén, Araucania, Los Lagos) and Argentina (Neuquén), between about 38 and 44° S, at elevations ranging from sea level to about 900 m on Juan Fernandez Is. Habitat: see under the species.

Representative specimens. ARGENTINA. NEUQUÉN: San Martin de los Andes, Kühnemann s.n. (BA). CHILE. Guaitecas I., Dusén s.n. (NY); Queule, Dusén s.n. (NY). AISÉN: Punto Puyuhuapi, Schwabe 14e (c. spor.) (JE); Termas de Puyehue, Schwabe 10/c (JE). ARAUCANIA: Cautin, near Temuco, Becker s.n. (U). JUAN FERNANDEZ: Masatierra, Skottsberg 93, 114, 1152 (S), 94, 96, 97, 99 (NY), Sparre 111277 (S), Hatcher & Engel 642 (NY). LOS LAGOS: Puerto Mont, Dusén 206 (NY); Corral, Dusén 125 (G, S, NY), Thaxter 70 (NY, WELC), Hosseus 35 (NY), Hosseus 36 (JE), Van Zanten 86-01-753 (U).

1b. Monoclea gottschei subsp. elongata Gradst. & Mues, Pl. Syst. Evol. 180: 133. 1992. Type. Bolivia. Ballívian: Serranía del Pilón Lajas, 650 m, 30 October - 3 November 1989, Gradstein 7191, male plants (holotype, U), Gradstein 7184, female plants, c. spor. (paratype, U). Figs. 158: A-F, 159, 160: I ? Monoclea elliottii Steph. ex Duss, Enum. Méth. Musc. Antilles Franç., Hép. 37. 1902, nom. inval.

Bischler et al. 202 Type. Martinique, St. Denis, Morne Rouge, Duss 578, 580 (n.v.). Antheridial receptacles 0.7-2(-2.5) mm wide, 1-5 times longer than wide, variable in shape, orbicular to oblong to linear, sometimes furcate and horse-shoe shaped. Receptacle length varying considerably within individual populations. Distribution and ecology. Monoclea gottschei subsp. elongata is widely distributed in the mountains of tropical America between 22° N and 30° S., ranging from Mexico and Cuba southwards to northern Argentina (Tucuman) and south-eastern Brazil at elevations from (150-)500 to the upper forest line at 4000 m, most commonly between 1000 and 3500 m. Habitat: see under the species. Representative specimens. MEXICO. CHIAPAS: Linden s.n. (PC); Jitotol, Breedlove 69045 (MEXU, NY); Villa Corzo, Breedlove & Thorne 30060 (NY). MEXICO: S of Convento, Delgadillo 4616 (MEXU); Lagunas de Zempoala, W of Tres Cumbres, Frahm 79-2338 (MEXU, U). MORELOS: Curnevaca, Barnes & Land 479 (NY); OAXACA: Sierra Juarez, above Tuxtepec, Sharp 3814 (MEXU). PUEBLA: along Cualtepec road, Barnes & Land 583 (c. spor.) (NY); Mun. Xochitlan S of Zapotitlan, Grimes et al. 2763 (MEXU, NY). SAN LUÍS POTOSÍ: above Xilitla, Sharp 5858 (MEXU). VERACRUZ: Zacuapan, Purpus 5533 (NY); Jalapa, Bischler et al. s.n. (MEXU). GUATEMALA. Mountains east of Tactic on road to Tamahu, Standley 71081 (NY). SACATEPÉQUEZ: Antigua, Standley 58984 (NY). SAN MARCOS: Standley 68558 (NY). BELIZE. CAYO: S. Maya Mts., Allen 15141 (MO). HONDURAS. Trail to Río Platano, Wilson 69 (NY). Various localities, Allen 14119, 14170, 14460, 17452, 17762, 17776 (MO). SAN SALVADOR. Monte Cristo, Thompson s.n. (c. spor.) (NY). NICARAGUA. ESTELI: highway to San Nicholas, 10 km W of La Trinidad, Stevens 14837 (MO). JINOTEGA: Macizos de Peñas Blancas, Stevens 11386 (MO). MATAGALPA: Fuente Pura, Moreno 18957 (c. spor.) (U). COSTA RICA. ALAJUELA: Peñas Blancas, Lyon 46, 51 (MO). CARTAGO: Tapantí, Gomez 18822, 18849, 18856 (MO); Turrialba, Maas 803 (c. spor.) (U); San Gerardo de Dota, Gradstein s.n. (GOET). HEREDIA: road between San Rafael and Rio Las Vueltas, Stevens 13920 (U). PUNTARENAS: Rincón, Gradstein & Dauphin 9342 (GOET); Monteverde Cloud Forest, abundant, Gradstein 9427, 9428 (c. spor.) (GOET), Lyon 22, 96, 105 (MO). SAN JOSÉ: La Hondura, King C91106 (MO); San Pedro, Centro Universitaria, along river, Gradstein s.n. (GOET). PANAMA. BOCAS DEL TORO: Hammel 6113 (MO). CHIRIQUÍ: Palo Alto, Hammel 7436C (MO, U); La Fortuna, Hammel 2105 (MO), 2286 (U), Salazar et al. 6245, 6250 (PMA, U), Spörle 1.20 (U), Valdespino et al. 21, 30, 105, 106 (PMA, U); Cerro Colorado, Antonio 2605 (NY), 4865 (MO). DARIÉN: between Cerro Pirre and Piji Vasal, Folsom 6418 (U).

Bischler et al. 203 CUBA. Without loc., Wright s.n. (NY); 7 localities (Reyes-Montoya, 1981). JAMAICA. Trail from Morce´s Gap to Vinegar Hill, Maxon & Killip 1301 (NY, PC); Hardwar Gap, Welch 17570, 17577B (NY); New Haven Gap, Underwood 890, 891 (c. spor.), 894, 1020 (c. spor.), 1081 (c. spor.) (NY), Evans 157 (NY); Clyde valley below Cinchona, Underwood 331 (c. spor.), 368 (c. spor.), 388, 408 (c. spor.) (NY, U), Evans 25 (NY); Tweedside, Underwood 1615 (c. spor.) (NY, U). HAITI. Mt. Casse, Nash 546 (NY). PUERTO RICO. Without loc., Sintenis H81 (NY); Luquillo Mts., Gradstein 6559, 6560, 6561 (U); El Yunque, Evans 33, 98 (NY), Pagán 1001, 1972 (NY); trail to Mt. Britton, Steere 4220, 4240 (NY), Mt. El Toro, Gradstein 6550 (U); Cordillera Central, above Villalba, Gradstein 6514, 6515, 6516 (U), Buck 15982 (NY). WINDWARD ISLANDS. GUADELOUPE: without loc., L’Herminier s.n. (NY, PC); Col de la grande Découverte et Ravine à Déjeuner, Duss 422 (c. spor.) (NY); Capesterre, Gradstein & Sastre 6609, 6610, 6611, 6612, 6613, 6614 (U); deuxième chute du Carbet, De Sloover 3876 (NY, U); cascade des Ecrevisses, De Sloover 23.463 (JE). MARTINIQUE: Camp de l’Alma, Duss 16 (NY). DOMINICA, Elliott s.n. (Spruce, 1895). ST. VINCENT: Elliott s.n. (Spruce, 1895). COLOMBIA. "Andes Bogotensis, " Weir s.n. (c. spor.) (NY). ANTIOQUIA: SE of Medellin, Sastre de Jesus & Churchill 916 (U). CASANARE: Aguirre et al. 2876 (COL, U). CUNDINAMARCA: San Antonio, Lindig 1708 (PC), Lindig 1733 (c. spor.) (BM, G, H, NY, S); Fusagasuga, Lindig s.n. (c. spor.) (W); Chicaque, Linares & Gradstein 3525 (c. spor.), 3529 (COL, U); San Bernardo, Van Reenen et al. 3040 (COL, U); Subachoque, Bulla & Linares s.n. (COL); El Tablazo, van Zanten & Gradstein 309 (c. spor.) (COL, U). HUILA: La Plata, Finca Merenberg, Aguirre et al. 63331, 6341 (c. spor.) (COL, U). MAGDALENA: San Lorenzo, Winkler C343 (Winkler, 1976). META: Van Reenen 2809 (COL, U). PUTUMAYO: King C1069a (Robinson, 1967). RISARALDA: Santa Rosa de Cabal, Van Reenen 3 colls. (COL, U), Gradstein 3569 (COL, U), Wolf 497 (COL, U); Cordillera Central, W of Mistrato, Gradstein 8376, 8394, 8487 (COL, GOET). SANTANDER: Suratá, Killip & Smith 16576 (JE); Charta, id. 19217 (JE); California, id. 16977, 17036 (JE). Charalà, Virolin, Castro et al. s.n. (COL), Gradstein 7020 (c. spor.) (COL, U). TOLIMA: Santa Isabel, Aguirre et al. 1514 (COL, U); Mt. Tolima, van der Hammen & Jaramillo M. 3395 (COL, U). VALLE: Farralones de Cali, near Pance, Hartman 232b (U). VENEZUELA. Without loc., Funk & Schlim s.n. (c. spor.) (NY). AMAZONAS: Cerro de la Neblina, Sastre-de Jesús 486 (JE). DISTRITO FEDERAL: Colonia Tovar, Steyermark et al. 127798 (U, VEN). LARA: Jiménez, Parque Nacional Yacambú, Davidse & Gonzáles 21283 (U). TACHIRA: Junín, Cerro San Isidro, Davidse & Gonzáles 22017 (c. spor.) (JE, MO, U); Uribante, Represa Las Cuevas, van derWerff & Gonzáles 5138 (U). TRUJILLO: Bocono, Guaramecal, Liesner et al. 13040

Bischler et al. 204 (MO, U). ZULIA: Sierra Pajarita, S. José de los Altos, Griffin s.n. (Griffin, 1977). GUYANA. Mt. Roraima, Gradstein 5255, 5312 (c. spor.), 5740 (U). SURINAME. Emmaketen, Daniels & Jonker 826 (U). FRENCH GUIANA. Mt. Galbao, Gradstein 6199 (U), Bekker 2305, 2307 (U). ECUADOR. CARCHI: Paramo el Angel, Gradstein & Mues 6846 (U). LOS RIOS: Rio Palenque, Gradstein & Mues 6945, 6955 (U). NAPO: Laguna Papallacta, Gradstein 6927 (U). PICHINCHA: W of Lloa, Gradstein 6704, 6705, 6781, 6798 (U); W of San Juan, Gradstein 6742 (U). PERU. AMAZONAS: prov. Chachapoyas, road Cajamarca-Chachapoyas, Frahm et al. 1118 (PC). LORETO: Pucallpa, Spichiger s.n. (G). HUANUCO: Tingo Maria (Proskauer, 1951). SAN MARTÍN: Mt. Guayrapurina, Spruce s.n. (W); road Yurimaguas-Tarapoto, Frahm et al. 1391, 1690, 1716, 2089 (PC); above Rioja, Frahm et al. 83, 642, 789, 2191 (PC), 429 (U). URUBAMBA: Macchu Picchu, Peyton 548 (F, U). BRAZIL. MINAS GERAIS: Caldas, 5 Sep 1873, Mosén s.n (W). RIO DE JANEIRO: Jardim Botánico, Pinheiro da Costa s.n. (RB); near Novo Friburgo, Gradstein 9809 (c. spor.) (GOET). RIO GRANDE DO SUL: Guaiba, Maria Pimentel, Vianna s.n. (ICN 37251 / Hep. 4251) (ICN, PC, U). SÃO PAULO: Rio Branco near Santos, Schiffner 293 (W); Serra São Joao near Santos, growing mixed with Dumortiera hirsuta, Schiffner 1996 (W); Alto da Serra, Schiffner 162, 163 (W), Apiai, Morro d'Oro, Schiffner 316 (W). BOLIVIA. BENI: Ballívian, Serranía del Pilón Lajas, Gradstein 7118, 7154, 7183, 7184 (paratype, c. spor.), 7191 (holotype), 7224 (U). COCHABAMBA: Carrasco, Lewis 83-1945M (BA); Chapare, E of Incachaca, Lewis 83-1238A, 85-1746 (BA); Río Tocorani, Herzog 4099 (W); Tablas Monte, Gradstein 7476, 7477 (U). LA PAZ: Inquisivi, W of Cajuata, Lewis 83-3826 (BA); Coroico, Lewis 86-1866 (BA), road La Paz-Coroico 3000 m, Gradstein 7069 (U). SANTA CRUZ: near Comarapa, Lewis 85-450 (BA). TARIJA: N of Entre Rios, Gradstein 7670, 7674 (U); road TarijaBermejo, N of El Marmore, Gradstein 7718 (U). PARAGUAY. Parque Nacional Ybicu’i, Zardini 12587 (GOET, MEXU, MO). ARGENTINA. TUCUMÁN: between Tafi and Tucumán, Steere 60-229, 60-310 (ster.) (NY); Sierra San Javier, Quebrada de Cainzo, Vervoorst s.n. (ster.) (Hässel de Menéndez, 1957); Chicligasta, Río Cochuna, Digilio-Grassi 229 (ster.) (Hässel de Menéndez, 1957). Monoclea gottschei subsp. elongata is rather similar to subsp. gottschei but differs in the different size and shape of the androecial receptacle and in its different geographical distribution. The antheridial receptacle in subsp. elongata is usually elongate (slightly to strongly so), rarely orbicular. When orbicular, the receptacle in subsp. elongata may distinguished (when mature) from that of

Bischler et al. 205 subsp. gottschei by its lesser diameter, measuring maximally up to 2.5 mm (3-5 mm in subsp. gottschei). The two subspecies have clearly non-overlapping, allopatric distributions: subsp. elongata has a wide neotropical range whereas subsp. gottschei occurs exclusively in warm-temperate regions of southern South America (Gradstein et al. 1992). The status of the two subspecies has been confirmed based on chloroplast DNA analysis (Meissner et al., 1998).

SPHAEROCARPALES by S. R. Gradstein

3. SPHAEROCARPALES Cav., New Phytol. 9: 84. 1910. Type. Sphaerocarpaceae (Dumort.) Heeg

Plants thalloid or more or less leafy, prostrate or erect. Thalli simple, without air chambers and pores, differentiated into a thickened median portion (axis or "midrib") and 1-layered wings. Oil-cells lacking. Rhizoids all with smooth walls. Antheridia and archegonia on the thickened axial surface of the thallus or (Riella) antheridia on the wing, always in involucres, each involucre containing one gametangium. Sporophytes protected by the female involucre, seta reduced. Capsules spherical, cleistocarpous, remaining completely enclosed in the involucres, opening through disintegration of the capsule wall and the involucre, the wall 1-layered. Spores large, 50-140 µm, often remaining in tetrads. Elaters lacking. Asexual reproduction by specialized devices absent or by gemmae. A small order of about 25 species, in 3 genera and 2 families. They are mostly plants of subtropical and Mediterranean regions with dry, hot summers and moist, rather cool winters. Sphaerocarpales are not yet known with certainty from the Neotropics but might occur in the region. The principal characteristics of the order are 1) delicate, simple, terrestrial thalli, without internal differentiation; 2) antheridia and archegonia surrounded singly by a conspicuous, ± pear-shaped involucre; 3) sporophyte with a reduced seta and a cleistocarpous capsule that is completely enclosed in an inflated involucre; and 4) lack of elaters.

KEY TO THE FAMILIES OF SPHAEROCARPALES

Bischler et al. 206 1. Thallus aquatic, erect, consisting of a narrow, thickened axis with a broad, undulate or spirally coiled wing on one side and with several rows of small leaf scales. Involucres in a row on the axis (female ones) or wing margins (male ones) .................................................................... 1. Riellaceae 1. Thallus terrestrial, prostrate, thickened in the middle, unistratose towards the margins, the margins sinuose-lobed. Involucres in dense groups on the median thallus surface, at maturity almost completely hiding the thallus .............................................................................. 2. Sphaerocarpaceae

12. RIELLACEAE Engler, Syll. Pflanzenfam., ed. 1: 45. 1892. Type. Riella Mont. A monogeneric family.

20. Riella Mont., Ann. Sci. Nat., Bot. sér. 3, 18 : 11. 1852. Type. Riella notarisii (Mont.) Mont. Genus dedicated to M. C. Durieu de Maisonneuve (1796-1878), Director of the Botanical Garden of Bordeaux. Thalli aquatic, erect growing (prostrate when growing emerged on wet substrate), green, 1-10 cm long, simple or furcate, laterally compressed, consisting of a narrow, thickened axis and a broad, unistratose, undulate or spirally coiled wing. Lateral and ventral scales present, originating from the axis, in two rows each. Dioecious, rarely monoicous. Female involucres large, originating from the axis, male involucres very small, usually in notches in the thallus wing, the involucres arranged in a single row. Spores large, not remaining in tetrads, 50-130 µm, surface covered with long, narrow spines. Asexual reproduction by caducous ventral scales. Riella is a subtropical-mediterranean genus of probably about 10-15 species (about 20 species have been described from Europe, Africa and America but several of these may prove to be synonyms; a monograph of Riella has never been undertaken). Five species have been reported from Argentina (Hässel de Menéndez, 1987), from the provinces Buenos Aires, La Pampa, Río Negro, and Neuquén, between about 35 - 40º S, and one species, R. americana Howe & Underw., is known from Chihuahua and Durango, northern Mexico, at about 30º N (Proctor, 1972; Schuster, 1992b). The genus has not yet been recorded from within the boundaries of the Neotropics. Riella is an aquatic genus and grows erect and submerged in fresh or usually somewhat brackish, alkaline standing waters, at the margins of ponds and lakes at low elevations. Exceptionally, it occurs in flowing water of shallow, intermittently dry streams (Proctor, 1972). The plants are easily recognized by their very unusual habit, viz., the erect growing thallus consisting of a narrow axis bearing a broad, undulate or spirally coiled wing and two rows of leaf scales. The laterally compressed thallus, the presence of a wing on one side of the thallus only, and the occurrence in intermittently dry bodies of water in arid areas are unique features of Riella, separating the genus from all other thalloid liverworts (A. Whittemore, pers.

Bischler et al. 207 comm.).

13. SPHAEROCARPACEAE (Dumort.) Heeg, Verh. Zool.-Bot. Gesellsch. Wien 41: 573. 1891. Type. Sphaerocarpos Boehm. This family consists of two genera, one of wide distribution (Sphaerocarpos), the other endemic to southern California (Geothallus).

21. Sphaerocarpos Boehm., in Ludwig, Definit. Gen. Pl. ed. 3, 501. 1760. After sphaero- = globose, carpus = fruit, referring to the swollen involucres surrounding the gametangia. Fig. 158: G-J Thalli prostrate, small and delicate, pale green, less than 1 cm long, without air chambers and pores, thickened in the middle, unistratose towards the margins, the margins sinuose-lobed, the median thallus surface of fertile plants covered by numerous swollen involucres. Dioecious; female and male thalli quite different in size and shape. Female thalli suborbicular, 5-8 mm long, involucres 1-2 mm long, obovate to cylindrical, with a very narrow mouth. Male thalli smaller, 1-3 mm long, elongate and forked-lobed, involucres tiny, less than 1 mm long, flask-shaped. Spores large, often in tetrads, yellow-brown, tetrads 100-180 µm in diameter, surface of individual spores covered by a fine network of areoles, the areoles 6-12 µm in diameter. Asexual reproduction by specialized devices lacking. Sphaerocarpos is a widespread subtropical-mediterranean genus of about 12 species. One species, S. muccilloi (Fig. 158: G-J) Vianna has been recorded from Rio Grande do Sul, southern Brazil (Vianna, 1981a; Gradstein & Pinheiro da Costa, 2003) and an unidentified species of Sphaerocarpos has been reported from the surroundings of Lima in Peru (Carillo & Chanco, 1971). Further species of Sphaerocarpos have been recorded from the southern U. S. A., Uruguay, Argentina, and Chile (e.g., Gradstein et al., 1983; Schuster, 1992b). The Peruvian record is of considerable importance as it is the only one of the order Sphaerocarpales within the boundaries of the Neotropics. The identity of the Peruvian specimen remains unknown, however; material could not be found in the Herbarium of the Universidad Nacional Mayor de San Marcos, Lima (Ms Jasmin Opisso, pers. comm., 8. April 2003). A field search in potentially suitable habitats in Peru should be undertaken to trace this important taxon. The species of Sphaerocarpos usually grow on temporarily moist, compact soils in slightly shaded, rather arid places. The small, prostrate thalli densely covered by numerous swollen involucres are unmistakable.

Bischler et al. 208 DOUBTFUL , EXCLUDED, OR ERRONEOUSLY REPORTED TAXA

Mannia fragrans (Balb.) Frye & L.Clark, distributed in Europe, western and eastern Asia, Canada, and in the U. S. A. south to Alabama, Arizona, Georgia, New Mexico, and Texas, has been erroneously recorded from Cuba (Müller, 1952 = Asterella elegans, see Grolle, 1975) and from north-western Peru (Hegewald & Hegewald, 1977 = Mannia hegewaldii Bischl. sp.nov., see this treatment). Marchantia androgyna L., lectotypified by the Italian syntype (see Grolle, 1968) is excluded by lectotypification. The Jamaican syntype belongs to M. chenopoda. Marchantia emarginata Reinw., Blume & Nees (Asia, Oceania), and M. platycnemos Schwägr., a synonym of M. emarginata, have been recorded by Montagne (1839a) from the Neotropics. The corresponding specimens belong to M. papillata subsp. papillata. Marchantia linearis Lehm. & Lindenb., a Himalayan species, has been recorded from the West Indies by Gottsche, Lindenberg & Nees (1846) and other authors. All specimens seen so far belong to M. inflexa. Marchantia macropora Mitt. (New Zealand) and M. wilmsii Steph. (Africa; synonym of M. pappeana Lehm.) have been erroneously cited from Bolivia (Stephani, 1916) and Colombia (Jack & Stephani, 1892). They do not occur in the Neotropics. Monoclea crispata Hook., Bot. Misc. 1: 117. 1829. Dendroceros crispatus (Hook.) Nees, in Gottsche et al., Syn. hepat. 579. 1846. Type. Windward Islands. St. Vincent: Guilding s.n., ex hb Hooker (isotype, NY, slide) = ? Dendroceros sp. (definitely a hornwort; not Monoclea). Preissia Corda. Genuine neotropical representatives of this genus are unknown. Preissia barbata and P. mexicana belong to Marchantia chenopoda, P. cucullata to Lunularia cruciata. Four specimens from Mexico (NY) identified as P. quadrata (Scop.) Nees (Svilha, 1941) belong to M. chenopoda; those of P. quadrata reported from Panama (Svilha, 1942) and Bolivia (Stephani, 1916) presumably belong to M. chenopoda as well (not seen). The species has been cited from Uruguay (Herter, 1933). Its occurrence in the Neotropics needs confirmation. Riccia bialbistrata Hässel, Opera Lilloana 7: 243. 1963. Type. Argentina. Buenos Aires: Delta del Paraná, Arroyo Abra Vieja, Sep 1956, Hässel de Menéndez 5 (BA 9383) (n.v.); the status of his species remains doubtful. It has close affinity with R. lindmanii and has been recorded also from Costa Rica (Jovet-Ast, 1981). Riccia bifurca Hoffm. The neotropical record (Müller, 1955) belongs to R. lindmanii (Hässel, 1963). Riccia campbelliana Howe subsp. austrigena R.M.Schust., Hep. North America 6: 583. 1992. Type. Argentina. Tucumán: Tafi, La Cienaga, near Casa Grande, 2500 m, 1950, Sleumer s.n. (holotype,

Bischler et al. 209 LIL, n.v.). Whether this taxon belongs to R. campbelliana remains questionable. Riccia canaliculata Hoffm., a holarctic species, has been cited from Brazil (Nees, 1833; Ångström, 1876; Schiffner & Arnell, 1964). Its occurrence in the Neotropics needs confirmation. Riccia chartacea K.I.Goebel, Organogr. Pfl. ed. 2, 2 (1): 630. 1915. Type. Brazil (n.v.). This species has not been recorded since its original description. Riccia fluitans L. The specimens recorded from Brazil (Bauer, 1899; Nees, 1833) and Argentina (Massalongo, 1906) belong to R. stenophylla (see Hässel, 1963; Schuster, 1992b), and also those recorded from the Galápagos Islands (Weber, 1975) and most of those of Central America (Frye & Clark, 1937). The species has furthermore been recorded from Costa Rica, Cuba, Haiti, and Jamaica by Howe (1923), from Brazil (Rio Grande do Sul) and Paraguay by Stephani (1897), and from Uruguay by Herter (1933) and recently from Pernambuco (Yano & Andrade-Lima, 1987), Goiás and Rio de Janeiro (Yano et al., 1987). Whether the species occurs in the Neotropics remains questionable. Riccia glauca L., a cosmopolitan but often confused species has been recorded from Mexico (Schlechtendahl & Chamisso, 1831), Dominica (Spruce, 1895), Brazil (Raddi, 1823), Chile (Montagne, 1850), and Argentina (Buenos Aires, Massalongo, 1917). Its presence in the Neotropics needs confirmation. Riccia glauca var. subinermis (Lindb.) Warnst. The specimen recorded from Argentina (Müller, 1955) belongs to R. sorocarpa (Hässel, 1963). Riccia hirta (Aust.) Underw. This North-American species has been recorded from Rio Grande do Sul (Vianna, 1985). The Brazilian record needs confirmation. Riccia nigrescens Mont., in A.D.Orbigny, Voy. Amér. mérid. 7, Bot(1): 15. 1839. Type. Argentina. Rio Negro: d'Orbigny 105 (PC). This species belongs to Anthoceros. Riccia reticulata Sw., Prodr. 146. 1788. Type. Jamaica. Swartz (S). The type material contains fern prothallia. Riccia subtilis (Steph.) Steph., Sp. hepat. 1: 39. 1898. Ricciella subtilis Steph., Bih. Kongl. Svenska Ventensk.-Akad. Handl. 23 (III, 2): 31. 1897. Type. Paraguay. Lindman B 357 (G). The specimen is sterile. Riccia sullivantii Aust., a synonym or a subspecies of Riccia hübeneriana Lindenb., has been reported once from Cuba (Howe 1923) (n.v.). The report needs confirmation.

ACKNOWLEDGMENTS The Directors and Keepers of the Herbaria cited in the text, including BA, BM, CAY, CINC, COL,

Bischler et al. 210 CORD, E, EGR, F, FH, FI, FLAS, G, GOET, H, HAC, HEID, ICN, JE, K, KSP, LIL, LISU, LPS, M, MADJ, MANCH, MEXU, MO, NICH, NY, OXF, PC, PMA, RB, S, SJ, SP, STR, TENN, TUB, U, UC, US, W, YU, and R. Düll, the late M. H. Fulford, P. & E. Hegewald, H. Hürlimann, A. SchäferVerwimp, the late O. H. Volk, A. T. Whittemore, and R. Wingfield are acknowledged for the loan of specimens and for assistance with herbarium work. We are very grateful to Riclef Grolle, Barbara Thiers and, especially, Alan Whittemore for carefully checking and correcting the manuscript. Hélene Bischler-Causse is grateful to Michèle Dumont who made the photographic prints and to Bryophytorum Bibliotheca and Cryptogamie, Bryologie-Lichénologie for permission of reproduction of parts of text and illustrations. Rob Gradstein is grateful to Elena Reiner-Drehwald for preparing the plate of Monoclea gottschei. Noris Salazar-Allen is grateful to Hélene Bischler for proposing to undertake the study of Cyathodium and for her valuable and unselfish support throughout her studies and during a short visit to the Muséum in Paris; to M. Sc. María Isabel Morales, University of Costa Rica, for providing assistance and laboratory space to process collections and for providing comments to improve the manuscript; to Dr. Gabriela Hässel for comments to the manuscript; to M. Sc. Mariela Bermúdez and Biólogo Rafael Chacón of the Museo Nacional de Costa Rica and Ing. Eduardo Lépiz (INBIO) for logistic support, field assistance and processing of samples in Costa Rica; to her assistant Lic. Clementina Chung for her patience and enduring field and laboratory assistance, to her students Betzaida Cuadra, Alex Espinosa (who first discovered Cyathodium in the Zoological Park in Costa Rica), Lic. Eduardo Montenegro, Néstor Rivera, Amauris Velásquez and Juan Carlos Villarreal for their kind assistance during field trips in Panama; to Mara Rodríguez whose collections and slides of Cyathodium for an undergraduate study were of great help; to Lic. Alberto Castillo for valuable help with illustrations and to Lic. Jorge Cevallos (STRI) who kindly suggested and tried out different techniques to improve the quality of fresh and previously fixed specimens for SEM; and to the Autoridad Nacional del Ambiente (A.N.A.M.) of Panama and the Ministerio de Ambiente y Energía, Sistema Nacional de Áreas de Conservación of Costa Rica for permission to do field work and collecting in Panama and Costa Rica respectively. Field work of N. Salazar Allen was supported by the Smithsonian Tropical Research Institute.

Bischler et al. 211

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Bischler et al. 227

INDEX OF SCIENTIFIC NAMES New names are in boldface, synonyms in italics. Primary page references are in boldface. An asterisk (*) indicates a page having an illustration or map. Aitonia chlorocarpa, 158 Aitonia elongata, 158 Aitonia intermedia, 151 Aitonia lanigera, 158 Aitonia mexicana, 158 Aitonia peruviana, 158 Aitonia subplana, 159 Aitonia valida, 158 Aitonia wrightii, 155 Angiocarpus, 75 Angiocarpus curtisii, 76 Anthoceros, 209 Anthocerotae, 21 Arnelliella, 143 Asterella, 8, 11, 14, 17, 19, 28, 120, 121, 122, 247 Asterella lateralis, 128 Asterella alpina, 137, 138, 247 Asterella austinii, 130 Asterella bolanderi, 133 Asterella boliviana, 134 Asterella chilensis, 122, 127 Asterella dominicensis, 122, 124, 125, 126, 136, 139, 247 Asterella echinella, 122, 128, 129, 131, 132, 247 Asterella elegans, 122, 127, 128, 129, 130, 131, 132, 136, 208, 247 Asterella evansii, 124 Asterella fissisquama, 134 Asterella grollei, 137 Asterella lateralis, 122, 129, 131, 132, 133, 136, 247 Asterella leptophylla, 124 Asterella lindenbergiana, 135, 136 Asterella macropoda, 122, 128, 134, 135, 247 Asterella marginata, 121 Asterella muscicola, 137 Asterella palmeri, 137, 138 Asterella pringlei, 133, 137, 138, 139, 247 Asterella reticulata, 130 Asterella rugosa, 138, 140, 247 Asterella saccata, 121, 137, 138 Asterella sect. Brachyblepharis, 124 Asterella sect. Lindenbergianae, 128

Asterella sect. Pappiae, 128 Asterella sect. Phragmoblepharis, 128 Asterella sect. Saccatae, 137 Asterella stahlii, 139 Asterella subgen. Asterella, 123 Asterella subgen. Asterella sect. Brachyblepharis, 124 Asterella subgen. Brachyblepharis, 124 Asterella subgen. Phragmoblepharis, 123, 127 Asterella subgen. Saccatae, 123, 137, 141 Asterella tenella, 121, 124 Asterella venosa, 122, 124, 126, 127, 247, 252 Asterella versicolor, 128, 136, 247 Asterella wrightii, 130 Astrella macropoda, 132 Athalamia, 11, 12, 27, 115, 116 Athalamia andina, 116, 117, 246, 247, 249 Athalamia hyalina, 117, 118 Athalamia pinguis, 116 Athalamia pygmaea, 117, 118, 247, 249 Athalamia robusta, 117 Athalamia spathysii, 117, 118 Aytonia, 119, 145, 156 Aytonia crenulata, 148 Aytonia evansii, 159 Aytonia jamaicensis, 152 Aytonia rupestris, 145, 156, 157 Aytoniaceae, 11, 12, 13, 16, 17, 25, 119, 120 Aytonioideae, 120 Boschia, 96 Boschia weddellii, 96, 98 Calobryales, 6 Chlamidium, 184 Chlamidium indicum, 184, 186 Clevea, 115, 116 Clevea andina, 117 Clevea hyalina, 116 Cleveaceae, 16, 17, 24, 115, 116 Corsinia, 11 13, 14, 26, 94, 95 Corsinia coriandrina, 14, 15, 95, 96, 234, 235, 244, 246 Corsinia coriandrina fo. gymnocarpa, 95 Corsinia marchantioides, 95 Corsiniaceae, 11, 12, 14, 16, 23, 94 Corsiniineae, 11, 12, 13, 14, 16, 23

Bischler et al. 228 Corsinioideae, 94 Cronisia, 3, 7, 9, 18, 26, 95, 96, 98 Cronisia fimbriata, 97, 244, 246 Cronisia mexicana, 97 Cronisia paradoxa, 96, 98 Cronisia weddellii, 96, 97, 98, 244, 246 Cronisioideae, 94 Cryptocarpus, 75 Cryptocarpus curtisii, 76 Cryptomitrium, 28, 121, 141 Cryptomitrium tenerum, 141, 248, 249 Cyathodiaceae, 24, 99 Cyathodium, 9, 10, 12, 16, 26, 99, 100, 102, 104 Cyathodium africanum, 103, 104 Cyathodium aureonitens, 102 Cyathodium barodae, 103 Cyathodium bischlerianum, 100, 101, 102, 105, 107, 108, 244, 245 Cyathodium cavernarum, 18, 99, 100, 101, 102, 103, 104, 105, 107, 108, 245 Cyathodium cf. foetidissimum, 106 Cyathodium foetidissimum, 18, 100, 101, 105, 106, 245 Cyathodium mexicanum, 103 Cyathodium spruceanum, 100, 101, 102, 104, 106, 107, 109, 244, 245, 246 Cyathodium steerei, 100, 101, 106, 108, 109, 246 Cyathodium subgen. Cyathodium, 100 Cyathophora, 142 Cyathophora. angustifolia, 142 Cyatophora mexicana, 187 Dendroceros, 208 Dendroceros crispatus, 208 Dumortiera, , 9, 16, 17, 22, 24, 28, 168, 169, 200 Dumortiera hirsuta, 15, 106, 169, 170, 173, 200, 235, 250, 253 Dumortiera hirsuta fo. depauperata, 169 Dumortiera hirsuta var. angustior, 169 Dumortiera hirsuta var. brasiliensis, 170 Dumortiera hirsuta var. inermedia, 170 Dumortiera hirsuta var. irrigua, 169 Dumortiera hirsuta var. latior, 170 Dumortiera hirsuta var. trichopus, 170 Dumortiera irrigua, 169 Dumortiera nepalensis, 169, 173 Dumortieroideae, 168 Duvalia, 142 Duvalia brevipedunculata, 141 Duvalia gayana, 141 Duvalia rupestris, 142 Duvalia tenera, 141

Exormotheca, 13, 14, 16, 27, 114 Exormotheca pustulosa, 15, 18, 114, 115, 235, 246, 247 Exormothecaceae, 11, 14, 16, 24, 114 Fimbraria, 121 Fimbriaria, 121 Fimbriaria alpina, 138 Fimbriaria arsenii, 132 Fimbriaria atrispora, 132, 133 Fimbriaria bakeri, 139 Fimbriaria boliviana, 134 Fimbriaria canelensis, 134 Fimbriaria cubanensis, 130 Fimbriaria echinella, 128 Fimbriaria elegans, 130, 132 Fimbriaria elegans var. cubensis, 130 Fimbriaria fissisquama, 134 Fimbriaria lindmannii, 126 Fimbriaria mandonii, 134 Fimbriaria mexicana, 132 Fimbriaria pringlei, 124, 138 Fimbriaria stahlii, 139 Fimbriaria subgen. Brachyblepharis, 124 Fimbriaria venosa, 126 Fissidens, 104 Fissidens flaccidus, 106 Funicularia, 96 Funicularia bischleriana, 98 Funicularia weddellii, 98 Geothallus, 207 Grimaldia, 142 Grimaldia californica, 143 Grimaldia chilensis, 119 Grimaldia debilis, 119 Grimaldia dichotoma, 142 Grimaldia hemisphaerica, 163 Grimaldia peruviana, 187, 191 Grimaldia stellaris, 113 Grimaldia swartzii, 186 Hygrophila, 169 Hygrophila irrigua, 169 Hygrophila nepalensis, 169 Hymenophyllaceae, 21 Hypenantron, 121 Hypenantron ciliatum, 121 Jungermanniales, 6, 21, 22 Jungermanniidae, 3, 6, 18 Lejeuneaceae, 104, 106 Lepidoziaceae, 21, 22 Liquidambar, 139 Lunularia, 10, 12, 13, 14, 16, 24, 27, 165 Lunularia alpina, 118 Lunularia cruciata, 165, 166, 168, 208, 235, 250, 253

Bischler et al. 229 Lunularia cruciata subsp. thaxteri, 166 Lunularia thaxteri, 166, 168 Lunularia. cruciata fo. thaxteri, 166 Lunulariaceae, 25, 165 Mannia, 28, 121, 142, 143 Mannia androgyna, 142, 143 Mannia californica, 24, 27, 143, 144, 248 Mannia fragrans, 208 Mannia hegewaldii, 143, 144, 145, 208, 248, 253 Mannia michelii, 142 Mannia subgen. Arnelliella, 143 Mannia subgen. Mannia, 143 Mannia subgen. Xeromannia, 143 Mannia triandra, 143, 145 Marchantia, 7, 9, 10, 11, 12, 13, 14, 16, 17, 29, 106, 168, 169, 174, 175 Marchantia anapodocarpos, 186 Marchantia androgyna, 142, 208 Marchantia berteroana, 18, 175, 177, 179, 250, 253 Marchantia berteroana var. anactis, 177 Marchantia berteroana var. biflora, 177, 178 Marchantia berteroana var. polylepida, 177, 178 Marchantia bescherellei, 187 Marchantia brasiliensis, 186, 191 Marchantia breviloba, 15, 176, 184, 185, 186, 191, 251 Marchantia caracensis, 192 Marchantia cartilaginea, 186 Marchantia cephaloscypha var. breviradiata, 179 Marchantia chenopoda, 168, 176, 184, 186, 188, 191,194, 208, 235, 251, 252, 253 Marchantia chenopoda var. cartilaginea, 186 Marchantia columbica, 190 Marchantia columbica, 187 Marchantia cruciata, 166 Marchantia dillenii, 186 Marchantia discoidea, 177, 178 Marchantia disjuncta, 191 Marchantia domingensis, 194 Marchantia ecuadorensis, 187, 190 Marchantia elliottii, 191 Marchantia emarginata, 208 Marchantia faxinensis, 197, 198 Marchantia flabellata, 187, 190 Marchantia foliacea, 175 Marchantia hemisphaerica, 163 Marchantia hirsuta, 169 Marchantia hyalina, 116 Marchantia inflexa, 176, 186, 191, 192, 194, 198, 208, 252, 253

Marchantia irrigua, 169 Marchantia lamellosa, 179, 180 Marchantia linearis, 208 Marchantia macropora, 208 Marchantia Marchantiotypus, 176 Marchantia martinicensis, 186 Marchantia notarisii, 187, 191 Marchantia paleacea, 176, 186, 194, 195, 196, 235, 252, 253 Marchantia paleacea subsp. diptera, 197 Marchantia paleacea subsp. paleacea, 194, 197 Marchantia papillata, 176, 194, 197, 198, 252, 253 Marchantia papillata subsp. grossibarba, 198 Marchantia papillata subsp. papillata, 197, 198, 208 Marchantia pappeana, 208 Marchantia peruviana, 187 Marchantia pittieri, 187, 189 Marchantia platycnemos, 208 Marchantia plicata, 175, 179, 181, 251 Marchantia polymorpha, 174, 175, 176, 181, 184, 235, 251, 253 Marchantia polymorpha subsp. montivagans, 184 Marchantia polymorpha subsp. polymorpha, 184 Marchantia polymorpha subsp. ruderalis, 184 Marchantia pusilla, 197, 198 Marchantia quinqueloba, 191 Marchantia sect. Astromarchantia, 176 Marchantia sect. Chlamidium, 184 Marchantia squamosa, 194 Marchantia Stellatae, 176 Marchantia subandina, 197, 198 Marchantia subgen. Chlamidium, 175, 184, 196, 198 Marchantia subgen. Marchantia, 175, 176, 184, 196 Marchantia swartzii, 186 Marchantia tenella, 121 Marchantia tenera, 141 Marchantia tholophora, 194, 196 Marchantia wilmsii, 208 Marchantiaceae, 12, 14, 24, 29, 168 Marchantiales, 3, 4, 6, 7, 8, 9, 10, 11, 12, 14, 15, 16, 18, 19, 21, 2225, 29, 32, 94, 168, 173 Marchantiidae, 3, 4, 6, 8, 10, 15, 18, 19 Marchantiineae, 11, 12, 15, 24 Marchantioideae, 168 Metzgeriales, 6, 22 Monoclea, 9, 10, 11, 12, 13, 14, 16, 17, 19, 22, 200, 208

Bischler et al. 230 Monoclea forsteri, 200 Monoclea gottschei, 173, 200, 253 Monoclea gottschei subsp. elongata, 201, 202, 204, 252, 253 Monoclea gottschei subsp. gottschei, 200, 201, 204 Monocleaceae, 199 Monocleae, 199 Monocleales, 3, 4, 6, 7, 9, 14, 15, 19, 22, 199 Myriorrhynchus, 96 Myriorrhynchus fimbriatus, 96, 97 Neesiella, 142 Neesiella chilensis, 119 Neesiella rupestris, 142 Notothylas, 104 Oxymitra, 9, 11, 12, 14, 25, 29 Oxymitra androgyna, 30 Oxymitra cristata, 30 Oxymitra incrassata, 30, 31, 235, 236, 246 Oxymitra paleaca, 30 Oxymitra pyramidata, 30 Oxymitraceae, 16, 22, 23, 29 Philonotis, 104 Pinus, 132 Plagiochasma, 7, 11, 12, 14, 17, 28, 119, 120, 145, 248 Plagiochasma argentinicum, 148, 154, 235, 248, 249 Plagiochasma bolivianum, 187, 191 Plagiochasma bornmülleri, 159 Plagiochasma brasiliense, 159 Plagiochasma chlorocarpum, 158 Plagiochasma cordatum, 145 Plagiochasma crenulatum, 15, 147, 148, 149, 248, 249 Plagiochasma crenulatum var. ß minus, 155 Plagiochasma cuneatum, 147, 150, 154, 249 Plagiochasma elongatum, 158 Plagiochasma elongatum var. ß leucolepis, 150 Plagiochasma evansii, 159 Plagiochasma hosseusii, 159 Plagiochasma intermedium, 147, 151, 152, 249 Plagiochasma jamaicense, 147, 152, 153, 155, 156, 249 Plagiochasma landii, 147, 153, 154, 249 Plagiochasma lanigerum, 158 Plagiochasma mexicanum, 158 Plagiochasma münchianum, 147, 154, 155, 156, 249 Plagiochasma peruvianum, 158 Plagiochasma rupestre, 14, 15, 24, 27, 117, 146, 156, 157, 234, 235

Plagiochasma rupestre fo. glaucum, 159 Plagiochasma rupestre var. rupestre, 158, 162, 249 Plagiochasma rupestre var. volkii, 158, 162, 249 Plagiochasma subgen. Micropylum, 146, 156 Plagiochasma subgen. Plagiochasma, 146 Plagiochasma subplanum, 159 Plagiochasma validum, 158 Plagiochasma validum. var. ß minus, 158 Plagiochasma wrightii, 147, 153, 155, 156, 249 Platanus, 139 Platycoaspis, 141 Platycoaspis tenera, 141 Pottiaceae, 104 Preissia, 168, 208 Preissia barbata, 187, 191, 208 Preissia cucullata, 166, 208 Preissia mexicana, 187, 208 Preissia quadrata, 208 Pteropsiella, 22 Pycnoscenus, 30 Pycnoscenus pyramidatus, 30 Quercus, 132, 139 Racopilum tomentosum, 104 Reboulia, 28, 120, 162 Reboulia chlorocarpa, 158 Reboulia hemipsphaerica, 144 Reboulia hemisphaerica, 14, 15, 162, 163, 165, 235, 250, 253 Reboulioideae, 120 Riccia, 3, 7, 9, 10, 11, 13, 14, 16, 17, 18, 26, 31, 32, 33, 96 Riccia natans var. brasiliensis, 94 Riccia aggregata, 59 Riccia albida, 35, 38, 39, 236 Riccia albopunctata, 35, 39, 40, 43, 67, 236 Riccia amazonica, 74, 243 Riccia americana, 55 Riccia andina, 57, 8 Riccia antarctica, 62 Riccia atromarginata, 15, 34, 40, 41, 54, 236, 243 Riccia atromarginata subsp. atromarginata, 41, 69 Riccia atromarginata var. atromarginata, 41, 69 Riccia austinii, 55 Riccia australis, 38, 41, 42, 236 Riccia bahiensis, 243 Riccia bahiensis, 79, 91, 241, 243 Riccia bialbistrata, 208 Riccia bifurca, 208

Bischler et al. 231 Riccia bistriata, 61 Riccia boliviensis, 36, 42, 43, 236, 237, 243 Riccia brandegei, 90 Riccia brasiliensis, 35, 43, 57, 237, 243 Riccia breutelii, 38, 44, 45, 48, 237, 243 Riccia brittonii, 37, 44, 45, 48, 237, 243 Riccia campbelliana, 35, 40, 45, 46, 237, 243 Riccia campbelliana subsp. austrigena, 46, 208 Riccia cana, 67 Riccia canaliculata, 209 Riccia canescens, 67 Riccia catalinae, 90 Riccia cavernosa, , 79, 89, 90, 91, 92, 242, 243 Riccia chartacea, 209 Riccia chiapasensis, 78, 80, 83, 241, 243 Riccia chilensis, 62 Riccia ciliata, 68 Riccia corcovadensis, 60, 72 Riccia coriandrina, 95 Riccia crassifrons, 79, 80, 81, 241, 243 Riccia crustata, 39 Riccia crystallina, 89, 91, 92, 242, 243 Riccia cubensis, 35, 46, 49, 237 Riccia curtisii, 75, 76, 77, 86, 241, 243 Riccia dorsiverrucosa, 57, 58 Riccia dussiana, 78, 81, 82, 241, 242, 243 Riccia echinatispora, 74 Riccia ekmanii, 36, 47, 58, 237, 238, 243 Riccia elliottii, 38, 44, 45, 48, 238, 243 Riccia enyae, 35, 48, 49, 237, 238, 243 Riccia erythrocarpa, 37, 49, 50, 238, 243 Riccia fimbriata, 96, 97 Riccia flavispora, 51 Riccia fluitans, 77, 84, 89, 209 Riccia frostii, 78, 82, 84, 235, 241, 242, 243 Riccia fruchartii, 38, 50, 51, 238, 243 Riccia gaumeri, 44 Riccia geissleriana, 78, 82, 83, 241, 242, 243 Riccia glauca, 33, 51, 209 Riccia glauca var. subinermis, 209 Riccia grandis, 38, 42, 51, 238, 243 Riccia grandisquama, 55 Riccia hegewaldiana, 78, 83, 84, 242, 243 Riccia hirta, 209 Riccia horrida, 35, 49, 52, 65, 68, 238 Riccia hosseusii, 87 Riccia howellii, 52, 53, 237, 238, 243 Riccia hübeneriana, 87, 209 Riccia ianthina, 37, 53, 54, 238, 243 Riccia incrassata, 30 Riccia indusiata, 55 Riccia iodocheila, 34, 41, 54, 238 Riccia jovet-astiae, 78, 84, 242, 243 Riccia lamellosa, 15, 37, 55, 56, 69, 235, 238,

243 Riccia lamellosa var. americana, 55 Riccia lanigera, 74 Riccia limicola, 79, 84, 85, 89, 242, 243 Riccia lindmanii, 35, 43, 56, 57, 208, 238, 239, 243 Riccia macrocarpa, 46 Riccia mauryana, 36, 57, 58, 238, 239, 243 Riccia membranacea, 74, 75, 86, 241, 243 Riccia montagnei, 90 Riccia natans, 92 Riccia natans var. brasiliensis, 92 Riccia nigrella, 36, 58, 59, 239, 243 Riccia nigrescens, 209 Riccia ochrospora, 91 Riccia oerstediana, 87 Riccia paradoxa, 98 Riccia paraguayensis, 84, 85, 86, 242, 243 Riccia paranaensis, 79, 86, 87, 242, 243 Riccia plana, 91, 92 Riccia plano-biconvexa, 38, 59, 60, 66, 239, 243 Riccia pyramidata, 30 Riccia rauhii, 62 Riccia reticulata, 209 Riccia ridleyi, 36, 47, 58, 60, 61, 72, 74, 239, 243 Riccia sanguineisporis, 37, 61, 239, 243 Riccia sect. Cavernosae, 89 Riccia sect. Riccia, 34 Riccia sect. Ricciella, 77, 78, 82 Riccia sect. Spongodes, 89 Riccia sorocarpa, 37, 45, 50, 51, 62, 63, 209, 239, 240, 243 Riccia sorocarpa subsp erythrophora, 63 Riccia sorocarpa subsp. sorocarpa, 63 Riccia spegazziniana, 76 Riccia spruceana, 80, 241 Riccia squamata, 36, 43, 64, 239, 240, 243 Riccia stenophylla, 78, 84, 85, 87, 88, 89, 209, 242, 243 Riccia subdepilata, 35, 52, 65, 239, 240, 243 Riccia subgen. Leptoriccia, 33, 34, 74 Riccia subgen. Riccia, 33, 34, 83 Riccia subgen. Ricciella, 33, 34, 77, 81, 82, 83 Riccia subgen. Ricciella sect. Ricciella, 77 Riccia subgen. Ricciella sect. Spongodes, 77 Riccia subgen. Thallocarpus, 33, 34, 75 Riccia subnigella, 64 Riccia subplana, 37, 53, 60, 65, 66, 240, 243 Riccia subsimilis, 76, 86 Riccia subtilis, 209 Riccia sullivantii, 209 Riccia synspora, 76

Bischler et al. 232 Riccia taeniaeformis, 38, 66, 67, 240, 243 Riccia tenuilimbata, 59, 60 Riccia trabutiana, 54 Riccia trichocarpa, 15, 35, 67, 68, 240, 243 Riccia velutina, 92 Riccia viannae, 37, 68, 69, 240, 243 Riccia violacea, 34, 41, 54, 69 Riccia violacea var. laevis, 70, 240, 243 Riccia violacea var. violacea, 47, 69, 70, 240 Riccia vitalii, 36, 58, 61, 71, 72, 74, 240, 243 Riccia weberbaueri, 50 Riccia weinionis, 36, 47, 58, 61, 60, 72, 73, 74, 240, 241, 243 Ricciaceae, 11, 12, 14, 16, 22, 23, 31, 32 Ricciella, 77 Ricciella fluitans, 77 Ricciella macropora, 85 Ricciella membranacea, 74 Ricciella subsimilis, 76 Ricciella subtilis, 209 Ricciineae, 11, 13, 16, 23 Ricciocarpos, 9, 26, 32, 92 Ricciocarpos natans, 92, 93, 94, 235, 243, 246 Ricciocarpos natans fo. terricola, 92, 94 Ricciocarpos velutinus, 92, 94 Riella, 21, 206 Riella americana, 206 Riella notarisii, 206 Riellaceae, 206 Ruppinia, 145, 156 Ruppinia chlorocarpa, 158 Ruppinia elongata, 158 Ruppinia intermedia, 151 Ruppinia lichenoides, 156

Ruppinia mexicana, 158 Ruppinia peruviana, 158 Ruppinia rupestris, 145, 157 Ruppinia valida, 158 Sauteria, 12, 27, 116, 118 Sauteria alpina, 118 Sauteria berteroana, 119 Sauteria chilensis, 119, 247, 249 Selaginella, 108 Sphaerocarpaceae, 205, 206, 207 Sphaerocarpales, 3, 4, 6, 7, 9, 19, 21, 205 Sphaerocarpos, 21, 31, 207 Sphaerocarpos muccilloi, 207, 252 Targionia, 26, 109, 110 Targionia bifurca, 110 Targionia convoluta, 111 Targionia hypophylla, 14, 15, 109, 110, 113, 235, 246 Targionia lorbeeriana, 111 Targionia mexicana, 110 Targionia robusta, 117 Targionia stellaris, 110, 113, 246 Targionia subgen. Prototargionia, 110 Targionia subgen. Targionia, 110 Targioniaceae, 17, 24, 109 Targioniaceae subfam. Cyathodioideae, 99 Taxiphyllum taxirameum, 106 Tessellina, 30 Tessellina pyramidata, 30 Thallocarpus, 75 Thallocarpus curtisii, 76 Thallocarpus synsporus, 76 Treubiales, 6 Weinmannia, 135

(CAPTIONS) Herbarium acronyms are given only for specimens not cited in the text. FIG. 1. Epidermal pores, thallus structure and scales. A. Plagiochasma rupestre, epidermal pore, a simple opening. B. Corsinia coriandrina, epidermal pore bounded by single ring of cells. C. Oxymitra incrassata, epidermal pore bounded by single ring of cells. D. Exormotheca pustulosa, epidermal pore strongly elevated above epidermis. E. Plagiochasma argentinicum, epidermal pore bounded by several rings of concentric cells, cuticle papillose. F. Lunularia cruciata, epidermal pore bounded by several concentric rings of cells. G. Marchantia paleacea, compound pore. H. Dumortiera hirsuta, dorsal surface without pores. I. Plagiochasma rupestre, cross section of thallus, with several layers of empty air chambers. K. Lunularia cruciata, cross section of thallus, with one layer of air chambers with chlorophyllose filaments. L. Reboulia hemisphaerica, arrangement of

Bischler et al. 233 scales in two rows on ventral side of thallus. M. Lunularia cruciata, oil cells in basal tissue. A from France, Bischler & Baudoin 75239 (PC), B from Corsica, Fridlender 94927 (PC), C from Portugal, Bischler 94788 (PC), D from South Africa, Perold 3056 (PC), E from Volk 6/669, F from France, Bischler & Boisselier 95747 (PC), G from Jovet-Ast s.n., H from Japan, Long & Yamasaki 24829 (PC), I from France, Bischler 93566 (PC), K from France, Bischler s.n. (PC), L from Corsica, Fridlender 93967 (PC), M from France, Bischler et al. 93613 (PC). (scale bars A-C, E-H = 10 µm, D = 50 µm, I-M = 100 µm).

FIG. 2. General characteristics, asexual reproduction and antheridia. A. Corsinia coriandrina, scales at thallus apex. B. Smooth rhizoid. C. Pegged rhizoid. D. Riccia frostii, male and female thalli. E. Riccia lamellosa, dry but still living thalli. F. Ricciocarpos natans. G. Lunularia cruciata, gemma cup. H. Marchantia polymorpha, immature gemma cup. I. Gemma. K. Exormotheca pustulosa, groove with antheridia. L. Marchantia chenopoda, papillae on male receptacle. M. Lunularia cruciata, male receptacles. A from Italy, Bischler 94888 (PC), B-C from Canada, Boudier 93680 (PC), D from Egypt, Jovet-Ast & Bischler 67015 (PC), E from Israel, Jovet-Ast & Bischler 64160 (PC), F from France, Noailles s.n. (PC), G from Israel, Jovet-Ast & Bischler s.n. (PC), H from Hungary, Horton s.n. (PC), I from New Zealand, Campbell s.n. (PC), K from South Africa, Perold & Koekemoer 2941 (PC), L from Israel, Jovet-Ast & Bischler 64210 (PC), M from Baudoin s.n. (scale bars A, E, G-H, K = 100 µm, B-C, L = 10 µm, D, F, M = 1 mm, I = 40 µm). FIG. 3. Gametangia and gametangiophores. A. Plagiochasma rupestre, male and female receptacles. B. Marchantia chenopoda, antheridiophore. C. Lunularia cruciata, female receptacles. D-E. Oxymitra incrassata, D. Involucres and scales, E. Thallus with involucres. F. Corsinia coriandrina, male and female gametangia in grooves. G. Marchantia chenopoda, pseudoperianths bounding sporophytes. H. Targionia hypophylla, mussel-shaped involucres. I-K. Archegoniophores. I. Plagiochasma rupestre. K. Reboulia hemisphaerica. A, I from France, Bischler & Baudoin 75239 (PC), B, G from Baudoin s.n., C from Israel, Jovet-Ast & Bischler 64210 (PC), D, E from Algeria, Baudoin s.n. (PC), F from Italy, Bischler 94888 (PC), H from Israel, Jovet-Ast & Bischler 64109 (PC), K from Spain, Jovet-Ast & Bischler 69039 (PC). (scale bars A-B, D, F, H-K = 1 mm, C, E = 2 mm, G = 100 µm). FIG. 4. Phylogram of the strict consensus tree of the 2080 most parsimonious trees derived from a data set of 43 morphological characters. CI = 0.458, RI = 0.749 (bootstrap values above lines, Bremer indices below lines). Bold line = Aytoniaceae set (from Boisselier-Dubayle et al., 2002). FIG. 5. Phylogram of the strict consensus tree of the six most parsimonious trees derived from the (LSU) rDNA gene data set. CI = 0.461, RI = 0.591 (bootstrap values above lines, Bremer indices below lines). Bold line = Aytoniaceae (from Boisselier-Dubayle et al., 2002).

Bischler et al. 234 FIG. 6. Oxymitra incrassata. A. Thallus with involucres. B. Thallus with antheridial groove. C. Cross section of thallus. D. Epidermal pores, surface view. E. Cross section of epidermal pore. F. Cross section of epidermis and air chambers. G. Arrangement of scales on ventral side of thallus, H. Scales. I. Appendages of scales. K. Scale of female receptacle. L. Cross section of involucre. M. Epidermal pore of involucre. A-C, F-G, K-M from Algeria, Jovet-Ast & Bischler 76083 (PC), D-E, H-I from U. S. A., Texas, Wheeler 1106 (PC). FIG. 7. Distribution of Oxymitra incrassata. FIG. 8. Riccia albida. A-B. Lobes, dorsal side. C. Epidermal cells inlaid with chalk. D-M. Cross sections of lobe, from apex to base; in K chlorophyllose tissue dotted, in K and L violet and pink spots shaded. N. Cross section of dorsal tissue. O-Q. Epidermal cells, in P and Q cleared from chalk. All from Whittemore 1599. FIG. 9. Riccia albopunctata. A. Thallus, dorsal side. B-D. Cross sections of lobe. E. Cross section of lobe margin. F. Scale arrangement.G. Cells and idioblasts. H. Scale cells. I. Cells of dorsal tissue, with thin, longitudinal strip. K. Cells of dorsal tissue, with one persisting epidermal cell. L. Dry lobe, margins incurved over dorsal side. M-P. Cross sections of lobes with idioblasts. All from Gradstein & Weber H218. FIG. 10. Riccia atromarginata. A-C. Lobes with papillae, dorsal side. D-G. Cross sections of lobes with papillae. H. Cells of dorsal epidermis, surface view. I. Epidermal cells in groove. K. Papillae of lobe flanks. L. Papillae of lobe margin. All from Mediterranean specimens. FIG. 11. Distribution of Riccia albida (diamonds), R. albopunctata (dots), and R. atromarginata (stars). FIG. 12. Riccia australis. A-B. Lobes, dorsal side. C-D. Lobe apices, ventral side. E-F. Cross sections of lobes. G. Cross section of dorsal tissue. H. Epidermal cells in groove. I. Scale cells. K. Dry thallus. L. Moist lobe. M-O. Cross sections of lobes. P. Cross section of dorsal tissue. A-I from Vianna 58731, K-P from Fruchart 65. FIG. 13. Distribution of Riccia australis. FIG. 14. A-K. Riccia boliviensis. A-C. Lobes, dorsal side. D. Lobe, ventral side. E-G. Cross sections of lobe. H. Cross section of lobe margin. I. Cross section of dorsal tissue. K. Scale cells, the marginal hyaline. L-S. Riccia brasiliensis. L-M. Lobes, dorsal side. N-O. Cross sections of lobe. P. Arrangement of cilia on lobe margin and flank. Q-R. Cilia. S. Scale cells. A-K from Hermann 24590, L-S from Schiffner 2395. FIG. 15. Riccia breutelii. A. Dry thallus. B-C. Lobes, dorsal side. D-E. Cross sections of lobe. F. Cross section of marginal cells of lobe. G. Cross section of part of lobe, dorsal and ventral tissues

Bischler et al. 235 indistinct. H-K. Cross sections of lobe from apex to base. L. Epidermal cells, dorsal view. A, D-F from Breutel s.n., B-C, G-L from Gaumer 1152. FIG. 16. Distribution of Riccia boliviensis (diamonds), R. brasiliensis (dots), and R. breutelii (stars). FIG. 17. Riccia brittonii. A-D. Lobes, violet flanks dotted. E-F. Cross sections of lobe; in e yellow parts dotted; in F violet spots shaded. G. Cross section of lobe margin, with violet external cells (dotted). H. Cross section of scale. I. Cross section of dorsal tissue, basal walls of epidermal cells thickened. All from Britton et al. 1749a. FIG. 18. Riccia campbelliana. A-B. Lobes, dorsal side. C. Dry lobe apex, the scales extending beyond lobe margins. D-I. Cross sections of lobe, from apex to base. K. Cross section of lobe margin. L. Epidermal idioblasts. M. Epidermal cells near groove. N-O. Dorsal and ventral tissue with idioblasts. P. Scale cells, violet (dotted) or orange (stippled). All from Campbell s.n. FIG. 19. Riccia cubensis. A. Thallus, dorsal side. B. Apex of lobe, ventral side, with transversal bands. C. Lobe apex, dorsal side. D. Lobe margin. E. Lobe apex with cilia. F. Cross section of lobe margin with cilia. G. Epidermal cells in groove. H-M. Cross sections of lobe, from apex to base. N. Cilia of lobe margins. All from Ekman 8348. FIG. 20. Distribution of Riccia brittonii, (diamonds) R. campbelliana (dots), and R. cubensis (stars). FIG. 21. A-K. Riccia ekmanii. A-B. Lobes, dorsal side. C. Lobe margin, tinged with violet. D. Lobe, ventral side. E-G. Cross sections of lobe. H. Cross section of lobe margin. I. Lobe margin seen from ventral side. K. Cross section of dorsal tissue, with longitudinal cell wall thickenings. L-Q. Riccia elliottii. L-M. Lobes, dorsal side. N-O. Cross sections of lobes showing the brown ventral side and the hyaline margin. P. Epidermal cells, dorsal view; the triangles are the openings of the air channels. Q. Epidermal cells in groove. A-K from Ekman 8432, L-Q from Elliott 984. FIG. 22. A-N. Riccia enyae. A. Dry thallus. B-C. Apices of moist lobes. D-F. Cross sections of lobes. G-H. Cross section of lobe margin with cilia. I-K. Cilia on lobe margin. L-M. Cilia. N. Thick walled scale cells. O-W. Riccia howellii. O. Thallus with capsules. P-Q. Lobes, dorsal side. R-V. Cross section of lobes. W. Cross section of lobe margin. A-N from Vianna 3075, O from Gradstein H 489, P-V from Gradstein H 459b, W from Howell 209. FIG. 23. Distribution of Riccia ekmanii (diamonds), R. elliottii (dots), and R .enyae (stars). FIG. 24. Riccia erythrocarpa. A. Dry lobe. B-G. Cross sections of lobes. H. Cross section of lobe margin. I-K. External layer of flank, ending in one or two hyaline cells. L. Cross section of dorsal tissue, with thickened basal wall of epidermal cells. M. Lobe margin seen from ventral side, surface view. N. Hyaline border of flank, surface view. O. Scale cells tinged with violet (outlined). All from Vital 7933.

Bischler et al. 236 FIG. 25. A-G. Riccia fruchartii. A-C. Lobes, dorsal side. D-E. Cross sections of lobe. F. Cross section of dorsal tissue. G. Scale margin and cells. H-l. Riccia grandis. H. Thallus rosette. I. Lobes, dorsal side. K-L. Cross sections of lobe. A-G from Fruchart 718, H, K-L from St. Hilaire s.n., I from Ule 219. FIG. 26. Distribution of Riccia erythrocarpa (diamonds), R. fruchartii (dots), and R. grandis (stars). FIG. 27. A-H. Riccia horrida. A. Rosette forming thallus. B. Branched lobe. C-G. Cross sections of lobe from apex to base. H. Apex of cilium. I-W. Riccia ianthina. I-K. Lobes, dorsal side. L-R. Cross sections of lobe. S. Cells of dorsal tissue with pitted strip. T-V. Cross sections of dorsal tissue. W. Scale cells. A-H from Vital 1869, I-W from Pike 2762. FIG. 28. Distribution of Riccia horrida (diamonds) and R. howellii (dots). FIG. 29. Riccia iodocheila. A-C. Cross sections of flanks. D-E. Margins of flank scales. F. Lobe, dorsal side. G. Apex of lobe, ventral side, with scales on flanks and ventral side. H-M. Cross sections of lobe. N-O. Cross sections of flanks. P-Q. Epidermal cells. R-S. Flank scales. T. Papilla of flank scale margin. U. Ventral scale at lobe apex. A-E from Howell 201, F-M, S-U from Düll 2/230, N-R from Steere 17562. FIG. 30. Distribution of Riccia ianthina (stars) and R. iodocheila (dots). FIG. 31. Riccia lamellosa. A-B. Moist thalli, dorsal side. C. Lobe with capsules. D-G. Cross sections of lobe. H-K. Cross sections of lobe margins and scales. L. Scale cells, pink cells dotted. A-B from Mediterranean specimens, C, H-L from Düll 2/98, D-G from Reveal 2663. FIG. 32. Distribution of Riccia lamellosa; records from U.S.A. not shown. FIG. 33. Riccia lindmanii. A. Thallus, dorsal side. B-E. Cross sections of lobe from apex to centre. F. Dry thallus with capsules. G. Cilia on lobe margin near lobe apex. H-K. Cross sections of lobes near apex. L. Cross section of lobe near base. M-N. Coss sections of central part of lobe near lobe base. O-Q. Cilia. R. Cross section of margin near lobe base. A-E, P-Q from Vital 9126, F-O, R from Lindman 250. FIG. 34. Riccia mauryana. A. Dry lobe. B-C. Cross sections of lobe. D. Lobe border, surface view, showing the hyaline margin and the smooth (stippled) and granulose (dotted) epidermal cells. E. Cross section of lobe margin, with hyaline cell. F. Cross section of dorsal tissue with hyaline, thin strips. G. Lobe, dorsal side. H-I. Cross sections of lobe near apex and in the centre. K. Lobe margin, surface view, with hyaline, smooth and granulose cells. L-M. Epidermal cells with granulose basal wall. N. Granules, surface view. O. Cross section of lobe margin with terminal hyaline cell. A-F from Sleumer 2141, G-O from Maury s.n. FIG. 35. Riccia nigrella. A. Dry thallus. B-C. Moist lobes. D. Lobe with capsules. E-I. Cross sections

Bischler et al. 237 of lobes. K. Epidermal cells in groove. L. Scale cells, the marginal pink, the interior (dotted), blackviolet. All from Mediterranean specimens. FIG. 36. Distribution of Riccia lindmanii (diamonds), R. mauryana (dots), and R. nigrella (stars); records from U.S.A. not shown. FIG. 37. Riccia plano-biconvexa. A. Dry lobe, dorsal side. B-F. Cross sections of lobe. G. Cross section of lobe margin. H. Cells of dorsal tissue with thin, longitudinal strip. All from Lindman 184. FIG. 38. Riccia ridleyi. A-C. Lobes, dorsal side. D. Lobe, ventral side. E-K. Cross sections of lobe from apex to base. L. Cross section of dorsal tissue. M. Cells of dorsal tissue with two thickened, longitudinal strips. N. Cross section of lobe margin. O. Marginal lobe cells, ventral view. P. Margin of lobe, ventral view. All from Ridley s.n. FIG. 39. Riccia sanguineisporis. A-B. Lobes, dorsal side. C-i. Cross sections of lobe. K. Cross section of lobe margin. L. Cross section of dorsal tissue, cell walls thickened. M. Thickened cells of dorsal tissue. N. Epidermal cells in groove. O. Scale cells. P. Part of flank, seen from above, with thickened cells. All from Vital 2335. FIG. 40. Distribution of Riccia plano-biconvexa (dots), R. ridleyi (diamonds), and R. sanguineisporis (stars). FIG. 41. Riccia sorocarpa. A. Thallus rosette. B-C. Lobes, dorsal side. D-G. Cross sections of lobe. H. Cross section of lobe margin, with hyaline marginal cell (partly shaded). I. Thickened basal walls of epidermal cells, horizontal section. K. Dry thallus. L-M. Cross sections of lobe. N. Cross section of dorsal tissue. A-K from Mediterranean specimens, L-O from Chile n° 73. FIG. 42. Riccia squamata. A-C. Lobes, dorsal side. D. Lobe, ventral side. E. Scales and transverse strips of ventral side. F-l. Cross sections of lobes. M. Cross section of dorsal tissue. N. Pitted cell of ventral strip. O. Dorsal tissue in compressed part of lobe. P. Cross section of groove. Q-R. Scale cells. S. Cross section of flank margin and scale. T. cross section of scale. All from Vital 2936. FIG. 43. A-K. Riccia subdepilata. A-B. Lobes, dorsal side, A dry, B moist. C-F. Cross sections of lobe from apex to base. G-H. Cross sections of lobe with capsules and antheridia. I-K. Marginal cilia. L-R. Riccia subplana. L-M. Lobes, dorsal side. N-O. Cross sections of lobe. P. Cross section of lobe margin. Q. Cross section of dorsal tissue, with pitted, longitudinal strips. R. Epidermal cells. A, C-F, I from Vital 8146, B, G-H, K from Vital 8147, L-R from Duss 579. FIG. 44. Distribution of Riccia sorocarpa (dots); records from U.S.A. not shown, R. squamata (diamonds), and R. subdepilata (stars). FIG. 45. Riccia taeniaeformis. A, C-D. Lobes, dorsal side. B. Lobe, ventral side. E-K. Cross sections

Bischler et al. 238 of lobe. L. Cross section of epidermis and dorsal tissue. M. Cross section of lobe margin. N. Scale cells. All from Bueno 4563. FIG. 46. Riccia trichocarpa. A-E. Lobes dorsal side. F. Cross section of lobe. G-H. Cilia. I-K. Setae. L. Cilia and setae above a capsule. All from Mediterranean specimens. FIG. 47. Distribution of Riccia subplana (dots), R. taeniaeformis (diamonds), and R. trichocarpa (stars); records from U.S.A. not shown. FIG. 48. Riccia viannae. A-C. Dry and moist thalli. D-F. Cross sections of lobe. G. Cross section of epidermis, with hyaline and chlorophyllose (dotted) dorsal tissue. H. Epidermis and hypodermal cells. I. Epidermal cells in groove. K-L. Cross sections of lobe margins with one or three hyaline marginal cells. M. Scale cells. N. Cells of scale margin. All from Vianna 1199. FIG. 49. Riccia violacea var. violacea. A-C. Cross sections of lobes. D. Granulose hyaline papillae. E-G. Lobes, dorsal side. H. Lobe, ventral side. I-K. Scales. L-M. Cross sections of lobe. N, S. Cross sections of flank with papillose cilia. O-R, T-U. Granulose papillae. V. Scale cells. A-D from Britton et al. 1749b, E-Q from Dietrich 1556, R-V from Dietrich 1554. FIG. 50. Riccia violacea var. laevis. A-D. Cross sections of lobe. E-H. Cross sections of flanks with papillae and scales. I-K. Papillae. A-D from Le Gallo 510, E-K from Le Gallo 1278. FIG. 51. Distribution of Riccia viannae (dots), R. violacea var. violacea (diamonds), and R. violacea var. laevis (stars). FIG. 52. Riccia vitalii. A-B. Dry lobes, dorsal side. C. Moist lobe, dorsal side. D. Lobe showing the scales, lateral side. E. Lobe bordered with scales. F. Cross section of lobe margin. G. Dorsal tissue, horizontal section. H. Cross section of dorsal tissue with longitudinal thickenings. I. Scale margin. K. Epidermal cells. L-N. Cross sections of lobe. All from Vital 5941. FIG. 53. .Distribution of Riccia vitalii. FIG. 54. Riccia weinionis. A-B. Thalli, dorsal side. C. Lobe, ventral side. D-E. Lobe margin, dorsal side. F. Lobe margin, ventral side. G-H. Cross sections of lobe. I. Scale cells, the marginal narrow and hyaline. K. Marginal cells of lobe, ventral side. L. Cross section of lobe margin. M. Dorsal epidermis, surface view. N. Cross section of dorsal tissue, with thickened longitudinal strips. O. Horizontal section of dorsal tissue. P. Epidermal cells in groove. A from Le Gallo 1274, B-P from Vital 3031. FIG. 55. Distribution of Riccia weinionis. FIG. 56. Riccia membranacea. A-C. Thalli, dorsal side. D. Epidermal pores. E. Cross section of lobe through capsule. F. Cross section of lobe with two layers of air chambers. G. Cross section of lobe

Bischler et al. 239 with single layer of air chambers. H. Cross section of lobe margin. I. Epidermal pore in older part of thallus. K-L. Dorsal epidermis and air chambers, surface view. A-C, I, L from Melendez-Howell s.n., D from different specimens, E, G from Vital, 1865, F, H from Jovet-Ast s.n, K from Bueno 6388. FIG. 57. Distribution of Riccia membranacea; records from U..S.A. not shown. FIG. 58. Riccia curtisii. A-B. Female thalli. C. Male thallus. D-F. Pores. G. Cross section of lobe near apex. H. Cross section of lobe margin. I, L. Cross sections of lobe in the middle. K. Cross section of air chambers. M-O. Regenerations from ventral side of lobe. A, B, G H from Vianna 4378, I, K, L from Vianna 4378, B, D, E, F from Vianna 4578, M, N, O from Breil s.n. FIG. 59. Distribution of Riccia curtisii; records from U.S.A. not shown. FIG. 60. A-F. Riccia bahiensis. A. Lobe, dorsal side. B. Cross section of lobe. C-E. Epidermal pores. F. Part of cross section of lobe with air chambers (dotted). G-N. R. Chiapasensis. G. Dry thallus. HK. Cross sections of lobes. L. Cross section of dorsal tissue. M. Epidermal pore. N. Cross section of epidermal pore. A-F from Martius s.n., G-N from Riba s.n. FIG. 61. Riccia crassifrons. A-B. Lobes, dorsal side. C. Cross section of lobe margin. D. Cross section of lobe. E-H. Epidermal pores. I. Lobes, dorsal side. K. Cross section of lobe with two layers of air chambers. L. Cross section of lobe margin. M. Cross section of lobe and capsule. N. Compact tissue beneath capsule. O-P. Epidermal pores. A-G from type of R. spruceana, Spruce s.n., H-P from type of R. crassifrons, Spruce s.n. FIG. 62. Distribution of Riccia bahiensis (dots), R. chiapasensis (diamonds), and R. crassifrons (stars). FIG. 63. Riccia dussiana. A-E. Thalli, dorsal side. F. Thallus, ventral side. G-l. Cross sections of lobes. M-N. Parts of cross sections of lobes with pores. O-P. Epidermal pores, surface view. Q. Cells with pitted strips in dorsal tissue. R. Section of scale. S. Margin at lobe apex. A, M from Duss 577, B-C, R, Q from Stehlé 4022, D-L, N-P, S from Le Gallo 3. FIG. 64. Riccia frostii. A-C. Female thalli. D. Male thalli. E. Cross section of lobe. A from Bachmann 10008, B-D from Mediterranean specimens. FIG. 65. Riccia geissleriana. A-C. Dry thallus and lobes. D. Lobe margin with scales. E-H. Cross sections of lobes. I. Cross section of dorsal tissue. K. Cross section of lobe margin. L. Dorsal epidermis, surface view. All from Geissler 8885. FIG. 66. Distribution of Riccia dussiana (dots), R. frostii (diamonds), and R. geissleriana (stars). FIG. 67. Riccia hegewaldiana. A-C. Thalli, dorsal side. D-F. Cross section of lobes. G. Cross section

Bischler et al. 240 of lobe margin. H. Cross section of dorsal tissue. I-K. Epidermal pores. L. Cross section of epidermal pore. All from Hegewald 5050. FIG. 68. Riccia jovet-astiae. A. Thallus, dorsal side. B-C. Lobe apices. D-F. Cross sections of lobes. G-H. Cross sections of lobe margins. I. Cross section of dorsal tissue and epidermal pore. K. Cells of dorsal epidermis, surface view. L. Cross section of part of epidermal pore. M-P. Epidermal pores, surface view. All from Vianna 665. FIG. 69. A-N. Riccia limicola. A-B. Lobes, dorsal side. C. Lobe with capsule on ventral side. D. Branching pattern of thalli and capsule position. E-G. Cross sections of lobes. H. Part of cross section of lobe showing the air chambers (dotted). I. Epidermal pore in old part of lobe. K-L. Epidermal pores in young part of lobe. M-N. Epidermal pores. O-R. R. Paraguayensis. O. Female thallus. P. Male lobe. Q. Lobe, dorsal side, and part of epidermal pore. R. Cross section of lobe, with single layer of air chambers. A-F, H-N from Weber B-41244, G from Weber 41238, O-R from Balansa 1280. FIG. 70. Distribution of Riccia hegewaldiana (dots), R. jovet-astiae (diamonds), and R. limicola (★). FIG. 71. Distribution of Riccia paraguayensis. FIG. 72. Riccia paranaensis. A-C. Thalli, dorsal side. D-L. Cross sections of lobes. M-O. Cross sections of dorsal tissue. P. Cross section of part of epidermal pore. Q-R. Epidermal pores, surface view. A-B, Q-R from Schäfer-Verwimp 8453, C-K from Schäfer-Verwimp 14992, L-N from Vital 9251, O-P from Vital 8957. FIG. 73. Distribution of Riccia paranaensis. FIG. 74. Riccia stenophylla. A-C. Thalli, dorsal side. D-F. Cross sections of lobes near apex. G. Cross section of lobe margin near base. H. Cross section of scale. I-K. Part of cross section of lobe near apex. L. Part of cross section of lobe near base. M-Q. Epidermal pores. A-B, D-I, M-Q from Balansa 3706, C, K, L from Hosseus 531. FIG. 75. Distribution of Riccia stenophylla. FIG. 76. A-G. Riccia cavernosa. A-C. Thalli with capsules. D-E. Cross sections of lobes. F-G. Epidermal pores. H-P. Riccia crystallina. H-M. Thalli, dorsal side. N. Cross section in narrow part of lobe. O. Cross section in wide part of lobe. P. Cross section of lobe margin. A-E from Mediterranean specimens, F, G from Hegewald 7387, H-P from Mediterranean specimens FIG. 77. Distribution of Riccia cavernosa (dots); records from U.S.A. not shown, and R. crystallina (stars). FIG. 78. Spores. A, M. Proximal faces, B-L. Distal faces. A. Riccia atromarginata. B. R. boliviensis.

Bischler et al. 241 C. R. brasiliensis. D. R. breutelii. E. R. brittonii. F. R. campbelliana. G. R. ekmanii. H. R. elliottii. I. R. enyae. K. R. erythrocarpa. L. R. fruchartii. M. R. grandis. A from a Mediterranean specimen, B from type, C from type, D from Hegewald 5449, E from type, F from a Californian specimen, G from Ekman s.n., H from type, I from type, K from type, L from Vital 9124, M from Vital 9250 (scale bars A-G, I-M = 10 µm, H=15 µm). FIG. 79. Spores. A-C, E-M. Distal faces. D. Proximal face. A. Riccia howellii. B. R. ianthina. C. R. lamellosa. D. R. lindmanii. E. R. mauryana. F. R. nigrella. G. R. plano-biconvexa. H. R. ridleyi. I. R. sanguineisporis. K. R. sorocarpa. L. R. squamata. M. R. subdepilata. A from Gradstein H 489, B from type, C from a Louisiana specimen, D from type, E from type, F from a Mediterranean specimen, G from type, H from Vital 8314, I from type, K from Hegewald 7258, L from type, M from type (scale bars = 10 µm). FIG. 80. Spores. A-G, L. Distal faces. H-I. Spore tetrads. K. Group of spores. M. Distal face and lateral view. A. Riccia subplana. B. R. taeniaeformis. C. R. trichocarpa. D. R. viannae. E. R. vitalii. F. R. violacea var. laevis. G. R. weinionis. H-I. R. curtisii. K. R. membranacea. L. R. bahiensis. M. R. chiapasensis. A from Melendez s.n., B from type, C from a Mediterranean specimen, D from type, E from Vital 6520, F from Le Gallo 1278, G from Vital 2875, H from Vital 9235, I from Vianna 4178, K from type of R. amazonica, L from type, M from type (scale bars A-I, L = 10 µm, K = 15 µm, M = 20 µm). FIG. 81. Spores. A, C-H, K. Distal faces. B. Distal and proximal faces. I, L-M. Groups of spores. A. Riccia crassifrons. B. R. dussiana. C. R. frostii. D. R. geissleriana. E. R. hegewaldiana. F. R. jovetastiae. G. R. limicola. H. R. paraguayensis. I. R. paranaensis. K. R. stenophylla. L. R. cavernosa. M. R. crystallina. A from type, B from Cremers 3642, C from Backmann 10008, D from Geissler 8885, E from type, F from Vianna 665, G from Weber B-41244, H from Vital 9251, I from Vital 9251, K from Balansa 3706, L from Hegewald 6522, M from France, Pierrot 1992(PC) (scale bars A, C-H, K = 10 µm, B = 15 µm, I, M = 20 µm, L = 30 µm). FIG. 82. Ricciocarpos natans. A. Thallus with archegonial groove. B. Thallus with antheridial groove. C. Cross section of thallus. D. Arrangement of air chambers. E. Epidermal pore, surface view. F. Cross section of epidermal pores and air chambers. G. Arrangement of scales on ventral side of thallus. H. Scale of floating form. I. Scales of terrestrial form. K. Apex of scale of terrestrial form. A-G, I from St. Hilaire s.n., H from Griffin IIl et al. 819. FIG. 83. Distribution of Ricciocarpos natans; records from U.S.A. not shown. FIG. 84. Corsinia coriandrina. A. Thallus male and female receptacles. B. Arrangement of scales on ventral side of thallus. C-D. Epidermal pores, surface view. E. Epidermal pores and air chambers,

Bischler et al. 242 cross section. F-G. Scales. H. Thallus with sporophyte enclosed in the calyptra, cross section. I. Involucre. K. Cross section of calyptra. L. Capsule wall. All from Morocco, Malençon s.n. (PC). FIG. 85. Distribution of Corsinia coriandrina. FIG. 86. Cronisia fimbriata. A. Thallus with involucres. B. Thallus with male groove. C. Arrangement of scales on ventral side of thallus. D. Cross section of thallus at level of an archegonium. E. Cross section of thallus at level of an antheridium. F. Epidermal pore, surface view. G. Cross section of epidermis and air chambers. H-I. Scales. K. Appendages of scales. L-M. Involucres and sporophytes. N. Involucral tissue with oil cells. O. Capsule wall. P. Spore mother cell and sterile cells. A-C, E-F, I, L, N-P from Vital 2924, D, G-H, K, M from Vital 8153. FIG. 87. Distribution of Cronisia fimbriata. FIG. 88. Cronisia weddellii. A. Thallus with involucres and antheridial grooves. B. Cross section of thallus. C. Epidermal pore, surface view. D. Cross section of epidermis and air chambers. E-F. Scales. G. Papilla on scale margin. H-I. Involucres and sporophytes. K. Involucral tissue with idioblasts. L. Capsule wall. M. Elaters. A, C-D, F-H, M from Vital 1877, B, E, I-L from Bischler 1671. FIG. 89. Distribution of Cronisia weddellii. FIG. 90. Cyathodium bischlerianum. A. Ventral view of thallus with sporophyte (sp) within involucre (in) and antheridial receptacles. B. Marginal cells of thallus. C. Dorsal epidermal cells with chloroplasts (ch) and oil cells (oc). D. Branched rhizoid. E. Pores. F. Ventral scales. G. Male receptacle. H. Scales of antheridial receptacles. I. Archegonium. J. Upper capsule cells. K. Seta. L. Elater. A, G, I from Salazar et al. 16612, B, D, J from Salazar et al. 16600, 14386, C from Salazar et al. 16600, D from Salazar et al. 14386, E from Salazar et al. 14392, 16600, F from Salazar et al. 16612, 14392, 14384, H from Salazar et al. 16613, K from Salazar et al. 16643, L from Rodríguez 216. FIG. 91. A-D. Cyathodium bischlerianum. A. Ventral epidermal cells. B. Upper marginal cells of involucre. C. Upper cells of involucre. D. Elaters. E-J. Cyathodium spruceanum. E. Inner involucral scales. F. Marginal cells of involucre near ventral insertion. G. Inner cells of involucre. H. Archegonium. I. Elater and (on right) detail of spiral bands. J. Branched elater. A from Salazar et al. 14386, B from Rodríguez 209, C from Salazar et al. 1439, D from Rodríguez 2162, E from Salazar et al.16660, F, I from Salazar et al. 16674, G from Salazar et al. 16610, H from Salazar et al. 16614, J from Salazar et al. 16655. FIG. 92. Distribution of Cyathodium bischlerianum.

Bischler et al. 243 FIG. 93. Cyathodium cavernarum. A. Dorsal view of thallus with sporophytes (sp) in involucre and anteridial receptacles. B. Marginal cells of thallus. C. Ventral view of upper epidermal cells. D. Rhizoids. E. Archegonia. F. Dorsal epidermal cells with chloroplasts (ch) and oil cells (oc). G. Ventral epidermal cells. H. Outer cells of operculum. I. Inner cells of operculum. J. Marginal cells of involucre. K. Inner cells of involucre near margin. L. Upper capsule cells. M. Elater. A, B, F, G, K from Salazar et al. 16547, C from Salazar et al. 15234, D from Salazar et al. 16116, 15236, E from Salazar et al. 14386, H from Salazar et al. 19109, 16572, 16547, 16110, I from Salazar et al. 16108, J from Salazar et al. 16572, L-M from Rodríguez 220. FIG. 94. Distribution of Cyathodium cavernarum (dots) and C. foetidissimum (stars). FIG. 95. Cyathodium foetidissimum. A. Adult plant, dorsal view. B. Ventral view of fertile plant. C. Ventral view of male receptacle and archegonia. D. Detail of archegonia in involucre. E. Upper epidermal cells (ch = chloroplasts, oc = oil cell). F. Dorsal pore. G. Marginal cells. H. Ventral epidermal cells. I. Part of transverse section of thallus. J. Ventral scales. K. Rhizoid. L. Upper capsule cells (ventral view). M. Upper capsule cells (dorsal view). N. Lower capsule cells. O. Elater. P. Spores. A-B, L-M from Salazar et al. 17048, C-H, J-K from Salazar, Lépiz & De Gracia 14747, I, N-O from Salazar, Lépiz & De Gracia 17049. FIG. 96. Cyathodium spruceanum. A. Female plant with involucres (in). B. Involucres. Upper one bearing 20-30 archegonia, lower one 5-10. C. Male plants with antheridial receptacles (o = receptacles, sc = scale). D. Ventral scales of thallus. E. Ventral scales of involucre, one with a rhizoid (ri). F. Marginal cells of thallus. G. Part of transverse section of thallus. H. Upper epidermal cells with chloroplasts (ch) and oil bodies (ob). I. Pores. J. Ventral scale of tuber. K. Cells of operculum. A from Salazar et al. 16610, B from Salazar et al. 16614, 14165, C from Salazar et al. 16080, D from Salazar et al. 16387, E from Salazar et al. 16648, 16655, 16610, F, from Salazar et al. 16614, G from Salazar et al. 16655, H from Salazar et al. 16633, I from Salazar et al. 15028, 16572, J from Rivera s.n., K from Salazar et al. 16129. FIG. 97. A-B. Cyathodium bischlerianum. A. Lateral view of spore with long acuminate spines and a cingulum. B. Antheridial receptacle with scales (arrows). C-D. Cyathodium cavernarum. C. Dorsal pore. D. Spore, distal face. E. Cyathodium foetidissimum, spore, distal face. F-H. Cyathodium spruceanum. F. Antheridial receptacle. G. Ventral scales at apex of female thallus. H. Two spores of the tetrad, note asymmetrical position of cingulum and distal face with lamellate-reticulate ornamentation. A from Salazar et al. 16612, B from Salazar et al. 16599, C from Salazar et al. 16184, D from Salazar et al. 16692, E from Salazar et al. 17048, F from Salazar et al. 16581, G from Salazar et al. 16549, H from Salazar et al. 16650. FIG. 98. Cyathodium steerei. A. Adult plant, dorsal view. B. Young plant, dorsal view. C. Ventral view of female plant with involucre. D. Ventral scales. E. Detail of pore. F. Ventral scale of tuber.

Bischler et al. 244 G. Marginal cells. H. Elater. I. Spore. J. Upper epidermal cells. K. Lower epidermal cells. A-B, C, D (two on the right side), E-F & H reproduced with permission from Hässel de Menéndez (1963), D (on the left side), G, I, J, K from type specimen. FIG. 99. Distribution of Cyathodium spruceanum (dots) and C. steerei (stars). FIG. 100. Targionia hypophylla. A. Thallus with apical involucre. B. Thallus with apical involucre, lateral view. C. Thallus with antheridial branches. D. Arrangement of scales on ventral thallus side. E. Cross section of thallus. F. Epidermal pore, surface view. G. Epidermal pore and air chambers, cross section (some chlorophyllose filaments only are drawn). H. Scales. I. Appendage of scale. K. Capsule wall. A-E, H-K from Geissler 8732, F-G from Volk 6/0698. FIG. 101. Targionia stellaris. A. Thallus, dorsal side. B. Thallus with young apical involucre, lateral view. C. Thallus with antheridia. D. Cross section of thallus. E. Epidermal cells. F. Epidermal pores, surface view. G. Epidermal pore, cross section. H. Epidermal pore and air chambers, cross section (some chlorophyllose filaments only are drawn). I. Scale. K. Appendage of scale. All from Sleumer 3596. FIG. 102. Distribution of Targionia hypophylla ( dots); records from U.S.A. not shown, and T. stellaris (diamonds). FIG. 103. Exormotheca pustulosa. A. Thallus with archegoniophore and male receptacle. B. Female receptacle with sporophytes. C. Arrangement of scales on ventral side of thallus. D. Cross section of thallus. E-F. Epidermal pores, surface view. G. Cross section of epidermis and air chambers. H-I. Scales. K. Appendage of scale. L. Cross section of archegoniophore stalk. M. Epidermal pore of female receptacle. N. Cross section of epidermis and air chambers of female receptacle. O. Capsule wall. A, C-H, L, O from Düll A 76, B, I-K, M-N from Spain, Bischler & Boisselier 1 (PC). FIG. 104. Spores. A, D, F, H, K-L. Distal faces. B, E, G, I, M. Proximal faces. C, group of spores. A-B. Oxymitra incrassata. C. Ricciocarpos natans. D-E. Corsinia coriandrina. F-G. Cronisia fimbriata. H-I. Cronisia weddellii. K. Targionia hypophylla. L-M. Exormotheca pustulosa. A-B from Algeria, Baudoin s.n. (PC), C from Japan, Savatier s.n. (PC), D-E from Vital 8153, F-G, from Bischler 1671, H-I from Tunisia, Jovet-Ast & Bischler 795 (PC), K from Israel, Jovet-Ast & Bischler 87 (PC), L-M from Düll 1966/2 (scale bars =10 µm). FIG. 105. Athalamia andina. A. Thallus with young female receptacle and archegoniophore. B. Female receptacle, ventral side. C. Thallus with male receptacle. D. Arrangement of scales on ventral side of thallus. E. Cross section of thallus. F. Epidermal pores, surface view. G. Cross section of epidermis and air chambers. H. Scales. I. Appendages of scales. K. Cross section of archegoniophore stalk. L. Capsule wall. A from Spruce s.n., B-L from Volk 6/0670.

Bischler et al. 245 FIG. 106. Athalamia pygmaea. A. Thallus with archegoniophore. B. Female receptacle with sporophytes. C. Thallus with male receptacle. D. Arrangement of scales on ventral side of thallus. E. Cross section of thallus. F. Epidermal pores, surface view. G. Cross section of epidermis and air chambers. H. Epidermal cells. I. Scale. K. Appendage of scale. L. Scale of female receptacle. M. Cross section of archegoniophore stalk. N. Capsule wall. A-B, L-M from Whittemore 2469, C-K from Whittemore 4096. FIG. 107. Distribution of Athalamia andina ( dots) and A. pygmaea (stars). FIG. 108. Sauteria chilensis. A. Thallus with archegoniophore and antheridial receptacle. B. Arrangement of scales on ventral side of thallus. C. Cross section of thallus. D. Epidermal pores, surface view. E. Cross section of epidermal pore and air chambers. F-G. Scales. H. Scale of female receptacle. I. Cross section of archegoniophore stalk. K. Female receptacle, ventral view. L. Epidermal pore of female receptacle, surface view. M. Open capsule in involucre. N. Capsule wall. All from Gay s.n. FIG. 109. Distribution of Exormotheca pustulosa ( dots) and Sauteria chilensis ( diamonds). FIG. 110. Distribution of Asterella dominicensis. FIG. 111. Distribution of Asterella venosa. FIG. 112. Distribution of Asterella macropoda (dots) and A. elegans (diamonds). FIG. 113. Distribution of Asterella echinella. FIG. 114. Distribution of Asterella lateralis. FIG. 115. Distribution of Asterella versicolor. FIG. 116. Asterella, A. dominicensis, A. venosa, A. macropoda, A. echinella, spores FIG. 117. Asterella, A. elegans, A. lateralis, A. pringlei, A. versicolor, spores FIG. 118. Asterella, A. rugosa, A. alpina, spores FIG. 119. Distribution of Asterella pringlei. FIG. 120. Distribution of Asterella rugosa (dots) and A. alpina (diamonds). FIG. 121. Cryptomitrium tenerum. A. Thallus with archegoniophore and anthridial receptacle. B. Thallus with female receptacle. C. Arrangement of scales on ventral side of thallus. D. Cross section of thallus. E. Epidermal pore, surface view. F. Cross section of epidermal pore and air chambers. G-

Bischler et al. 246 H. Scales. I. Cross section of archegoniophore stalk. K. Female receptacle, dorsal side. L. Female receptacle, ventral side. M. Epidermal pore of female receptacle, surface view. N. Epidermal pore of female receptacle, cross section. O. Margin of involucre. P. Sporophyte with open capsule in involucre and lid of capsule. Q. Capsule wall. All from Dusen 149. FIG. 122. Mannia californica. A. Thallus with young female receptacle and archegoniophore. B. Thallus with antheridial receptacle. C. Arrangement of scales on ventral side of thallus. D. Cross section of thallus. E. Epidermal pore, surface view. F. Cross section of epidermal pore and air chambers. G. Scales. H. Appendage of scale. I. Cross section of archegoniophore stalk. K. Female receptacle, ventral side. L. Epidermal pore of female receptacle, surface view. M. Epidermal pore of female receptacle, cross section. N. Sporophyte with open capsule in involucre. O. Capsule wall. All from U. S. A., Arizona, Frahm s.n. (PC). FIG. 123. Distribution of Cryptomitrium tenerum ( dots), Mannia californica ( stars) and Mannia hegewaldii (diamonds). FIG. 124. Mannia hegewaldii. A. Thallus with archegoniophore and antheridial receptacle. B. Cross section of thallus. C. Epidermal pore, surface view. D. Cross section of epidermal pore and air chambers. E. Scales. F. Appendages of scale. G. Cross section of archegoniophore stalk. H. Female receptacle, dorsal side. I. Female receptacle, ventral side. K. Scale of female receptacle. L. Epidermal pore of female receptacle, surface view. M. Epidermal pore of female receptacle, cross section. N. Capsule wall. A-B, D-E, H-I from Hegewald 5138, C, F-G, K-N, from Hegewald & Hegewald 5286. FIG. 125. General characteristics of the genus Plagiochasma. A. Thallus with dorsal androecium, young female receptacle, and archegoniophore. B. Thallus with successive adventitious branches, with androecia. C. Arrangement of scales on ventral side of thallus. D. Cross section of thallus. E. Cross section of epidermis, epidermal pores and assimilatory layer. F. Antheridium and paraphyses. G-H. Mature female receptacles. I. Epidermal pore of female receptacle, surface view. K. Epidermal pore of female receptacle, cross section. L. Cross section of archegoniophore stalk. M. Urn of open capsule. N. Lid of capsule. O. Capsule wall. From different specimens. FIG. 126. A-E. Plagiochasma argentinicum. A-B. Epidermal pores, surface view. C. Epidermal pore, cross section. D. Scales. E. Appendages of scales. F. Apices of appendages. G-N. Plagiochasma crenulatum. G-H. Epidermal pores, surface view. I. Epidermal pore, cross section. K. Scales. L. Appendages of scales. M. Apex of appendage. N. Margin of appendage. A-C from Volk 6/669, D-F from Hosseus 163, G-H, K from Barnes & Land 659, I, L-N from Conzatti & Gonzalez 1130ï. FIG. 127. Distribution of Plagiochasma argentinicum (stars) and P. crenulatum (dots).

Bischler et al. 247 FIG. 128. A-G. Plagiochasma cuneatum. A-B. Epidermal pores, surface view. C. Epidermal pore, cross section. D. Scales. E. Appendages of scales. F. Apices of appendages. G. Margin of appendage. H-N. Plagiochasma intermedium. H. Epidermal pore, surface view. I. Epidermal pore, cross section. K. Scales. L. Appendages of scales. M. Apex of appendage. N. Margin of appendage. A-B, D-E from Volk 6/717, C, G from Deppe & Schiede s.n., F from Mc Allister s.n., H, K-L from Heyde & Lux 6292, I from Standinger s.n., M-N from Liebmann 532b. FIG. 129. A-G. Plagiochasma jamaicense. A. Epidermal pore, surface view. B. Epidermal pore, cross section. C. Scales. D. Appendages of scales. E. Margin of appendage. F-G. Apices of appendages. H-P. Plagiochasma landii. H-I. Epidermal pores, surface view. K. Epidermal pore, cross section. L. Scales. M. Appendages of scales. N-O. Appendices of appendages. P. Margin of appendage. A-B, D-G from Underwood 1173, C from Pringle 10682, H-K, N from Amable 1652, L-M, O-P from Barnes & Land 466. FIG. 130. Distribution of Plagiochasma cuneatum (dots), P. intermedium (diamonds) and P. jamaicense (stars). FIG. 131. A-G. Plagiochasma muenchianum. A. Epidermal pore, surface view. B. Epidermal pore, cross section. C. Scales. D. Appendages of scales. E-F. Apices of appendages. G. Margin of appendage.H-O. Plagiochasma wrightii. H-I. Epidermal pores, surface view. K. Epidermal pore, cross section. L. Scales. M. Appendages of scales. N. Apices of appendages. O. Margin of appendage. A-G from Münch 5607, H-O from Wright 9. FIG. 132. Distribution of Plagiochasma münchianum. (stars) and P. wrightii (dots). FIG. 133. A-E. Plagiochasma rupestre var. rupestre. A. Epidermal pore, surface view. B. Epidermal pore, cross section. C. Scales. D. Appendages of scales. E. Apex of appendage. F-L. Plagiochasma rupestre var. Volkii. F. Epidermal pore, surface view. G. Epidermal pore, cross section. H. Scales. I. Appendages of scales. K. Margin of appendage. L. Apex of appendage. A, C, E from Bertero 1129, B, D from France, Bischler & Baudoin 75239 (PC), F-L from Goodspeed 33011. FIG. 134. Distribution of Plagiochasma landii (dots) and P. rupestre var. volkii (diamonds). FIG. 135. Distribution of Plagiochasma rupestre var. rupestre. FIG. 136. A-K. Spores. A-B, D-E, G-H. Distal faces. C, lateral view. F, I-K. Proximal faces. A. Athalamia andina. B. Athalamia pygmaea. C. Sauteria chilensis. D. Cryptomitrium tenerum. E-F. Plagiochasma argentinicum, G. Plagiochasma wrightii, H-I. Plagiochasma intermedium. K. Plagiochasma landii. L-M. elaters. L. Plagiochasma intermedium. M. Plagiochasma crenulatum. A from Spruce s.n., B from Whittemore 2469, C from Gay s.n., D from California, Blasdale s.n. (PC), E-F fromVolk 6/669, G from Nicolas 5612, H-I from Standinger s.n., K from Amable 1952, L from

Bischler et al. 248 Barnes & Land 640, M from Volk 6/718 (scale bars A-E, G-K, M = 10 µm, F = 20 µm, L = 5 µm). FIG. 137. Reboulia hemisphaerica. A. Thallus with young female receptacle, archegoniophore and antheridial receptacles. B. Arrangement of scales on ventral side of thallus. C. Cross section of thallus. D. Epidermal pores, surface view. E. Cross section of epidermal pore and air chambers. F. Scales. G. Appendage of scale. H. Scale of antheridial receptacle. I. Scale of female receptacle. K. Cross section of archegoniophore stalk. L. Female receptacle, ventral side. M. Epidermal pore, surface view. N. Epidermal pore of female receptacle, cross section. O. Sporophyte with open capsule in involucre. P. Capsule wall. All from Herter 94773. FIG. 138. Distribution of Reboulia hemisphaerica; records from U.S.A. not shown. FIG. 139. Lunularia cruciata. A. Thallus with archegoniophore. B. Thallus with antheridial receptacle and gemma cup. C. Thallus with young female receptacle. D. Arrangement of scales on ventral side of thallus. E. Cross section of thallus. F. Epidermal pore, surface view. G. Cross section of epidermal pore and air chambers (some chlorophyllose filaments only are drawn). H. Scale. I. Appendage of scale. K. Gemma. L. Scales of female receptacle. M. Cross section of archegoniophore stalk. N. Female receptacle. O. Capsule wall. All from Gay s.n. FIG. 140. Distribution of Lunularia cruciata. FIG. 141. Dumortiera hirsuta. A. Thallus with archegoniophore and antheridiophore. B. Young archegoniophore. C. Arrangement of bristles on ventral side of thallus. D. Epidermis, surface view. E. Epidermis with papilliform cells, cross section. F. Cross section of part of thallus. G. Scales. H. Scales of female receptacle. I. Apex of scale of female receptacle. K. Cross section of archegoniophore stalk. L. Female receptacle, dorsal side. M. Female receptacle, ventral side. N. Capsule wall. O. Elater and spores. All from Ule 128. FIG. 142. Distribution of Dumortiera hirsuta; records from U.S.A. not shown. FIG. 143. Marchantia berteroana. A. Thallus with archegoniophore and thallus with antheridiophores. B. Arrangement of scales on ventral side of thallus. C-D. thallus margin. E. Epidermal pore, outer and inner openings. F. Epidermal pore, cross section (some chlorophyllose filaments only are drawn). G. Median scale. H. Appendage of median scale. I. Laminal scale. K. Marginal scale. L. Lobe of cupule margin. M. Female receptacle. N. Cross section of archegoniophore stalk. O. Margin of involucre. P. Male receptacle. Q. Cross section of antheridiophore stalk. A-B, P from New Zealand, Campbell s.n. (PC), C-D, O from Mayor 57, E-G, I-K, Q from Troll 2087, H, N from Bertero s.n., L from Castellanos 586, M from Perez Moreau 4602. FIG. 144. Distribution of Marchantia berteroana.

Bischler et al. 249 FIG. 145. Marchantia plicata. A. Thallus with archegoniophore and thallus with antheridiophore. B. Arrangement of scales on ventral side of thallus. C. Thallus margin. D. Epidermal pore, outer and inner openings. E. Epidermal pore, cross section (some chlorophyllose filaments only are drawn). F. Median scale. G. Appendage of median scale. H. Laminal scales. I. Marginal scale. K. Lobe of cupule margin. L. Female receptacle. M. Cross section of archegoniophore stalk. N. Margin of involucre. O. Male receptacle. P. Cross section of antheridiophore stalk. A-C, E from d'Orbigny 209, D, G from Spruce s.n., I, L from Benoist 2501, F, H, K, M-P from Bischler 378. FIG. 146. Distribution of Marchantia plicata. FIG. 147. Marchantia polymorpha. A. Thallus with archegoniophores and thallus with antheridiophores. B. Arrangement of scales on ventral side of thallus. C. Thallus margin. D. Epidermal pore, outer and inner openings. E. Epidermal pore, cross section (some chlorophyllose filaments only are drawn). F. Median scale. G. Appendage of median scale. H. Laminal scale. I. Marginal scale. K. Lobe of cupule margin. L. Female receptacle. M. Cross section of archegoniophore stalk. N. Lobe of margin of involucre. O. Cross section of female ray with papillae. P. Male receptacle. Q. Cross section of antheridiophore stalk. A-B, K, M, O from France, Bischler s.n. (PC), C from Espinosa 4020, D, L, Q from France, Hébrabrd s.n. (PC), E-I from U. S. A., Roell s.n. (PC), N from Killip & Smith 15548, P from Castellanos 1314. FIG. 148. Distribution of Marchantia polymorpha. FIG. 149. Marchantia breviloba. A. Thallus with archegoniophore and thallus with antheridiophore. B. Arrangement of scales on ventral side of thallus. C. Mucilage cavity, sclerotic cells and oil cells in basal thallus tissue. D. Epidermal pore, outer and inner openings. E. Epidermal pore, cross section (some chlorophyllose filaments only are drawn). F. Median scales. G. Appendage of median scale. H. Laminal scales. I. Cupule margin. K. Female receptacle. L. Cross section of archegoniophore stalk. M. Margin of involucre. N. Male receptacle. O. Cross section of antheridiophore stalk. A-F, H, K from Evans 175, G, N from Underwood 2234, I from Benoist 3125, L-M from Underwood 2251, O from Bischler 1713. FIG. 150. Distribution of Marchantia breviloba. FIG. 151. Marchantia chenopoda. A. Thallus with archegoniophore. B. Thallus with antheridiophore. C. Epidermal pore, outer opening. D. Inner openings of epidermal pores. E. Epidermal pore, cross section (some chlorophyllose filaments only are drawn). F. Median scales. G. Appendages of median scales. H. Laminal scales. I. Cupule margin. K. Female receptacle. L. Cross section of archegoniophore stalk. M. Margin of involucre. N. Male receptacle. O. Cross section of antheridiophore stalk. P. Cross section of male ray. A-B, H-I, P from Baudoin s.n., C, K, L from Guilielmus s.n., D from d'Orbigny 226, E from Jovet-Ast s.n., F, N, from Harling 3427, G, O from

Bischler et al. 250 Moritz 47b, M from Glaziou 6348. FIG. 152. Distribution of Marchantia chenopoda. FIG. 153. Marchantia inflexa. A. Thallus with archegoniophore. B. Thallus with antheridiophore. C. Epidermal pore, outer opening. D. Epidermal pore, cross section (some chlorophyllose filaments only are drawn). E. Inner openings of epidermal pores. F. Median scales. G. Appendage of median scale. H. Laminal scales. I. Cupule margin. K. Female receptacle. L. Cross section of archegoniophore stalk. M. Margins of involucres. N. Male receptacle. O. Cross section of antheridiophore stalk. P. Cross section of male ray. A-H, L, N-P from Baudoin s.n., I from U. S. A., Sullivant exs. 128 (NY), K, M from Martinique, lectotype of M. inflexa. FIG. 154. Distribution of Marchantia inflexa. FIG. 155. Marchantia paleacea. A. Thallus with archegoniophore. B. Thallus with antheridiophore. C. Arrangement of scales on ventral side of thallus. D. Epidermal pore, outer opening. E. Inner openings of epidermal pores. F. Epidermal pore, cross section (some chlorophyllose filaments only are drawn). G. Median scales. H. Appendage of median scale. I. Laminal scale. K. Lobe of cupule margin. L-M. Female receptacles. N. Cross section of archegoniophore stalk. O. Margin of involucre. P. Male receptacle. Q. Cross section of antheridiophore stalk. A-C, K, O from Jovet-Ast s.n., D, L-M from type of M. domingensis, E, H-I from type of M. tholophora, F from Italy, Raddi s.n. (PC), G from type of M. squamosa, N from France, Dismier s.n. (PC), P from Den Held & Van Rijn HH 41, Q from India, Gollan 1302 (PC). FIG. 156. Marchantia papillata. A. Thallus with archegoniophore. B. Thallus with antheridiophore. C. Epidermal pore, outer opening. D. Inner openings of epidermal pores. E. Epidermal pore, cross section (some chlorophyllose filaments only are drawn). F. Median scales. G. Appendage of median scale. H. Laminal scales. I. Cupule margins. K. Female receptacle. L. Cross section of archegoniophore stalk. M. Margin of involucre. N. Scale of female receptacle. O. Male receptacle. P. Cross section of antheridiophore stalk. A-B, D, F, H, O-P from Spruce s.n., C from d'Orbigny 104, E, K, M from type of M. papillata, G, I from Gaudichaud s.n., L from Balansa 4006, N from Schiffner 2349. FIG. 157. Distribution of Marchantia paleacea (dots) and M. papillata (stars). FIG. 158. A-E. Monoclea gottschei subsp. elongata. A. Habit with androecia, dorsal view. B. Habit with gynoecium and sporophyte, dorsal view. C. Elater and spores. D. Portion of elater and spores. E. Habit with gynoecium and sporophyte with opened capsule. F. Portion of cross sction through thallus. G-j. Sphaerocarpos muccilloi. G. Habit with androecia, dorsal view. H. Habit with gynoecia, dorsal view. I. Tetrad of spores. J. Thallus margin. K-R. Asterella venosa. K. Cross

Bischler et al. 251 section of thallus. L. Schema of female receptacle in longitudinal section (a = archegoniophore, b = calyptra, c = capsule, i = involucre, p = pseudoperianth). M. Thallus pore. N. Thallus with scales in two rows, ventral view. O. Cross section of thallus middle. P. Thallus pore, cross sction. Q. Female receptacle (i = involucre, p = pseudoperianth). R. Habit with groups of androecia at base of archegoniophores, dorsal view (reproduced with permission from Gradstein et al., 2001). FIG. 159. Distribution of Monoclea gottschei subsp. elongata. FIG. 160. Spores and reproductive structures. Spores. A. Mannia hegewaldii, lateral view. B. Reboulia hemisphaerica, proximal and distal faces. C. Lunularia cruciata, distal face. D. Dumortiera hirsuta, distal face. E. Marchantia berteroana, distal face and lateral view. F. Marchantia chenopoda, distal face. G. Marchantia paleacea, distal face. H. Marchantia papillata, distal face. I. Monoclea gottschei, distal face. K. Part of elater of Marchantia inflexa. L. Capsule wall thickenings of Marchantia paleacea. M. Papillae on rays of female receptacle of Marchantia polymorpha. A from Hegewald 5138, B from Israel, Jovet-Ast & Bischler 118 (PC), C from Israel, Herrnstadt s.n. (PC), D from Borneo, Menzel et al. 4235 (PC), E from St. Paul, Segonzac s.n. (PC), F from Bischler 365, G from Wright s.n., H from Spruce s.n., I from Frahm et al. 1690, K from Baudoin s.n., L from Frye 2852, M from Marivault s.n. (scale bars A-B, E, K-L = 10 µm, C, I = 1 µm, D, F-H, M = 5 µm).

Bischler et al. 252

(Author CV’s)

HELENE BISCHLER-CAUSSE

Helene Bischler-Causse was born in Switzerland in 1932. She attended the University of Genève where she obtained a Ph.D. in natural sciences in 1957. She has held research positions at the Universidad Nacional, Bogotá (Colombia), during 1956-1959, at the Conservatoire Botanique, Genève, during 1960-61, and at the Centre National de la Recherche Scientifique (C.N.R.S.), Paris, from 1961 onwards where she was appointed to several research teams (liverworts of the Neotropics, liverworts of the Mediterranean area, evolution of cryptogams, biodiversity, molecular systematics). Dr. Bischler’s research interest was first systematics of neotropical liverworts, then ecology of Mediterranean liverworts, then systematics and evolution of the liverwort subclass Marchantiidae. She did extensive field work in Colombia and in the Mediterranean area, and contributed to many floras. She introduced many research students from all over the world to liverwort studies and was the editor of the scientific periodical Cryptogamie, Bryologie-Lichénologie for more than 30 years. She is a co-author of several volumes of Index Hepaticarum and published more than 120 scientific research papers and several books, including taxonomic monographs of the general Marchantia and Plagiochasma, and a summary on systematics and evolution of the genera of the Marchantiales. For her research, Dr. Bischler was awarded the prize Plantamour-Prévost of the Université de Genève in 1953, the P.Bertrand prize of the Académie des Sciences, Paris in 1974, the Geneva Sayre award of the Harvard University in 1985, and the Hattori Prize of the International Association of Bryologists in 1995.

Bischler et al. 253

STEPHAN ROBBERT GRADSTEIN

Stephan Robbert Gradstein was born in The Netherlands in 1943. He attended the University of Utrecht, where he obtained a Ph.D. in botany in 1975. He has held positions as a botany lecturer and curator of cryptogams at Utrecht university during 1969-1995, as a graduate research assistant at the University of Cincinnati, U.S.A. during 1970-71, as a Fullbright fellow at the University of Colorado in 1976, and as a professor in tropical botany at the university of Michigan during 1986-87. In November 1995 followed his appointment as professor in botany and director of the Herbarium and Botanical Garden of the University of Göttingen. Dr. Gradstein's research interest is the flora and vegetation of the tropics, especially of tropical America. His research speciality are liverworts. He had done extensive fieldwork in all parts of the Neotropics, has taught several field courses in tropical bryology, has directed numerous research projects, and has published more than 200 research papers and several books, including the "Guide to the Bryophytes of Tropical America" (2001) coauthored by S. P. Churchill (Missouri Botanical Garden) and N. Salazar Allen (Smithsonian Tropical Research Institute). Dr. Gradstein is an editor of Flora Neotropica and a Deputy-Director of the Organization for Flora Neotropica (UNESCO), and a founding member and president of the International Association of Bryologists (IAB). He is a member of the Academy of Sciences of Göttingen. For his botanical research Dr. Gradstein was awarded the Candolle Prize of the Societé de Physique et d'Histoire Naturelle de Genève in 1994.

Bischler et al. 254

SUZANNE JOVET-AST

Suzanne Jovet-Ast was born in Paris in 1914. She attended the Faculty of Sciences of Paris where she obtained a Ph.D. in 1943 with the mention “très honorable”. She held the positions of assistant (1945-1946), assistant director (1957-1975), then director of the Laboratoire de Cryptogamie, Muséum National d’Histoire Naturelle (1975-1982). Dr. Jovet’s research interest focuses principally on liverworts. She did extensive field work in the Mediterranean area and tropical America. She directed many botanical and ecological research projects, dealing especially with semiarid and arid regions in the Mediterranean, and published over 150 papers on liverwort systematics and taxonomy. She has worked for many years on a worldwide taxonomic revision of the large and difficult genus Riccia. Dr. Jovet was the editor of Revue Bryologique and Lichénologique for many years. She was a member of nearly all bryological societies over the world and took part in many international meetings. For her research she was awarded the Coincy Price of the Botanical Society of France in 1948 and the Desmazière Prize of the Académie des Sciences, Paris, in 1964.

Bischler et al. 255

DAVID GEOFFREY LONG

David Geoffrey Long was born in 1948 in Scotland and graduated in Botany at Edinburgh University in 1966. He obtained his Ph.D. at Trinity College, Dublin in 2001. Since 1975 he has been employed as a botanist at the Royal Botanic Garden Edinburgh where he has held a variety of posts. From 1975 until 1995 he was co-author and editor of Flora of Bhutan, and from 1990 to 1995 head of the Himalayan Botany Section. From 1988 to 1990 served as Curator of the Edinburgh Herbarium. From 1995 onwards his work has been exclusively on bryophytes, especially those of the Sino-Himalayan region, and on taxonomy and evolution of Marchantiales, with a monographic interest in Asterella. Since 1997 he has held the position as Head of Cryptogamic Botany at the Royal Botanic Garden, Edinburgh. During these years Dr. Long has undertaken extensive botanical fieldwork overseas, especially in the Himalayas and China, but also in Europe, Macaronesia, the Arctic, the Middle East, tropical Africa, Central and South America. Collections have been both of herbarium material of flowering plants and bryophytes, and more recently of material for molecular phylogenetic investigation, particularly of liverworts. In addition to co-authoring several volumes of the Flora of Bhutan, he has published over 170 botanical papers, latterly mostly on bryological topics, including taxonomy, floristics and phylogeny. He teaches and supervises postgraduate students at the University of Edinburgh where he is an Honorary Fellow, and leads several research projects on bryophytes. He is also active in promotion of bryology through the British Bryological Society, of which he was President during 2002 and 2003, and is involved in bryophyte recording and conservation initiatives in Scotland. He serves on the editorial boards of several Journals including Edinburgh Journal of Botany and Journal of Bryology.

Bischler et al. 256

NORIS SALAZAR ALLEN

Noris Salazar Allen was born in the Republic of Panama in 1947. She attended the University of Panama and Trinity college (B.A., 1969), the State University of New York (S.U.N.Y.), College at Geneseo (M.Sc., 1973) and the University of Alberta, Canada (Ph.D. in bryology, 1986). She was awarded the Benito Juárez (University of Michigan, Institute of Modern Languages, 1967) and LASPAU Scholarships (Trinity College and S.U.N.Y.), a Dissertation Fellowship (University of Alberta, 1984) and a Postdoctoral Fellowship from the Smithsonian Tropical Research Institute (1989-1990). She has done research at the Missouri Botanical Garden, Smithsonian Institution, New York Botanical Garden, National Museum of Natural History (Laboratoire de Cryptogamie) in Paris, and in The Netherlands. She has been on the reaching staff of the University of Panama and has been the Curator of the Cryptogamic Herbarium since 1969. She holds a joint appointment with the University of Panama and the Smithsonian Tropical Research Institute (Panama, 1990). Dr. Salazar Allen’s main research interests are the biosystematics, floristics and biogeography of tropical rain forest bryophytes and lichens. She has done field work in the Neotropics, Vanuatu and Fiji and has published many research papers. She is co-author with S. R. Gradstein and S. P. Churchill of the “Guide to the Bryophytes of Tropical America.” She has organized and taught in various bryological workshops in Central America and the Caribbean. She has served as main editor of Briolatina, the newsletter of the Sociedad Latinoamericana de Briología (1992-1998), she is in the directorship of the Red de Herbarios de Mesoamérica y el Caribe and is a member of the Commission for Flora Neotropica.