(Marchantiophyta) of Colombia

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ranges of the Andes and in isolated hills of Sierra Nevada de Santa Marta and .... SPECIMENS EXAMINED: COLOMBIA: Boyacá: Páramos al NW de Belén, ...
Nova Hedwigia Vol. 94 issue 1–2, 97–127 Stuttgart, February 2012

Article

Taxonomic revision of Balantiopsaceae (Marchantiophyta) of Colombia Laura Victoria Campos-Salazar* and Jaime Uribe-M. Instituto de Ciencias Naturales, Facultad de Ciencias, Universidad Nacional de Colombia, apartado 7495 Bogotá, D.C. Colombia. With 12 figures

Campos-Salazar, L.V. & J. Uribe-M. (2012): Taxonomic revision of Balantiopsaceae (Marchantiophyta) of Colombia. – Nova Hedwigia 94: 97–127. Abstract: This paper presents the taxonomic treatment of Balantiopsaceae for Colombia. In Colombia, the family is represented by three genera: Isotachis, Neesioscyphus and Ruizanthus. A re-evaluation of the morpho-anatomical characters used to delimit species within these genera is presented. Seven species are recognized of which four belong to Isotachis, two to Neesioscyphus and one to Ruizanthus. Two species, I. lindigiana and I. mascula are treated as synonyms of I. serrulata, and I. tenax is synonymous to I. multiceps. Keys for the three genera and seven species, descriptions, illustrations and distribution maps are provided. Key words: Balantiopsaceae, Colombia, Isotachis, Marchantiophyta, Neesioscyphus, Ruizanthus.

Introduction Colombia harbors 60% of the known liverwort species of the Neotropical region. The richness of the Colombian liverwort flora (840 species in 136 genera) is in contrast with the scarcity of detailed taxonomic, morphological and phylogenetic knowledge of Colombian liverworts (Uribe & Gradstein 1998). As part of our ongoing studies of the Colombian bryoflora we have focused on the study of the leafy liverwort family Balantiopsaceae. A discussion of the morpho-anatomical characters traditionally used to delimit genera is presented along with descriptions of all Colombian species of the Balantiopsaceae.

*Corrresponding author; e-mail: [email protected] © 2012 J. Cramer in Gebr. Borntraeger Verlagsbuchhandlung, Stuttgart, Germany www.borntraeger-cramer.de DOI: 10.1127/0029-5035/2012/0094-0097 0029-5035/2012/0094-0097 $ 7.75

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TAXONOMIC HISTORY. The Balantiopsaceae include seven or eight genera: Anisotachis R.M.Schust., Acroscyphella N.Kitag. & Grolle, Balantiopsis Mitt., Eoisotachis R.M.Schust., Hypoisotachis (R.M.Schust.) J.J.Engel & G.L.Merril, Isotachis Mitt., Neesioscyphus Grolle and Ruizanthus R.M.Schust. (Crandall-Stotler et al. 2009a, b; Frey & Stech 2009). The family is currently divided into three subfamilies: Isotachidoideae, Balantiopsidoideae and Ruizanthoideae; the monophyly of these infra-familiar divisions has been supported by several molecular studies (Hentschel et al. 2006; He-Nygren 2007 and Masuzaki et al. 2010). The Balantiopsaceae are mainly distributed in the temperate zones of the Southern Hemisphere (South America, New Zealand and New Caledonia). In the Neotropics, four genera and thirteen species have been reported (Gradstein et al. 2001). The monotypic genus Ruizanthus is distributed in Colombia, Costa Rica and Venezuela. Isotachis (7 species) is widely distributed in Tropical America, Neesioscyphus (5 species) in Brazil, Colombia, Costa Rica, Bolivia, Peru and Ecuador, and Balantiopsis (2 species) in southern South America (Gradstein et al. 2001). In Colombia, representatives of Isotachis, Neesioscyphus and Ruizanthus have been reported. Isotachis was established by Mitten in 1855 with two species from New Zealand, I. lyallii Mitt. and I. subtrifida (Hook. & Taylor) Mitt. The latter species was later transferred to the genus Triandrophyllum Fulford & Hatcher (Herbertaceae) (Fulford & Hatcher 1958). A more complete treatment of the genus was published by Gottsche (1863) in his work on the liverworts of Mexico. Gottsche (1864) transferred in his treatment of the liverworts for the "Prodromus Florae-Novogranatensis" several species of Jungermannia L. to Isotachis. In addition, he described two new species based on specimens collected by Lindig in the vicinity of Bogotá: I. lindigiana Gottsche and I. mascula Gottsche. The type specimen of the latter species was destroyed in B. by the fire in the Second World War. Stephani (1909) treated Isotachis on a world-wide basis and included 49 species, 21 for tropical America. The genus was placed in the family Ptilidiaceae by Evans (1939). Schuster (1953) proposed the family Isotachidaceae and later (Schuster 1972) placed Isotachis along with the segregate genus Eoisotachis R.M.Schust. and Neesioscyphus Grolle in the subfamily Isotachidoideae. The concept of the family was expanded by Hatcher (1960, 1961). Currently, there is no taxonomic treatment of the genus Isotachis for Colombia. Hatcher (1960, 1961) undertook a monographic study of all Latin American species of the genus recognizing five species for Colombia: I. haematodes (Lehm. & Lindenb.) Gottsche, I. lindigiana Gottsche, I. madida (Hook.f. & Taylor) Mitt., I. multiceps (Lindenb. & Gottsche) Gottsche and I. serrulata (Sw.) Gottsche, and dealt with the great variability and proposed species boundaries with emphasis on the leaf and underleaf characters. Fulford (1963) followed the treatment of Hatcher. Schuster (1985) re-evaluated the species concepts of Hatcher and recognized three genera in Hatcher’s treatments. Gradstein et al. (1977) acknowledged that I. haematodes and I. madida are conspecific with I. serrulata. Gradstein & Hekking (1979) reported eight species for Colombia, adding I. lacustris Herzog, I. mascula Gottsche

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and I. tenax Steph. In Uribe & Gradstein (1998), six species are reported for Colombia: I. lacustris , I. lopezii (R.M.Schust.) Gradst., I. lindigiana, I. multiceps, I. serrulata and I. tenax. Grolle (1964) established the genus Neesioscyphus with three species: N. carneus (Nees) Grolle and N. argillaceus (Nees) Grolle from the Neotropics, and N. phoenicorhizus Grolle from New Zealand. The latter species was transferred to Acroscyphella N.Kitag. & Grolle (N.Kitagawa & Grolle 1985). Grolle (1966) added three neotropical species to Neesioscyphus: N. homophyllus (Nees) Grolle (Brazil), N. bicuspidatus (Steph.) Grolle, and N. allionii (Steph.) Grolle, and placed (Grolle 1972) the genus in Balantiopsaceae. Neesioscyphus is mostly neotropical, with one species, N. homophyllus, reported for the north-east of Argentina (ReinerDrehwald & Drehwald 1995). Isotachis and Neesioscyphus are also reported from Ecuador: Isotachis with six species, and Neesioscyphus with three species (León-Yáñez et al. 2006); from Bolivia: Isotachis with five species, and Neesioscyphus with two species (Churchill et al. 2009). The genus Ruizanthus was proposed by Schuster (1984), and based on two species: R. venezuelanus R.M.Schust. and R. lopezii R.M.Schust. from the Sierra Nevada de Mérida in Venezuela. Gradstein transferred R. lopezii to Isotachis, as I. lopezii, because of the bifid leaves and twisted capsule valves (Gradstein et al. 2001). Materials and methods About 500 specimens of Isotachis, Neesioscyphus and Ruizanthus were examined, including type specimens and fresh-collected material. Dissecting and compound light microscopes were used to scrutinize morphological and anatomical characters suggested in previous taxonomic treatments for the family (Hodgson 1949; Hatcher 1960, 1961; Fulford 1963; Schuster & Engel 1997; Engel & Merril 1997). Leaf length was measured from the basal insertion on the stem to the apex, leaf width at the widest point perpendicular to the length and cell size according to the lumen. Cell length was measured as the longest distance of the cell and the width was measured perpendicularly to the length. All measurements represent minimum and maximum values. Illustrations were done using a drawing tube. The terminology used in this treatment follows Magill (1990), Malcolm & Malcolm (2000), Gradstein et al. (2001) and Paton (1999). The morphological species concept used in this study refers to groups of individuals that present a continuous spectrum of phenotypic variation and that differ from other groups by a combination of at least two exclusive characters. For example: quadrifid leaves and bisbifid underleaves in Ruizanthus venezuelanus R.M.Schust. and the presence of a cuspidate leaf apex, presence of conspicuous trigones and bordered leaves, as in I. lopezii. Characters discussed are those traditionally employed in the taxonomy of the family for separation of genera and species as used by Hatcher 1960, 1961. These characters are derived mainly from the gametophyte, and include the shape of leaves and underleaves, degree of dentition, cell shape and size, and coloration of plants. In more recent studies of New Zealand species, some authors have remarked on the variability of gametophytic characters according to gradients of rainfall and light intensity (Schuster & Engel 1997) and a suite of new sporophytic characters has been proposed by Schuster & Engel (1997) and Engel & Merril (1997). The absence of sporophytes in a large percentage of specimens makes the use of these characters difficult in routine taxonomic identifications.

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Results and discussion Morphological and anatomical characters HABIT: Species are procumbent to suberect to erect in growth and occur in tufts or turfs. In Colombia, Isotachis multiceps has a procumbent habit, although sometimes it can also have an ascending habit as suggested by earlier authors (Hatcher 1961). PIGMENTATION: Generally, the species possess a reddish tint, with different intensities, to the point that this character has been used by some authors to differentiate species. For example, Hatcher (1960, 1961) separated species groups of Isotachis, because they are dark brown instead of red (e.g., I. obtusiloba and I. lacustris). In Colombian specimens, it has been observed that color range varies from green-reddish, to intense red, to brown, to dark-brown, both within and between species. Grolle (1966) separated Neesioscyphus argillaceus from other species of the genus by the color of the plants. However, field observations show that color intensity in these plants is due to the light intensity in the places where they grow. Plants growing in more exposed habitats have a more intense red color, whereas those growing in shadier places are more greenish. Because of the variation displayed and because it is caused by environmental conditions, this character will not be considered in the taxonomic treatment. STEMS: In cross section stems are very variable. A differentiation between cortex and medulla is usually found (Figs 1J, 2J, 3S, 4N, 5L, 6P, 7K, 8K, 9H). The cortex has one to three strata of isodiametric cells with thick brown-colored walls. Cells of the medulla are usually larger, with thin and colorless walls. Generally, the stem cuticle in the family may be smooth or papillose. BRANCHING: In Balantiopsaceae there are branches of the ventral-intercalary type and occasionally terminal branches of the Frullania type. Although Hatcher (1960, 1961) established that in Isotachis there are only branches of the ventral-intercalary type (Hatcher 1961). Schuster & Engel (1997) reported the presence of Frullania type terminal branches. In Colombian specimens only the ventral-intercalary type was found. LEAVES: Leaf shape is a very variable but a useful character. In Isotachis, the majority of species have ovate leaves that vary in the proportion of length to width. In Neesioscyphus leaves are ovate to almost orbicular or subquadrate (Figs 7D, 8C–D). In Ruizanthus the leaves are subquadrate to rotund (Fig. 9C). In some species of Isotachis, leaves are strongly asymmetric (I. serrulata, Figs 5C–D, 6B–F), but in others they are symmetrical (I. multiceps, Fig. 3G–K). In Neesioscyphus there are asymmetric leaves in which the dorsal segment is larger than the ventral or vice versa (Figs 7D–E, 8C–D). Isotachis and Ruizanthus have strongly concave leaves, whereas Neesioscyphus has flat leaves. Lobing. The degree of leaf lobing is a useful character but difficult to delimit. All species of the family (except Ruizanthus venezuelanus) have bifid leaves. In Isotachis, leaves are divided from more than 1/3 up to 1/2 of the leaf length (Figs 1C–G, 3G, 4D). The only species with a rather stable degree of the sinus depth is I. lacustris, in

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which leaves are bifid up to 1/6 their length (Fig. 1C–F). In the other species of Isotachis there is broad variation, even in leaves of the same plant. In Neesioscyphus, leaves are only slightly bifid, to 1/5 of the leaf length, and in the specimens examined there was not much variation (Fig. 8C–D). Ruizanthus represents the exception within the family, because their leaves are quadrifid (Fig. 9C), with long-acuminate to piliferous segments. Leaf segments vary from rotund or obtuse to long- acuminate, including acute. LEAF INSERTION: Two types of leaf insertion have been described in the family, transverse and oblique; the latter can be incubous or succubous. Hatcher (1960, 1961) described Isotachis with an incubous insertion. Gradstein et al. (2001) noted Isotachis as having leaves with a transverse insertion, but sometimes slightly succubous or incubous. Colombian Isotachis all possess a slightly incubous insertion, and when all leaves are removed from the plant, a transverse insertion line is clearly observed. Isotachis multiceps is the exception in this genus since the insertion is slightly succubous. In Neesioscyphus the leaf insertion is clearly succubous, being a diagnostic character within the group. The presence of teeth on the leaves is no discrete character, but appears in a complex range of variation. Branches with clearly dentate leaves can be found along with branches with entire leaves in the same specimen (Fig. 2C–D). Isotachis serrulata is the only Colombian species of the family with strongly dentate to serrulate leaf margins but this character varies greatly within and among individuals (Figs 5C–D, 6B–F). Teeth are small and reduced in number in I. lopezii and I. multiceps (Figs 2C–D, 3I, 4C). In Neesioscyphus and Ruizanthus, toothed leaf margins are rare and are restricted to underleaves and gynoecial bracts and bracteoles (Figs 7F–H, 9D–F). Leaf cells present discrete variation in size, shape, wall thickness, and trigones between the cells of the upper and median leaf lamina. Median cells are rectangular, large and clearly thick-walled as in Isotachis multiceps and I. serrulata (Figs 3Q–R, 4L, 5K, 6Q), or more or less thin-walled as in the other species of Isotachis and the other genera of the family (Figs 1L, 7J, 8M, 9I). Upper cells are generally isodiametric, with incrassate walls and trigones (e.g., Ruizanthus venezuelanus and I. serrulata). In I. lopezii the presence of a margin of elongated cells is an important character to define this species (Fig. 2K). Cuticle ornamentation is a species specific character within the family. Some species have a smooth cuticle, e.g., I. lacustris. A strongly papillose cuticle, almost verruculose, is found in Ruizanthus venezuelanus. Species of Neesioscyphus have clearly a striate-papillose cuticle, a character shared with Isotachis lopezii and I. serrulata. UNDERLEAVES: Size, shape, degree of teething and lobing are important taxonomic characters. For example, species of Isotachis have underleaves similar to the leaves in shape and dentition, but are slightly smaller. Underleaves in Neesioscyphus and Ruizanthus are smaller than in Isotachis, and are only 1/3 the leaf length. Isotachis lopezii has underleaves armed with strong, long, tooth-like spines, but the leaf armature, however, is reduced in size and number (Fig. 2G–I). In Neesioscyphus the underleaves appear armed with a few small teeth, not exceeding two cells in length

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(Figs 7F, 8E–G). In Ruizanthus, on the other hand, the underleaves are strongly armed with long and piliferous teeth, up to 5 cells long, and are also unique in the family in being clearly bisbifid (Fig. 9D–E). In the majority of species of the family, the insertion line of underleaves is slightly curved to straight, with the exception of I. lacustris and I. serrulata having them deeply curved, to the point of forming a cordate or short decurrent base, as in I. lacustris (Fig. 1H–I). As in leaves, segments of the underleaves vary between rotund or obtuse to longacuminate, to acute. In Isotachis, there is variation in shape, even in specimens of the same species. Ruizanthus is unique within the family, showing stability in size and shape (Fig. 9D–E). In Isotachis, the degree of lobing is a highly variable character within species, ranging from more than 1/2 to less than 1/6 the length of the underleaf, with the exception of I. lacustris, in which the sinus is consistently 1/6 the length of the underleaf. In Neesioscyphus and Ruizanthus, on the other hand, lobing is invariably at or close to 1/2 the underleaf length. Cell size, shape, cell wall and cuticle of the underleaves are the same as those of leaves. It is important to emphasize the presence of marginal papillae in underleaves of N. argillaceus (Fig. 8H), a diagnostic character in this species, since the other Colombian species, N. allionii, lacks papillae. RHIZOIDS: In Balantiopsaceae, the presence of rhizoids is an important character associated with the growth form. Prostrate to suberect plants of Isotachis multiceps (Figs 3A, 4A), I. lopezii (Fig. 2A–H), Neesioscyphus argillaceus (Fig. 8A), N. allionii and Ruizanthus venezuelanus (Fig. 9D) have numerous rhizoids. The rhizoids grow in tufts at the base of the underleaves, and are distributed throughout the plant. They are generally colorless with the exception of Neesioscyphus. There they are slightly red-pigmented. ANDROECIA: The absence of sex organs in most collections precludes the use of antheridial features as a taxonomic character at the species level. In the few specimens studied of Isotachis multiceps and Neesioscyphus argillaceus, one and two to three antheridia per bract were found, respectively. Schuster (1985) implemented a key for the family, based on gynoecial and androecial characters, and he mentioned the presence of 4–5 antheridia per bract in Ruizanthus, 2 in Neesioscyphus and in three species of Isotachis. In the present study these observations are confirmed. GYNOECIA: Gynoecia are formed by specialized leaves and underleaves, bracts and bracteoles respectively. In Balantiopsaceae the bracts generally conserve the shape, dentition, degree of lobing and cellular structure of leaves and underleaves, but are larger (Fig. 3L–N). Schuster (1985) described the Isotachis-type gynoecia as terminal on the main axis (Fig. 5B), or on innovations. One known exception is found in I. multiceps where gynoecia are terminal on short ventral-intercalary branches (Fig. 3A). Ruizanthus and Neesioscyphus have gynoecia terminal on the main axes. PERIANTH-PERIGYNIUM COMPLEX: In the present work the term "perianth-perigynium complex" is used to define the protective structure of the sporophyte (Figs 2B, 3A, 5B, 7C, 8B, 9A). In the great majority of leafy liverworts a perianth is formed by gynoecial bracts and therefore it is unistratose. A perigynium is formed mainly of 102

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gametophyte stem tissue and it is pluristratose. In Balantiopsaceae, the proportion of perigynium to perianth varies. In Isotachis the perigynium covers 90% and the perianth is restricted to a laciniate apical portion, and, in such cases, the term TRUE PERIGYNIUm is adopted. In Neesioscyphus and Ruizanthus, the perigynium is 50% or less of the structure, in which case the term TRUE PERIANTH is adopted. A special case appears in I. serrulata, where there are vestigial, scale-like structures on the perigynium, originating from the perianth. Generally, for the family, the perigyniumperianth complex has a conical-cylindrical shape (Figs 2B, 3A, 5B, 7C, 8B). Taxonomic treatment Balantiopsaceae H.Buch, Thüring. Bot. Ges. 1(2–3): 23. 1955 TYPE: Balantiopsis Mitt., in Hooker f., Handb. New Zealand Fl. 751. 1867.

Plants small, medium to large, erect to prostrate on soil or rock or aquatic; strongly anisophyllous, pigmented, from green to intense red, to brown and yellow depending on the species. Ramification generally ventral-intercalary. Stems with cortical cells clearly differentiated from the medullar ones by size and color. Rhizoids at base of underleaves, colorless to reddish or absent. Leaves transverse, succubous to slightly incubous, flattened to concave, bifid or quadrifid, margin entire to strongly toothed. Median cells are rectangular, the upper leaf cells are slightly smaller. Trigones present or absent. Cuticle smooth to striate-papillose. Underleaves smaller than leaves, but generally conserving their shape and ornamentation. Androecia with 1–5 antheridia per bract. Gynoecia terminal on short or long branches. Perianth-perigynium complex dominated by the perianth or the perigynium. Capsule generally with spirally twisted valves. Key to the Colombian genera of Balantiopsaceae 1

Leaves with longitudinal line of insertion, clearly succubous, flat. Gynoecia with a perianthperigynium complex dominated by perianth (true perianth)...................................... 2 Neesioscyphus 1' Leaves with transverse insertion, slightly incubous, strongly concave. Gynoecia with a perianthperigynium complex dominated by the perianth or perigynium.......................................................... 2

2

Leaves quadrifid with long-acuminate segments. Perianth-perigynium complex dominated by the perianth (true perianth).................................................................................................. 3 Ruizanthus 2' Leaves bifid with acute to rotund segments. Perianth-perigynium complex dominated by the perigynium (true perigynium).................................................................................... 1 Isotachis

1 Isotachis Mitt. Isotachis Mitt., in Hooker, J.D., Bot. Antarct. Voy. II. 2: 148. 1855 TYPE: Isotachis lyallii Mitt.

PLANTS small to large, up to 9 cm long, 0.5–2 mm wide, erect or procumbent, strongly anisophyllous to subisophyllous (I. lacustris), brown to reddish. STEMS dark brown; in cross section a clear difference between the cells: the cortical ones (small 103

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and dark brown colored), in 2–3 rows, the medullar ones larger and colorless. RHIZOIDS colorless, in tufts or absent. LEAVES strongly asymmetric, subquadrate to ovate, concave, channeled, imbricate to approximate; bifid to 1/2 to 1/4, segments triangular, with cuspidate to acute tips, sinus acute to rotund; margins entire, dentate to serrulate; insertion transverse to slightly incubous, succubous in I. multiceps. LEAF CELLS isodiametric to clearly rectangular. Trigones absent to conspicuous; walls thickened to thin; cuticle striate-papillose to smooth. UNDERLEAVES smaller than leaves, subquadrate to ovate, imbricate to proximal, extended to reflexed; sinus rotund to acute; margins entire, dentate to serrulate; bases rotund, line of insertion arcuate. ANDROECIA diandrous, with exception of I. multiceps, which is monandrous. GYNOECIA terminal. BRACTS AND BRACTEOLES larger than leaves and underleaves, but similar in shape, concave, narrowing towards the base, bifid; segments widely triangular, acute, to cuspidate. PERIANTH-PERIGYNIUM COMPLEX multistratose in 90% of its length, rigid, erect, with a single cell layer only at the apex (true perigynium). CAPSULE ovoid with spirally twisted or straight valves. DISTRIBUTION: Southern Hemisphere, higher elevations in the Tropics; Asia to China. In Colombia, Isotachis is represented by four species, distributed in the three mountain ranges of the Andes and in isolated hills of Sierra Nevada de Santa Marta and Serranía de la Macarena. Key to the species of Isotachis in Colombia 1 Plants very small up to 1.0 cm long, suberect to procumbent. Rhizoids present................................. 2 1' Plants medium to large, 3–9 cm long, erect. Rhizoids absent............................................................. 3 2

Leaves bordered by elongate cells; leaf segments cuspidate by 2–3 cells, rarely with only one. Leaf cells mainly isodiametric; trigones bulging. Androecia diandrous. Gynoecia terminal on foliose main axes. Microphyllous stolons absent. Underleaves with numerous long-acuminate teeth, some bisbifid................................................................................................................................ 2 I. lopezii 2' Leaves non-bordered; leaf segments non-cuspidate, acute to obtuse. Leaf cells mainly rectangular to elongate, trigones inconspicuous to cordate. Androecia monandrous. Gynoecia terminal on short ventral-intercalary branches. Microphyllous stolons present................................ 3 I. multiceps 3

Leaves bifid to 1/6 of their length, entire, symmetrical; apex of the segments rounded. Median cells of leaves (–3)5 to 10 times longer than wide. Plants associated with bodies of water... 1 I. lacustris 3' Leaves bifid 1/4 to 1/2 of their length, strongly toothed to entire, asymmetric; apex of the segments acute to obtuse. Median cells of the leaves up to 3 times longer than wide. Plants growing on ground or cliffs.............................................................................................................. 4 I. serrulata

1 Isotachis lacustris Herzog, Hedwigia 74: 94. 1934.

Figs 1, 10

TYPE: Bolivia, Illampu peak, monte Sorata, cordillera Real, Troll 56 (holotype, JE!).

PLANTS large, erect, up to 8 cm long, 3–4 mm wide, including leaves¸ subisophylous, yellowish to dark brown, ramified or not. STEMS brown dark; in cross section 14–18 cells in diameter; the cortical cells smaller and dark brown, 20–27 × 12–25 µm, distributed in two rows, the medullar cells larger and colorless, 42–60 × 17–35 µm. RHIZOIDS absent. LEAVES strongly symmetrical to weakly asymmetric, ovate to orbicular, 2.3–3.0 × 1.6–2.5 mm, proximal, bifid up to 1/6 or less of their length, 104

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segments triangular, with apices obtuse to round, sinus obtuse, margins entire, base rounded; insertion line of ventral portion transverse, insertion line of dorsal portion longitudinal. LEAF CELLS near the apex slightly isodiametric to subquadrate, with uniformly incrassate walls, of 42–50 × 17–40 µm, the median cells elongate to rectangular, 68–145 × 15–17 µm, walls thin, trigones absent; the basal cells are equal to the median; cuticle smooth. UNDERLEAVES rounded to ovate, contiguous, symme-trical 1.4–1.8 × 1.0–1.7 mm, bifid up to 1/6 of their length; segments triangular; apex rotund to obtuse; sinus obtuse; margins entire; line of insertion curved. ANDROECIA and GYNOECIA not seen. HABITAT AND DISTRIBUTION: Tropical America: Bolivia and Perú, in Colombia the distribution of this species is restricted to the Eastern Cordillera (Departamentos of Meta, Cundinamarca and Boyacá), with a single record in the Central Cordillera in Nevado del Tolima. Its altitudinal distribution is mainly high-Andean, from 3400 to 3950 m. Plants generally submerged aquatics. DIAGNOSTIC FEATURES: Isotachis lacustris can be clearly differentiated from other species of the genus even though it is similar to I. serrulata (see below for diagnosis of I. serrulata ). The diagnostic characters are: leaves with median cells up to 10 times longer than wide, with thin walls and trigones absent. Underleaves bifid to up to 1/6 of their length, segments distant from one another. Aquatic habitat. SPECIMENS EXAMINED: COLOMBIA: Boyacá: Municipio de Duitama, Páramo de La Rusia, Laguna Negra, 3745 m, 14 Dec. 1972, A.M.Cleef 7225 (COL, GOET, U); Sierra Nevada del Cocuy, Alto Valle de Lagunillas, 3950 m, 24 Sep. 1972, P.A.Florschütz, 4000, 5936 (GOET, U, COL); Municipio de Belén, Páramos al NW de Belén, Quebrada Minas, Laguna El Alcohol, 3785 m, 26 Mar. 1972, A.M.Cleef 1905 (COL). Cundinamarca: Municipio de Cógua, Páramo entre Cógua y San Cayetano, Laguna Verde, 3600 m, 30 Nov. 1971, A.M.Cleef 84 (COL). Meta: Macizo de Sumapaz, Laguna La Guitarra (lado nor-oeste), 3420 m, 5 Jul. 1981, J.Aguirre 2169 (COL), 5 Jun. 1981, J.Aguirre 2169a (COL). Tolima: Municipio de Santa Isabel, el Ochoral al Nevado del Tolima, 3580 m, 30 Jul. 1980, J.Aguirre 1655 (GOET). ECUADOR: Provincia de Carchi, El Ángel, 3730 m, 22 Dec. 1983, W.R.Buck 10255 (NY, MO); carretera Tulcán-El Ángel, 3630 m, 4 Apr. 1976, S.R.Gradstein et al. 3451, 3458, 6876 (U, GOET). BOLIVIA: Cochabamba, carretera Chapare-Incachasa, 3400 m, 8–13 Nov. 1989, S.R.Gradstein 7387 (U, GOET), Cordillera Real, cerca de Illampu, 4700 m, 1928, C.Troll s.n. (GOET, U).

2 Isotachis lopezii (R.M.Schust.) Gradst., Mem. New York Bot. Gard. 86: 84. 2001. Figs 2, 10 TYPE: Venezuela, Mérida: Sierra de Santo Domingo, Páramo de Mucubaji, 3600 m., Schuster & RuizTeran 76-870 (holotype, MERF!). Ruizanthus lopezii R.M.Schust., Phytologia 39(4): 240, 1978.

PLANTS prostrate to suberect, very small, up to 5 mm long, 1–2.5 mm wide, including leaves; weakly ramified, strongly anisophylous, pale green, except in the underleaves which are slightly pink pigmented. STEMS in transverse section with 13 cells on average; the cortical cells smaller and brown, 11–28 × 5–13 µm, distributed in 2–3 rows; the medullar cells larger and colorless, 34–45 × 33–40 µm. RHIZOIDS conspicuous, colorless, numerous, in tufts at base of underleaves. LEAVES clearly orbicular to slightly ovate, slightly wider than long, concave to channeled 1.5–2.0 × 2.0–2.5 mm, imbricate, bifid up to 1/5 of their length; segments short, widely triangular, with apex cuspidate, consisting of a column of 2–3 cells, the distal cell thin and acuminate; sinus acute to 105

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Fig. 1. Isotachis lacustris. A–B. Part of shoot, ventral view. C–G. Leaves. H–I. Underleaves. J. Stem, cross section. K. Marginal and apical leaf cells. L. Median leaf cells. (All from the type, Troll 56.)

lunate; margins entire or with 1–2 small teeth on each side; segments symmetrical, bases rotund; insertion of the leaves incubous, insertion line not deeply arcuate. 106

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LEAF CELLS forming a border of a clear row with uniformly incrassate walls, averaging 61 × 24 µm, the apical ones clearly isodiametric, with uniformly thin walls; trigones bulging, conspicuous; the median cells slightly larger as they approach the base, 28–51 × 17–25 µm, trigones conspicuous, the basal cell ones are equal to the median ones; cuticle clearly papillose. UNDERLEAVES smaller than leaves, almost twice as wide as stem, ovate, plane, imbricate, 0.8–1.1 × 0.5–0.9 mm, bifid up to 1/2 of their length; segments long to short acuminate, triangular; apex acute; sinus rotund to acute, with short to almost piliferous teeth, one or up to four teeth on both sides, line of insertion curved. ANDROECIA not seen. GYNOECIA bearing bracts greater than leaves, ovate to oblong, concave, bifid up to 1/3 of its length; segments broadly triangular; sinus round; 5 × 3 mm; bracteoles smaller than bracts, ovate, concave, bifid up to 1/3 of their length, 2.2 × 1.8 mm. PERIGYNIUM TRUE, rigid, erect to slightly arcuate, conicalcylindrical, narrowing towards the apex, ending in a ciliate mouth, covered by more than 50% of its length by bracts and bracteoles. Capsule ovoid, with spiral valves. H ABITAT AND DISTRIBUTION : Previosly reported from Ecuador and Venezuela. Colombia: Departamentos of Boyacá, Cundinamarca, Nariño and Tolima. Mainly in zones of peatlands, at altitudes between 3100 and 3800 m. Plants growing on ground and sometimes on rock. DIAGNOSTIC FEATURES: Isotachis lopezii has some characters similar to I. multiceps (see the latter), but it is possible to differentiate the two by the bordered leaves of I. lopezii. A suite of characters is also present in R. venezuelanus including leaves orbicular, underleaves long acuminate and clearly dentate, conspicuous presence of rhizoids in fascicles, and the procumbent to suberect habit. SPECIMENS EXAMINED: COLOMBIA: Boyacá: Páramos al NW de Belén, Quebrada Minas, Laguna El Alcohol, 3800 m, 26 Feb. 1972, A.M.Cleef 1887 (COL). Cundinamarca: Municipio de Guasca, Páramo de Guasca, 3150 m, 7 Aug. 1980, S.R.Gradstein & J.Aguirre 3679 (NY); Municipio de Fómeque, P.N.N.Chingaza, 3200 m, 5 Oct. 1982, E.Santana & J.Aguirre 309, 379, 397, 398 (COL); Páramo de Palacio, 3230 m, 12 May. 1972, A.M.Cleef 3686 (COL); Municipio de Subachoque, Cuchilla El Tablazo, 3500 m, 20 Oct. 1988, R.M.Schuster 88-980, 88-989 (COL). Nariño: Municipio de Pasto, Volcán Galeras, 4200 m, 29 Jun. 1991, R.Ramírez 3820 (MO); Túquerres, Páramo Volcán Azufral, 3800 m, 25 Feb. 1995, R.Ramírez 6888 (MO). Tolima: Municipio de Ibagué, Corregimiento de Juntas, Finca El Rancho, sector Filtros, 3500 m, 28 Jun. 2002, L.V.Campos 150 (TOLI). JAMAICA: near pass Buna-Buna, 2700 ft, 1 may. 1903, Underwood 2725 (NY); VENEZUELA: Municipio Libertador, Estado de Mérida, teleférico de Mérida, estación Loma Redonda, 4070–4100 m, 26 Feb. 1997, T.Pócs et al. 9715/AN (MERC); ECUADOR: Municipio de Carchi, Páramo del Ángel, 3400 m, 3 Apr. 1976, S.R.Gradstein et al. 3381 (NY); Provincia de Napo, Cordillera Oriental, Páramo de Papallacta, 4200 m, 20 Sep. 1982, W.C.Steere 25689 (NY). PERÚ: Chachapoyas, Calla-Calla, 3100 m, 31 Aug. 1973, P. & E.Hegewald 6940 (MO).

3 Isotachis multiceps (Lindenb. & Gottsche) Gottsche, Mexikanske Levermoser 105. 1863. Figs 3, 4, 11 TYPE: México, Sempoaltepec, Liebmann 137b, 1848, (holotype, S-PA!, isotype, BM!). Jungermannia multiceps Lindenb. & Gottsche, in Gottsche, Lindenberg & Nees, Syn. Hep.: 687. 1847. Jungermannia conduplicata Lindenb. in Gottsche, Lindenberg & Nees, Syn. Hep.: 680. 1847. Type: Brasil, "ad Rio Janeiro (Luschnath, Hb Ac. Petr.)". Isotachis tenax Steph., in: Urban, I.Symb. Antill. 3: 277. 1902.

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Fig. 2. Isotachis lopezii. A. Part of shoot, dorsal view. B. Perigynium with bracts. C–D. Leaves. E. Bract of perigynium. F. Bracteole of perigynium. G–I. Underleaves. J. Stem, cross section. K. Marginal and apical leaf cells. L. Bulging trigones of median leaf cells. (All from T.Pócs et al. 9715.)

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TYPE: Guadalupe, s. l. Dussén 550, 19 Jul. 1901 (holotype, G!, isotype, FH!, NY!), SYN. NOV. Isotachis parva Steph., Arq. Mus. Rio de Janeiro 13:113. 1903. TYPE: Serra Itataia, inter Retiro do Ramos et Maciera do Couto, 2100m, P. Dúsen s.n. Isotachis coilophylla Herzog, Repert. Sp. Nov. Regni Veg. 21:25. 1925. TYPE: Brazil, Serra dos Orgaos, Morro Assu, Luetzelburg 6035b.

PLANTS small, prostrate to ascendant, up to 15 mm long, 0.5–2 mm wide, only plant tips are observed as ascendant, strongly anisophylous, and pale green to reddish. STEMS reddish, 0.7–1.0 mm in diameter in transverse section, with 8–13 cells of diameter; the cortical cells smaller and brown, 12–25(–27) × (5)10–20 µm, in two to three rows; the medullary cells larger and colorless, 25–35 × 10–25 µm. RHIZOIDS colorless, numerous, in fascicles at the base of the underleaves. LEAVES subquadrate to rectangular, almost as long as wide or wider than long, concave, 1.0–1.3 × 0.5–1.5 mm, imbricate to strongly imbricate, bifid up to 1/2 their length; segments triangular, the apex acute, round to obtuse; sinus acute to lunate; margins entire to dentate, with 1–2 teeth on each side, 1–4 cells in length, acute or obtuse, line of insertion slightly curved. LEAF CELLS very variable as far as the shape and size, the apical ones irregularly isodiametric to slightly longer than wide, (17–)25–32 × (10–)22–25(–27) µm, with thickened walls; trigones inconspicuous; the median cells elongate to rectangular, (17–)34–50 × 11–23(–25) µm, thickened walls, trigones inconspicuous to cordate; cuticle smooth to striate-papillose. UNDERLEAVES smaller than leaves, subquadrate to more or less rectangular, concave, 420–620 × 450–570 µm, bifid up to 1/2 of their length; segments divergent, triangular, the apex round to acute, sinus lunate to acute; margins entire or dentate, with teeth on one or both sides, line of insertion arcuate. ANDROECIA with a single antheridium per bract. GYNOECIA on very short, ventral-intercalary branches; bracts progressively larger as they approach the perigynium, ovate to subquadrate, concave, bifid up to 1/3 of its length; segments triangular to broadly triangular, acute, sinus acute, 0.7–0.8(–1.7) × 0.7–0.9(–1.8) mm; bracteoles slightly smaller than bracts, ovate to subquadrate, concave, bifid up to 1/3 of their length, 0.7 × 0.7 mm. PERIGYNIUM TRUE, rigid, erect, conical-cylindrical, plicate, narrowing towards the apex, terminating in a short-ciliate mouth, pink to intense pink with off-white apex. SETA 5 mm long and 3 mm broad. CAPSULE oblong long-cylindrical, 1.0 mm long, spiral valves, with rotund apex, 2.2 mm long. SPORES slightly equinulate, reddish, 7 µm in diameter; ELATERS intense brown to yellowishreddish, double spiral, 180–270 µm long. HABITAT AND DISTRIBUTION: Mexico, Costa Rica, Puerto Rico, Guadalupe, Jamaica, Dominican Republic, Ecuador and Brazil. In Colombia distributed in the three Andean Cordilleras, the Sierra Nevada de Santa Marta and the mountainous area of La Macarena, mainly in moors over 3500 m, although it is also found in Andean and high-Andean forests, at altitudes between 2400 and 2700 m. Explanation of the presence in lower altitudes in La Macarena (500 m) and in the Sierra Nevada de Santa Marta (2200 m) is unsolved. DIAGNOSTIC FEATURES: Isotachis multiceps is clearly distinct from the remaining Isotachis species present in Colombia by its size, the weakly succubous leaf insertion, the presence of a single antheridium per bract, by microphyllous stolons. The leaves 109

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Fig. 3. Isotachis multiceps. A. Part of shoot with perigynium and microphyllous stolons. B. Part of shoot, ventral view. C–F. Underleaves. G–K. Leaves. L. Perigynium bracteoles. M–N. Perigynium bracts. O–P. Apical leaf cells. Q–R. Median leaf cells. S. Stem, cross section. (All from type, Liebmann 137b.)

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Fig. 4. Isotachis multiceps. A. Part of shoot, ventral view. B–G. Leaves. H–K. Underleaves. L. Median leaf cells. M. Marginal leaf cells. N. Stem, cross section. (All from type of I. tenax, Dussen 550).

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have small marginal teeth, one to two at each side, the bases of the underleaves have abundant rhizoids distributed in fascicles and habit is prostrate to ascendant. The last two characters appear to be correlated. Type of growth and distribution of rhizoids in this species can be compared with I. lopezii but the two species cannot be confused. I. multiceps has leaves with marginal cells larger than the laminar ones, the leaf segments are very broad and cuspidate with an apex of 1–3 cells, underleaves are bifid, with long and numerous spine-like projections on both sides. Engel & Merrill (1997) proposed Hypoisotachis as a separate genus, but this proposal is not accepted here, considering the lack of an exhaustive study of neotropical specimens, necessary to determine if the characters supporting Hypoisotachis as a genus are stable or not. SPECIMENS EXAMINED: COLOMBIA: Boyacá: Sierra Nevada El Cocuy, alto Valle Lagunillas, quebrada La Bocatoma, 4200 m, 28 Sep. 1972, P.A.Florschütz 4053 (COL); Páramo de la Rusia, NW de Duitama, 1 km. SE de Laguna Negra, 3745 m, 15 Dec. 1972, A.M.Cleef 7275 (COL). Cundinamarca: Municipio de Fómeque, P.N.N.Chingaza, 3400 m, 23 Nov. 1982, S.R.Gradstein & E.Santana 4251 (NY); Páramo de Cruz Verde, Alto de La Viga, 3560 m, 28 Apr. 1972, A.M.Cleef 3280 (COL); Páramo de Palacio, Lagunas de Buitrago, 3560 m, 25 May. 1972, A.M.Cleef 4126 (COL); Páramo de Palacio, 3485 m, 16 Dec. 1971, A.M.Cleef 331 (COL). Huila-Cauca: Macizo Colombiano, Páramo de las Papas, Lagunas La Magdalena y Santiago, 3530–3630 m, 5–19 Nov. 1958, H.Bischler 786 (COL). Magdalena: Sierra Nevada de Santa Marta, al norte de San Lorenzo, 2200 m, 12 Jan. 1967, S.Winkler C319 (COL). Meta: Páramo de Sumapaz, 4015 m, 13 Jan. 1973, A.M.Cleef 7761, 7761b (COL); La Macarena, 500 m, 10 Mar. 1957, R.E.Schultes 11668 (COL). Putumayo: Municipio de Sibundoy, Páramo La Rejoya, 2600 m, 30 Mar. 2010, L.V.Campos 696, 697. Risaralda: Municipio de Santa Rosa de Cabal, entre Termales y Volcán Otún, 2400 m, 16–19 Jul. 1980, S.R.Gradstein 3517 (COL). MÉXICO: Sempoaltepec, Liebman s.n. (BM00918036, BM00918037); valle de Bravo, 1800 m, 22 Oct. 1966, R.Düll s. n (NY). COSTA RICA: Provincia Limón, P.N.Braulio Carrillo, quebrada Gonzales, 450 m, 19 Apr. 2001, G.Dauphin 2956 (NY); Puntarenas, Cantón de Coto Brus, 2300 m, 7 Feb. 1995, G.Dauphin 1491 (NY). GUADALUPE: s.l. 1903, Pére Dussen 60 (NY). JAMAICA: Blue Mountain peak, 21 Jul. 1903, A.W.Evans 234 (NY); 7400 ft, 12 Apr. 1945, N.W.Simmonds 150 (NY); New Haven gap, 16 Jul. 1903, A.W.Evans 154 (NY). REPÚBLICA DOMINICANA: Provincia La Vega, La Cotorra, Pico Duarte, 5000 ft, 16 Jan. 1987, W.R.Buck 14293 (NY); arroyo Los Flacos, cerca de Pirámides, valle Nuevo, 7400 ft, 15 May. 1982, W.R.Buck 8635 (NY).

4 Isotachis serrulata (Sw.) Gottsche, Ann. Sci. Nat. Bot. ser. 5. 1:121. 1864. Figs 5, 6, 12 TYPE: Jamaica, s.l. Swartz, s.n. (Syntype, BM!). Jungermannia serrulata Sw., Prodromus Flora Ind. Occ.: 143. 1788. Isotachis haematodes (Lehm. & Lindenb.) Gottsche, Ann. Sci. Nat. 1: 122. 1864. TYPE: Guadelupe, without collector, ex Hb. Lehmann (isotypes, BM, G) Isotachis madida (Hook. f. & Tayl.) Mitt., in: Hooker, J. D. Bot. Antarctic Voy. 2(2): 149. 1855. TYPE: Tierra del Fuego, Cape Horn, Hooker, s n. (isotypes, NY, BM). Isotachis lindigiana Gottsche, Ann. Sci. Nat. Ser. V. 1: 123. 1864. TYPE: Colombia, Bogotá, Lindig 1702, (holotype S-PA!, isotypes G!, BM!), SYN. NOV. Isotachis mascula Gottsche, Ann. Sci. Nat. Bot. V. 1: 199. 1864. TYPE: Colombia, Bogotá, Fusagasuga, 1800 m (sintypes, Lindig 1722, 2104, 2140, 2904 not seen), SYN. NOV.

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PLANTS large, erect, up to 9 cm long, 0.5–2 mm broad, including the leaves, strongly anisophylous, brown to reddish, young sprouts pink, ramified or not. STEMS brown dark, in transverse section 13–15 cells in diameter; the cortical cells smaller and dark brown, 15–30 × 15–27 µm, in 1–3 rows; the medullary cells larger and colorless, 40–60 × 25–37 µm. RHIZOIDS absent, in some specimens a few were observed, scattered and colorless. LEAVES strongly asymmetrical, subquadrate to ovate, concave, canaliculate, 1.1–2.7 × 0.9–2.4 mm, imbricate to adjacent, bifid from 1/4 to 1/2 their length; segments triangular, the apex obtuse to acute, sinus acute to rotund; margins entire, dentate to serrulate; teeth 1-3 cells long, acute, present in the margin of the leaves including the sinus; apical cell may be pink-pigmented and triangular (associated with leaves with serrulate margins); dorsal segment larger than ventral; insertion incubous; ventral portion line of insertion transverse, dorsal portion line of insertion longitudinal. Leaf CELLS near apex ± isodiametric to slightly longer than wide, 27– 62 × 20–52 µm, with thickened walls; trigones conspicuous; the median and basal cells rectangular, 42–80 × 15–22 µm, incrassate or thin walled; trigones generally conspicuous; cuticle striate-papillose. UNDERLEAVES smaller than leaves, subquadrate to ovate, imbricate to approximate, plain to reflexed, symmetrical 1.0–2.5 × 0.8–1.5 mm, bifid 1/4–1/3 of their length, apex obtuse to acute, sinus rounded to acute; margins entire, dentate to serrulate; bases slightly auriculate; line of insertion arcuate. ANDROECIA not seen. GYNOECIA terminal; bracts larger than leaves but conserving their shape, ovate to subquadrate, concave, narrowing towards the base, bifid, acute; segments widely triangular, sinus acute, 2.1–3.0 × 1.8–2.2 mm; bracteoles slightly smaller than bracts, ovate to subquadrate, concave narrowing towards the base, bifid, 2.0–2.2 × 1.7–1.9 mm. PERIGYNIUM TRUE, rigid, erect, longoblong, narrowing towards the apex, which terminates in a narrow mouth with small and short scales; this terminal part is of less intense color than the rest. SETA 2 cm long. CAPSULE ovoid, 1.5 mm, spiral valves, with round apex, 2.2 mm long. SPORES smooth, reddish. ELATERS of intense brown to reddish, bispiral, 300 µm long. HABITAT AND DISTRIBUTION: Widely distributed in tropical America. In Colombia it is the most widely distributed species of the family, from 1300 to 4500 m, distributed in the three Andean Cordilleras, from the south in Nariño and Putumayo to the páramos in the north of Boyacá and Santander. In the Western Cordillera the species is present in the highest hills of San Jose del Palmar, Tatamá hill. In the Central Cordillera it has been found in the departamentos of Tolima and Risaralda, and it is likely to be in Los Nevados National Park. It has also been reported from the Sierra Nevada de Santa Marta. Mainly on soil and along roads and cliffs. DIAGNOSTIC FEATURES: Isotachis serrulata is unmistakable in relation to the rest of Isotachis species present in Colombia, owing to the fact that plants are large, up to 9 cm long, reddish to yellowish-brown, with asymmetric leaves that are clearly ovate to subquadrate, bifid up to 1/2 of their length, and the median cells of the leaves are up to two times longer than wide. Most important is the presence of indented to serrulate margins; in some cases they may be entire, but if this is the case I. serrulata can be differentiated from I. lacustris by the dark brown to blackish pigmentation of I. lacustris, by symmetrically ovate-lanceolate leaves, which are bifid up to 1/6 of their length and median leaf cells four or more times longer than wide.

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Isotachis serrulata is the species with greatest morphological variability among neotropical members of the genus, particularly the presence of teeth on the margins of leaves and underleaves, even within a single individual. Leaves without teeth as well as those that are strongly dentate to serrulate are generally present on young branches. These branches coincide with the type specimen of I. lindigiana, but older branches in the same specimen have leaves that morphologically and anatomically correspond to I. serrulata. The two types represent the ends of a broad range of morphological and anatomical variation, and therefore I. lindigiana is treated here as synonymous with I. serrulata. Gottsche (1864) described I. mascula based on specimens collected in Colombia, from the municipality of Fusagasuga (Lindig 1722, 2104, 2140, 2904) 1800 m. These specimens were lost in the Second World War in Berlin. However, it is clear from the original description that I. mascula belongs to I. serrulata. Gottsche (1864) described it as new because of the presence of scales throughout the perigynium. Scales frequently occur in many specimens of I. serrulata. These scales are vestiges of the small perianth. SPECIMENS EXAMINED: COLOMBIA: Arauca: Sierra Nevada del Cocuy, quebrada El Playón, Patio Bolos, 4190 m, 8 Mar. 1973, A.M.Cleef 8927 (COL). Boyacá: Municipio de Belén, Páramos al NW de Belén, quebrada Minas, Laguna El Alcohol, 3785 m, 26 Feb. 1972, A.M.Cleef 1881, 1905 (COL); Municipio de Duitama, Páramo de la Rusia, 3500 m, 26 Oct. 1988, G.Hassel de Menéndez 5744, 5764 (COL); Municipio de Villa de Leyva, Santuario de fauna y flora de Iguaque, camino a las lagunas, 2920 m, 23 Nov. 1992, J.Uribe 2726 (COL); Cauca: Volcán Puracé, Laguna de San Rafael, 3300 m, 6 Jan. 1972, A.M. Cleef & A. Fernández 586 (COL); Macizo colombiano, Páramo de las Papas, entre el Boquerón y la Hoyola, 3200–3510 m, 7–27 Nov. 1958, H.Bischler 789 (COL). Casanare: Municipio de Sácama, carretera Socha-Sácama, Páramo de Pisba, 3050 m, 15 Aug. 1982, J.Aguirre et al. 2853 (COL) Cundinamarca: Municipio de Cógua, páramo entre Cógua y San Cayetano, laguna Verde, 3625 m, 19 Aug. 1972, A.M.Cleef et al. 5170 (COL); Municipio de Choachí, Páramo de Cruz Verde, 3350 m, 22 Apr. 1972, A.M.Cleef 3001 (COL); Municipio de Guasca, Páramo de Guasca, 3150 m, 29 Dec. 1971, A.M.Cleef 376 (COL); Páramo de Sumapaz, valle Quebrada Honda, Andabobos, 3715 m, 10 Feb. 1972, A.M.Cleef 1559 (COL). Chocó: Municipio de Anserma Nuevo, San José del Palmar, 1542–2050 m, 15 May. 1984, J.Luteyn 10554 (NY). Magdalena: Sierra Nevada de Santa Marta, San Lorenzo, 2000 m, 16 Jan. 1967, S.Winkler C97 (COL). Meta: Páramo de Sumapaz, Cerro Nevado, 3590 m, 12 Jan. 1973, A.M.Cleef 7715 (COL). Nariño: Municipio de Pasto, Corregimiento El Encanto, Km. 28 vía Sibundoy, 2900–3000 m, 21 Jun. 1993, B.Ramírez 5382 (COL). Putumayo: Municipio de Mocoa, carretera a Sibundoy, 1150 m, 4 May. 1994, Giraldo-Cañas 2197 (MO). Risaralda: Municipio de Pereira, vereda El Cedral, Parque Ucumarí, 2500 m, 10–17 Jun. 1989, J.Uribe & J.Aguirre 308, 364 (COL); Municipio de Mistrató, 1500 m, 25 Jul. 1992, S.R.Gradstein 8411, 8480 (COL). Santander: Municipio de Suaita, 1500 m, 1 Aug. 2001, J.Uribe et al. 3943 (COL). Tolima: Municipio de Santa Isabel, finca El Ochoral, quebrada Las Lomas, 2950 m, 28 Jul. 1980, J.Aguirre 1580 (COL); COSTA RICA: Puntarenas, Cantón de Coto Brus, 1000–1200 m, 30 Oct. 1995, G.Dauphin 2037 (NY). GUADALUPE: Saint Louis, 100–200 m, 1899, Pére Dussen 269 (NY); Soufriere, 1300–1470 m, 1898, Pére Dussen 371 (NY). JAMAICA: Swartz (BM000918038) (NY); Near Hardwar gap, 4000 ft, 15 Abr. 1903, Underwood 2222 (NY); Road from Cinchona to Morce's gap, 5000 ft, 30 Ene. 1903, Underwood 312 (NY). MARTINICA: Montagne Pelée, 1250–1350 m, 1868, Husnot s.n. (BM000918039) (NY); Deux-choux, Sep. 1901, Pére Dussen 605 (NY); Bon de Colson, Sep. 1903, Pére Dussen 1209 (NY). ECUADOR: Provincia de Napo, Sierra de Guacamayas, 2100 m, 12 Ene. 1981, W.C.Steere E 121, E 147 (NY). BOLIVIA: Departamento de Yungas, 6000 pies, H.H.Rusby 5037. BRASIL: Provincia de Sao Paulo, Santos, alto da Serra, 25 Feb. 1875, Mosén 33 (NY); Rio de Janeiro, Burchell 3713 (NY); Brasilia, Montagne s.n. (NY). PERÚ: Departamento del Amazonas, Chachapoyas, 2900 m, 4 Sep. 1982, G.Philippi 754 (NY).

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Fig. 5. Isotachis serrulata. A. Part of shoot, ventral view. B. Perigynium with bracts and bracteoles. C–D. Leaves. E–F. Perigynium bracts. G–I. Underleaves. J. Apical and marginal leaf cells. K. Median leaf cells. L. Stem, cross section. (All from type, Swartz s.n.)

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Fig. 6. Isotachis serrulata. A. Part of shoot, ventral view. B–F. Leaves. G–M. Underleaves. N–O. Bracts and Bracteoles. P. Stem, cross section. Q. Median leaf cells. (All from type of I. lindigiana, Lindig 1702.)

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2 Neesioscyphus Grolle Neesioscyphus Grolle, Oesterr. Bot. Z. 111: 19. 1964.

Figs 7, 8

TYPE: Neesioscyphus carneus (Nees) Grolle, Oesterr. Bot. Z. 111: 19. 1964.

PLANTS very small, prostrate, strongly anisophyllous, pale green to reddish, with ventral-intercalary ramification. STEMS in transverse section with small, brown cortical cells in 2–3 rows, the medullary cells larger and colorless. RHIZOIDS colorless to slightly red pigmented, in fascicles at the base of some underleaves. LEAVES subquadrate to ovate, flat, with acute to rotund apex, sinus lunate to obtuse; margins entire; insertion clearly succubous. Leaf CELLS with thin walls, without trigones; the median cells elongate to rectangular; cuticle striate-papillose. UNDERLEAVES smaller than leaves, subquadrate to more or less rectangular, flat, bifid up to 1/2 of their length: segments emarginate, triangular, lanceolate, cuspidate; margins entire or with a single teeth on each side; line of insertion arcuate. ANDROECIA bearing bracts with 2 antheridia. GYNOECIA bearing a true perianth as the protective structure of the sporophyte. CAPSULE ovoid and with spiral valves. HABITAT AND DISTRIBUTION: Neesioscyphus includes 6 species, 5 in the Neotropics and 1 in New Zealand. The center of diversity of this genus is the southeast of Brazil (Gradstein et al. 2001). In Colombia there are 2 species in the Eastern Cordillera, in the Departamento of Casanare, in the foothills of the Eastern Plains, and in the Western Cordillera, at the boundary between Risaralda and Chocó. Growing on steep banks, on earth and on thin soil. Key to the species of Neesioscyphus in Colombia 1

Dorsal segment of the leaf larger than the ventral one. Underleaves more than 1.5 to 2 times as wide as the stem. Mouth of the perianth with short teeth........................................................ 1 N. allioni 1' Ventral segment of the leaf larger than the dorsal one. Underleaves as wide as the stem. Mouth of the perianth with numerous spines............................................................................. 2 N. argillaceus

1 Neesioscyphus allionii (Steph.) Grolle, Rev. Bryol. Lichénol. 34 (1–2): 185. 1966. Figs 7, 10 Isotachis allionii Steph., Sp. Hepat. 6: 350. 1922. TYPE: Ecuador, Río San José, V.Bomboiza, 1300 m, 1910, Allionii 226 (holotype, G!)

PLANTS very small, prostrate, up to 9 mm long, 3 mm broad, including the leaves, strongly anisophyllous, pale green to reddish, ramification ventral-intercalary. STEMS in transverse section on average 10 cells in diameter; the cortical cells smaller and brown, 17–23 × 25–35 µm, in 2–3 rows; the medullary cells larger and colorless, 22–37 × 37–64 µm. RHIZOIDS slightly red pigmented, scarce, in fascicles at the base of some underleaves. LEAVES subquadrate to ovate, flat, 1.5–2.0 × 1.1–1.5 mm broad, bifid up to 1/3 their length, segments triangular, the apex acute; sinus lunate; margins entire; dorsal lobe larger than the ventral one; insertion clearly succubous. Leaf CELLS of apex irregularly rectangular, 17–25 × 32–60 µm, with thin walls,

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Fig. 7. Neesioscyphus allionii. A. Part of shoot, ventral view. B. Part of shoot, dorsal view. C. Perianth with bracts. D-E. Leaves. F. Underleaf. G. Bracteole. H. Perianth bract. I. Apical and marginal leaf cells. J. Median leaf cells. K. Stem, cross section. (All from Gradstein 8391.)

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trigones absent; median cells rectangular 28 × 55 µm, thin walls, trigones absent; the basal cells equal to the median ones, trigones inconspicuous; cuticle striate-papillose. UNDERLEAVES smaller than leaves, subquadrate to more or less rectangular, flat, 0.6– 1.5 × 0.4–0.8 mm, bifid up to 1/2 of their length, triangular, lanceolate-cuspidate; segments emarginate; margins with a single tooth on each side; insertion line arcuate. ANDROECIA not seen. GYNOECIA with bracts larger than the leaves, 3 × 1.3 mm, ovate, concaves, bifid up to 1/3 of its length; segments triangular, emarginate; sinus acute; bracteoles smaller than the bracts, 1.3 × 0.8 mm, ovate, concave, bifid up to 1/3 of their length. TRUE PERIANTH, 4 mm long, membranous, erect, conical-cylindrical, the mouth dentate-ciliate. DISTRIBUTION: The only specimens are the type from Ecuador and a single Colombian specimen collected in the Western Cordillera, near the border between Risaralda and Chocó, at 1500 m. DIAGNOSTIC FEATURES: Neesioscyphus allionii is similar to N. argillaceus, from which it can be distinguished by the following characters: dorsal segment of the leaf larger than the ventral one; segments apiculate; underleaves almost double the width of the stem, with small teeth, only one on each side, and perianth mouth dentate-ciliate. N. allionii was first described as a species of Isotachis by Stephani (1922). Grolle (1966) proposed to transfer it to Neesioscyphus. The clearly succubous insertion of the leaves and a true perianth in the perianth-perigynium complex are diagnostic characters of the latter genus. SPECIMENS EXAMINED: COLOMBIA: Risaralda: Cordillera Occidental, municipio de Mistrató, entre Jeguadas y Puerto de Oro, 1500 m, 25 Jul. 1992, S.R.Gradstein 8391 (COL).

2 Neesioscyphus argillaceus (Nees) Grolle Oesterr. Bot. Z. 111: 24. 1964 Figs 8, 10 TYPE: Brasil, Minas Geraes, Serra de Estrella, without collector (holotype, G!). Jungermannia argillacea Nees in: Martius, Flora Brasil. 1:338. 1833. Gymnomitrium argillaceum (Ness) Gottsche in Gottsche, Lindenderg & Nees, Syn. Hep.: 5. 1844. Acolea argillacea (Nees) Trevisan, Mem. Real. Istit. Lombard. Sci. Mat. Nat. Ser. 3, 13: 395. 1877. Jungermannia heteracria Spruce, Trans. Proc. Bot. Soc. Edinburgh 15: 512. 1885. TYPE: Perú. "Circa Tarapoto", 500–800 m. Notoscyphus argillaceus (Nees) Spruce, Rev. Bryol. 15: 34. 1888. Cesiusa argillacea (Nees) Kuntze, Revisio Gen. Plant.: 833. 1891.

PLANTS medium, prostrate, up to 1cm long, 1–1.5 mm broad, including the leaves, plants strongly anisophyllous, brown to pink; ramification ventral intercalary. STEMS in transverse section 10 cells in diameter on average; the cortical cells smaller and brown, 11–18 × 13–28 µm, in 1–2 rows; the medullary cells larger and colorless, 15–20 × 20–28 µm. RHIZOIDS slightly red pigmented, abundant, in fascicles at the base of the underleaves. LEAVES round to ovate, flat, bifid, 0.6–0.8 × 0.5–0.8 mm, slightly bifid (up to 1/6 of their length) segments triangular, the apex obtuse to round, sinus round; margins entire; dorsal lobe shorter than the ventral one; insertion clearly succubous. Leaf CELLS: apical cells subquadrate to rectangular, 17–27 × 119

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Fig. 8. Neesioscyphus argillaceus. A. Part of shoot, ventral view. B. Perianth with bracts and bracteoles. C–D. Leaves. E–G. Underleaves. H. Underleaves with marginal papillae. I. Bracteole. J. Perianth bract. K. Stem, cross section. L. Apical and marginal leaf cells. M. Median leaf cells. (All from Aguirre 3033.)

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Fig. 9. Ruizanthus venezuelanus. A. Perianth with bracts and bracteoles. B. Part of shoot, dorsal view. C. Leaf. D–E. Underleaves. F. Perianth bracteole. G. Perianth bract. H. Stem, cross section. I. Median leaf cells. J. Apical and marginal leaf cells. (All from Orrego 156.)

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Fig. 10. Distribution of Isotachis lacustris, I. lopezii, Neesioscyphus allionii, N. argillaceus and Ruizanthus venezuelanus in Colombia.

37–50 µm, walls thin, trigones absent; median cells elongate to rectangular, 18–28 × 44–65 µm, walls thin, trigones absent; cuticle striate-papillose. UNDERLEAVES much smaller than leaves, not much wider than stem, rectangular, flat, 0.3–0.4 × 0.1–0.3 mm, bifid up to 1/2 of their length, some times more than half; segments divergent, triangular, lanceolate, cuspidate, rarely with marginal teeth; line of insertion arcuate; marginal papillae present. ANDROECIA not seen. GYNOECIA with bracts almost double the leaves, 1.7 × 1.4 mm, ovate, concave, bifid up to 1/4 of their length; segments triangular, emarginated, sinus broad; bracteoles smaller than the bracts, 0.6 × 0.3 mm, the same shape as the underleaves, concave, bifid up to 1/2 of their length. TRUE PERIANTH, membranous, erect, 4 mm long, terminating in a mouth with numerous spine-like projections. DISTRIBUTION: Costa Rica, Panama, Venezuela, Ecuador, Bolivia, Brazil and Peru. In Colombia this species occur in the Departamentos of Casanare, Boyacá and Valle del Cauca. DIAGNOSTIC FEATURES: Neesioscyphus argillaceus can be confused with N. allionii, but the ventral segment of the leaf is larger than the dorsal one, the segments are 122

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Fig. 11. Distribution of Isotachis multiceps in Colombia.

obtuse to rotund, and the underleaves are almost the width of the stem and the perianth mouth is spinulose. SELECTED SPECIMENS EXAMINED: COLOMBIA; Boyacá: Municipio de Santa María, quebrada La cristalina, 900 m, 23 Apr. 2005, L.V.Campos 236 (COL). Casanare: Municipio de Sácama, quebrada Macueque, rio Sácama, 1300 m, 17 Aug. 1982, J.Aguirre, S.R.Gradstein & E.Santana 3033 (COL). Valle del Cauca: Municipio de Buenaventura, carretera entre Buenaventura y Málaga, 100 m, 15 Jul. 1993, Croat 75738 (COL, MO). VENEZUELA: Río Negro, Amazonas, Cañón Grande, río Agua Blanca, 150 m, 3 Feb. 1985, W.R.Buck 12765 (NY). ECUADOR: Islas Galápagos, San Cristóbal, Tres Palacios hacia El Junco, 400 m, 22 May. 1976, S.R.Gradstein & J.Lanier H 303, 307 (NY). BOLIVIA: Apolo, 4800 ft, 15 Apr. 1902, R.S.Williams 2229 (NY). BRASIL: Río de Janeiro, Glaziovi 9093, 9193 (NY). PERÚ: San Martín, provincia de Rioja, carretera entre Moyabamba y Chachapoyas, 1300 m, 28 Aug. 1982, G.Philippi 36 (NY).

3 Ruizanthus R.M.Schust. Ruizanthus R.M.Schust., Phytologia 39 (4): 240. 1978. TYPE: Ruizanthus venezuelanus R.M. Schust., Phytologia 39(4): 240. 1978. One species.

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Figs 9, 10

Fig. 12. Distribution of Isotachis serrulata in Colombia.

Ruizanthus venezuelanus R.M.Schust., Phytologia 39(4): 240. 1978. TYPE: Venezuela, Mérida, Sierra Nevada de Mérida, at Loma Redonda station of the Telepherico. R.M.Schuster & L.Ruiz-Terán 76-1462 (Holotype MERF!).

PLANTS small, procumbent, up to 15 mm long, 1–2 mm broad, including leaves, strongly anisophyllous, green to pale pink, more intensely pink in the underleaves, with intercalary ventral ramification. STEMS in transverse section 9 cells in diameter, with a slight differentiation in coloration between cortical and medullary cells, although the cortical ones are smaller, 10–20 × 15–30 µm, distributed in 1–2 rows; medullary cells larger and slightly colorless, 15–20 × 22–30 µm. RHIZOIDS conspicuous, colorless, numerous, in fascicles at the base of underleaves. LEAVES almost orbicular, concave, 1.0–1.4 × 0.8–1.3 mm, imbricate to contiguous, quadrifid up to 1/2–1/3 of their length; segments piliferous, with ranks of up to 5 cells, the last one of these more elongate and acuminate, sinus clearly rotund; margins entire; bases round; dorsal portion insertion transverse, line of insertion weakly arcuate. LEAF CELLS: apical cells subisodiametric to rectangular; the median ones slightly larger, 22–30 × 17–20 µm, with thin walls, trigones inconspicuous; cuticle strongly papillose. UNDERLEAVES smaller than leaves, as wide as the stem, ovate, planar, imbricate,

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0.6–0.8 × 0.3–0.5 mm, bifid up to 1/2 of their length or bisbifid, sinus round, with long piliferous teeth, 1–3 at each sides; line of insertion straight. ANDROECIA not seen. GYNOECIA bearing bracts larger than leaves, 2.0 × 1.2 mm, ovate to oblong, concave, bifid up to 1/4 of their length; segments similar to those of the leaves; sinus round; bracteoles smaller than bracts, 1.7 × 0.8 mm, conserving the same shape as the underleaves, concave, bifid up to 1/2 of their length. TRUE PERIANTH conicalcylindrical, narrowing towards the apex, terminating in a cut-ciliate mouth, covered by bracts and bracteoles over more than 50% of its length, multistratose only in the basal part. CAPSULE ovoid with straight valves. SPORES 15 µm in diameter, yellow, surface somewhat reticulate. ELATERS bispiral, 220 × 5 µm, brown. DISTRIBUTION: Costa Rica and Venezuela. In Colombia known from a specimen collected in Sierra Nevada de Santa Marta at 3500 m. DIAGNOSTIC FEATURES: Ruizanthus venezuelanus is an unmistakable species within the family, with the following diagnostic characters: quadrifid leaves with piliferous segments, bisbifid underleaves and capsules with of straight valves. SPECIMENS EXAMINED: COLOMBIA: Magdalena: Sierra Nevada de Santa Marta, alto La Cumbre, 3500 m, Jul. Aug. 1977, O.Rangel 897. VENEZUELA: Mérida, municipio Libertador, P.N.Sierra Nevada, estación La Aguada, 3340 m, 05 Feb. 1997, Pócs et al. 9702/VI (MERC), 22 Feb. 1997, Pócs, León & Frahm TP 9720/A.B. (MERC); teleférico de Mérida, estación Loma Redonda, 4045 m, 26 Feb. 1997, O.Orrego 156 (COL). Acknowledgements We are thankful to Robbert Gradstein for discussions on the taxonomy of the family and comments on nomenclatural issues. Thanks to the Colombian National Herbarium (COL) of the National University of Colombia, where the present study was done, and to the curators of the following herbaria: British Museum (BM), University of Harvard Herbarium (FH), University of Göttingen Herbarium (GOET), University of Jena Haussknecht Herbarium (JE), Missouri Botanical Garden (MO), New York Botanical Garden (NY), Herbarium of the Swedish Museum of Natural History (S) and the Universidad de los Andes Herbarium in Mérida (MERC), for loan of specimens. Special thanks to John Engel (Field Museum, Chicago), Juan Carlos Villarreal (University of Connecticut) and Lauren Raz (National University of Colombia) for comments on the manuscript. References CHURCHILL, S.P., N.N. SANJINES & C. ALDANA (2009): Catálogo de los briofitos de Bolivia: Diversidad, distribución y ecología. – Missouri Bot. Gard. Museo Hist. Nat. Noel Kempff Mercado. – Santa Cruz, Bolivia. 340 pp. CRANDALL-STOTLER, B., R.E. STOTLER & D.G. LONG (2009a): Morphology and classification of the Marchantiophyta. – In: GOFFINET, B. & A.J. SHAW: Bryophyte biology, pp. 1–54. – Cambridge Univ. Press, Cambridge. CRANDALL-STOTLER, B., R.E. STOTLER & D.G. LONG (2009b): Phylogeny and classification of the Marchantiophyta. – Edinburgh J. Bot. 66: 155–198. ENGEL, J.J. & G.L.S. MERRIL (1997): Austral Hepaticae. 22. The genus Balantiopsis in New Zealand, with observations on extraterritorial taxa and a phylogeny of Balantiopsis and the family Balantiopsaceae (Jungermanniales). – Fieldiana, Bot., n.s. 37: 1–62. EVANS, A.W. (1939): The classification of the Hepaticae. – Bot. Rev. 5: 49–96.

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FREY, W. & M. STECH (2009): Marchantiophyta, Bryophyta, Anthocerotophyta. – In: FREY, W.: Syllabus of Plant Families, Part 3: 9–263. – Gebr. Borntraeger; Berlin, Stuttgart. FULFORD, M.H. (1963): Isotachis. Manual of the leafy Hepaticae of Latin America. Part I. – Mem. New York Bot. Gar. 11(1): 63–79. FULFORD, M. & R.E. HATCHER (1958): Triandrophyllum, a new genus of leafy Hepaticae. – Bryologist 61: 276–285 GOTTSCHE, C.M. (1863): De Mexikanske Levermosser, pp. 1–284. – Kongelige Danske Videnskab.Selskabs, Kjobenhavn GOTTSCHE, C.M. (1864): Hepaticae. – In: TRIANA, J.J. & J.E. PLANCHON (eds.): Prodromus florae novo-granatensis. – Ann. Sci. Nat. Bot. 5, 1: 95–198. GRADSTEIN, S. R. & W.H.A. HEKKING (1979): Studies on Colombian Cryptogams IV. Catalogue of the Hepaticae of Colombia. – J. Hattori Bot. Lab. 45: 93–144. GRADSTEIN, S. R., A. M. CLEEF & M. H. FULFORD (1977): Studies on Colombian Cryptogams II. Hepaticae: Oil body structure and ecological distribution of selected species of tropical Andean Jungermanniales. – Proc. Kon. Ned. Acad. Wetensch., Ser. C 80: 377–420. GRADSTEIN, S.R., S.P. CHURCHILL & N. SALAZAR-ALLEN (2001): Guide to the bryophytes of Tropical America. – Mem. New York Bot. Gard. 86: 1–577. GROLLE, R. (1964): Neesioscyphus - eine neue Lebermoosgattung mit gedrehten Sporogonklappen. – Oesterr. Bot. Z. 111: 19–36. GROLLE, R. (1966): Notulae hepaticologicae XIV. Zwei weitere Neesioscyphus-Arten. – Rev. Bryol. Lichenol. 34: 182–186. GROLLE, R. (1972): Die Namen der Familien und Unterfamilien der Lebermoose (Hepaticopsida). – J. Bryol. 7: 201–236. HATCHER, R.E. (1960): A monograph of the genus Isotachis. I (Hepaticae). – Nova Hedwigia. 2: 573–608. HATCHER, R.E. (1961): A monograph of the genus Isotachis. II (Hepaticae). – Nova Hedwigia. 3: 1–35, pl. 1–30. HENTSCHEL, J.R. WILSON, M. BURGHARDT, H.-J. ZÜNDORF, H. SCHNEIDER & J. HEINRICHS (2006): Reinstatement of Lophocoleaceae (Jungermanniopsida) based on chloroplast gene rbcL data: exploring the importance of female involucres for the systematic of Jungermanniales. – Pl. Syst. Evol. 258: 211–226. HE-NYGRÉN, X. (2007): Multi-gene phylogeny supports single origin of jungermannioid perigynium. – Ann. Bot. Fe4nnici 44:450–462. HODGSON, E.A. (1949): A review of the New Zealand species of the genus Isotachis. – Rev. Bryol. Lichen. 18: 25–31. KITAGAWA, N. & R. GROLLE (1985): A new name for Acroscyphus N. Kitag., Hepaticae. – Acta Phytotax. Geobot. 36(1–3): 58. LEÓN-YÁÑEZ, S., S. R. GRADSTEIN & C. WEGNER (2006): Hepáticas (Marchantiophyta) y Anthoceros (Anthocerotophyta) del Ecuador. Cátalogo. Herbario QCA, Ponitficia Universidad Católica del Ecuador, Quito. 101 pp. MAGILL, R.E. (ed.). (1990): Glossarium Polyglottum Bryologiae. A multilingual glossary for bryology. – Monogr. Syst. Bot. Missouri Bot. Gard. 33: 1–297. MALCOLM, B. & N. MALCOLM (2000): Mosses and other bryophytes: An illustrated Glossary. Micro-Optics Press. New Zealand. 220p.

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Manuscript received January 27, 2011; accepted August 10, 2011.

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