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Sinopsis de la familia Frullaniaceae (Marchantiophyta) para Colombia.

INTRODUCTION

The liverwort family Frullaniaceae consists of two genera, Frullania with about 300-350 species worldwide and Neohattoria with one species in eastern Asia (Gradstein et al., 2001, with updates). In Colombia, Frullania is the second-largest liverwort genus (after Plagiochila) with 59 species. The first records from Colombia were by Alexander von Humboldt and Aimee Bonpland who collected Frullania atrata

(Sw.) Nees (as Jungermannia atrata Sw.) and F. riojaneirensis Raddi (as Jungermannia obscura Sw.) between Popayan and Almaguer, Dept. Cauca (Kunth, 1822). Gottsche (1864) reported 17 species from various localities based on specimens collected by Alexander Lindig and by Jose Jeronimo Triana. The majority of these early collections from Colombia have not been re-examined critically; those studied by Gottsche were presumably destroyed in 1944 with the bombing of the herbarium of Berlin. A further 30+ species were reported as new to Colombia by Jack & Stephani (1892), Stephani (1901-1905), Herzog (1942, 1955), Bonner (1965), Robinson (1967) and others. Stotler (1969) in his revision of the neotropical species of Frullania subgenus Thyopsiella Spruce (as subg. "Frullania") accepted 1 0 species for Colombia and reduced several others to synonymy. Yuzawa (1991) in his world monograph of subgenus Chonanthelia Spruce recorded 19 species from Colombia.

The first edition catalogue of the liverworts and hornworts of Colombia by Gradstein & Hekking (1979) listed 58 species of Frullania for the country. This number has remained almost unaltered in subsequent editions (Uribe & Gradstein, 1998; Gradstein & Uribe, in press) in spite of intensive recent collecting activities and discovery of species new to the country, such as F. kunzei (Pinzon et al., 2003) and F. dulimensis Uribe (Uribe, 2006). Thus, the second edition of the catalogue lists 58 species (Uribe & Gradstein, 1998) and the third edition 59 species (Gradstein & Uribe, in press). The lack of increase in the number of species is explained by the reduction of various species to synonymy, thus counterbalancing the new floristic records. Examples of recent new synonymy include F. atrosanguinea Tayl. (= F. peruviana; Uribe, 2008), F. crinoidea Spruce ex Steph. (= F. atrata; Uribe & Gradstein, 2003), F. mathanii Steph. (= F. peruviana; Uribe, 2008) and F. mucronata (Lehm. & Lindenb.) Lehm. & Lindenb. (= F. brasiliensis; Uribe & Gradstein, 2003).

In this paper, keys are provided for the identification of 42 species of Frullania from Colombia which we have been able to recognize with certainty. A few well-defined species known from neighbouring countries but not yet from Colombia are also included in the keys. For each species from Colombia a brief characterization is given of its distribution, habitat and elevation in the country, its world distribution, as well the main diagnostic morphological characters separating the species from related ones. The remaining species recorded from Colombia are little known taxa that need more study and may turn out to be synonyms; they are listed under "Doubtful taxa" or mentioned briefly in the discussions under the species.

The species are keyed according to the six subgenera to which they are traditionally assigned (Spruce, 1884; Uribe, 2008): subg. Chonanthelia with 17 species from Colombia, subg. Diastaloba with 2 species, subg. Frullania (= Trachycolea) with 1 species, subg. Homotropantha with 1 species, subg. Meteoriopsis with 4 species and subg. Thyopsiella with 17 species. A first morphological-phylogenetic analysis of the genus Frullania (Uribe, 2008) recovered four of these subgenera as monophyletic s.l. (incl. paraphyly; Horandl & Stuessy, 2010): subg. Frullania, subg. Homotropantha, subg. Meteoriopsis and subg. Thyopsiella. The subgenera Chonanthelia and Diastaloba, however, proved to be polyphyletic. Hentschel et al. (2009) in a molecular-phylogenetic study based on a broader sampling confirmed the polyphyly of subg. Diastaloba and the monophyly of subg. Frullania and subg. Homotropantha; in addition, subg. Chonanthelia turned out to be monophyletic in the latter study. Subgenus Thyopsiella, however, was recovered as two poorly supported clades, a "temperate" clade containing the type of subg. Thyopsiella (F. tamarisci) and a few other species, and a "tropical" clade which included the majority of the Thyopsiella species and the type of subg. Meteoriopsis (F. peruviana). Accordingly, the temperate clade was called subg. Thyopsiella and the tropical clade was called subg. Meteoriopsis. In the present treatment, however, we have retained the six subgenera of neotropical Frullania as traditionally circumscribed. The new defintions of Thyopsiella and Meteoriopsis proposed by Hentschel et al. (2009) must be considered premature due to the lack of robust support for the two clades and the very limited sampling of the subgenus Meteoriopsis. A reclassification of the polyphyletic subgenus Diastaloba, finally, should await the urgently needed systematic revision of this group, especially of the neotropical species which have not been studied critically.

DISTRIBUTION PATTERNS

About half of the species of Frullania recorded from Colombia (22) have very wide geographical ranges and occur throughout tropical America, from Mexico to southern Brazil (see below). Six of them also occur in the Old World tropics and are pantropical species. The remaining species (20) are largely restricted to the Andes; some of these extend to the mountains of Central America and Mexico and one species (F. standaertii) occurs also in East Africa. Eight Andean species occur only in the northern Andes, from northern Peru to Venezuela (sometimes also in Costa Rica) and two species (F. bogotensis, F. dulimensis) are only known from Colombia. The geographical distribution patterns of the Colombian species of Frullania are as follows:

Pantropical: apiculata, arecae, ecklonii, ericoides, nodulosa, riojaneirensis

Neotropical: atrata, beyrichiana, brasiliensis, caulisequa, confertiloba, convoluta, cuencensis, dusenii, ecuadorensis, gibbosa, intumenscens, kunzei, macrocephala, montagnei, pittieri, setigera

Andes, sometimes extending to Central America and Mexico: grandifolia, laxiflora, paradoxa, peruviana, pluricarinata, sphaerocephala, standaertii, tetraptera, tunguraguana, winteri

Northern Andes, sometimes extending to costa Rica: bicornistipula, clandestina, formosa, jelskii, lobato-hastata, meridana, mirabilis, ringens

Colombia only: bogotensis, dulimensis

The distribution of the species within Colombia is still incompletely known. The largest number of species is known from Cundinamarca (26 spp.), followed by Boyaca (21 spp.), Risaralda (19 spp.) and Santander (17 spp.). Eight departments have 5-15 species (Casanare, Cesar, Choco, Huila, Magdalena, Meta, Quindio, Tolima, Valle de Cauca), 5 departments have 2-4 species (Amazonas, Caldas, Narino, Norte de Santander, Putumayo) and 12 departments are without any species record (see Table 1). The departmental species richness reflects on the one hand the very uneven collecting intensity in the different departments, on the other hand it shows the typically montane distribution of Frullania in Colombia with more than 95% of the species occurring in the Andean region, above 1000 m.

The elevational distribution of the species diversity of Frullania in Colombia is typically hump-shaped which the highest diversity (75% of the species) occurring at mid elevations, between 1000- 2000 m (Table 2). The elevational distribution pattern of Frullania in Colombia is similar to that of liverworts in general (Uribe & Gradstein. 1999; data from 750 spp.) and apparently reflects the prevalence of epiphytism in liverworts and the preference of liverworts for humid ecological conditions. Similar hump-shaped richness patterns with peaks at mid-elevations have been reported for other plant groups, for example Melastomataceae and ferns, which like liverworts have strong ecological preferences for humid conditions (Kessler, 2000; Kluge et al., 2006). The elevation pattern of species richness detected in Frullania may thus be similar to that of many other plant groups of Colombia.

The number of Frullania species decreases to the paramo (17 spp.) but no species of Frullania is exclusive to the paramo belt even though F. decidua has been found only in paramo at 3800. The latter species occurs in Ecuador at much lower elevation in the forest belt (type location!), indicating that its distribution in Colombia is insufficiently known. Species richness is lowest in the submontane and lowland areas with only 10 and 13 species, respectively. Frullania kunzei and F. nodulosa, are only known from Amazonia below 500 m. The latter two are common and widespread lowland taxa of tropical America and F. nodulosa occurs also in tropical Africa and Asia. A third species recorded from Amazonia, F. caulisequa, is also known from the Andean region where it is reported up to 3000 m. The high-Andean records of this species should be checked, however, because elsewhere in tropical America F. caulisequa is a lowland or lower montane species (Gradstein & Costa, 2003; Leon-Yanez et al., 2006). Possibly, confusion with some other member of the subgenus Diastaloba to which F. caulisequa belongs has occurred. Such confusion may well have happened since the neotropical species of subg. Diastaloba have not been revised and are still very poorly known.

KEYS AND DESCRIPTIONS

Frullania Raddi

Plants small to large, reddish-green to reddish-brown or purplish, sometimes pure green, creeping, ascending or pendent, regularly or irregularly 1-3-pinnately branched. Branches Frullania-type; innovations normally lacking. Stems rigid, made up of thick-walled cells. Leaves incubous with a very short, subtransverse insertion, divided into a large dorsal lobe, a smaller ventral lobule, and a small stylus between lobule and stem. Leaf lobe usually ovate-orbicular, apex rounded to acute to acuminate, margins entire, rarely toothed. Leaf lobule almost free from the dorsal lobe, rounded to elongate, transformed into a sac or flattened, or both. Stylus usually linear and minute, rarely large, foliar. Cells isodiametrical to (more often) elongate, usually with trigones and intermediate thickenings, the trigones often confluent and cell walls often irregularly undulate-sinuose, cuticle smooth; oil bodies finely granular; ocelli sometimes present. Underleaves small or large, 2-lobed, rarely undivided. Rhizoids in bundles from underleaf bases. Dioicous or autoicous, rarely paroicous. Androecia usually on a short, globose male branch with a few pairs of bracts and bracteoles. Gynoecia on short or elongate shoots, each with 2-5 archegonia. Perianths flattened or inflated, with 0-14 keels, the mouth contracted into a beak. Seta of numerous rows of cells. Foot of the sporophyte not penetrating into the stem. Capsule globose, wall 2-layered. Elaters attached to the capsule valves, arranged vertically inside the capsule, with 2-3-spirals. Spores large, multicellular, germination endosporic. Vegetative reproduction rare, by caducous or fragmenting leaves.

Keys to the species of Frullania from Colombia

Some species recorded from Ecuador and/or Venezuela that may occur in Colombia but are not yet known from the country are also included in the key [in brackets].
1. Introductory Key

1. Lobules distant from the stem, space
between stem and lobule wider than the
width of the lobule. Plants very small,
creeping                                       Key 2 (subg. Diastaloba)

1. Lobules close to the stem, space between
stem and lobule narrower than the width of
the lobule. Plants small or large                                     2

2. Lobules pendent, opening of the lobule
positioned toward the stem or the apex
of the plant. Amazonian lowlands (subg.
Homotropantha)                                              F. nodulosa

2. Lobules upright, opening of the lobule
positioned toward the base of the plant                               3

3. Lobules with a long or short flattened lower
portion (flattened portion sometimes very
short!). Ventral surface of perianth with 2-6
keels                                        Key 3 (subg. Chonanthelia)

3. Lobules without flattened lower portion.
Ventral surface of perianths with 0-1(-4)
keels                                                                 4

4. Leaves distinctly squarrose when moist,
brittle. Lobules as long as wide. Surface
of perianth rough by scattered tubercles
or scale-like outgrowths (subg. Frullania)
                                                           F. ericoides

4. Leaves not squarrose, not brittle. Lobules
longer than wide. Surface of perianth
without tubercles or scale-like outgrowths,
smooth                                                                5

5. Plants long-pendent from branches in cloud
forest. Leaf bases with 2 very large auricles,
the auricles similar in size. Leaves convolute
around the stem both when dry and when
moist                                        Key 4 (subg. Meteoriopsis)

5. Plants not long-pendent. Auricles at leaf
bases lacking or not similar in size, the
dorsal auricle distinctly larger than the
ventral auricle. Leaves [+ o -] convolute when
dry, spreading when moist                     Key 5 (subg. Thyopsiella)

Key 2. Frullania subgenus Diastaloba

1. Leaf apex always rounded. Plants creeping,
regularly or irregularly branched                         F. caulisequa

1. Leaf apex at least in some leaves apiculate.
Plants regularly (bi)pinnate, ascending from
the substrate                                                         2

2. Dorsal leaf base distinctly auriculate.
Female bracts strongly toothed. Known
from Ecuador, not yet recorded from
Colombia                                          [F. serrata Gottsche]

2. Dorsal leaf base not auriculate. Female
bracts entire                                              F. apiculata

Key 3. Frullania subgenus Chonanthelia

1. Lobules on the main stem with a very long
laminate (= flattened) portion, which extends
downwards beyond the ventral margin of
the leaf lobe (sect. Cladocarpicae)                                   2

1. Lobules on the main stem with a short
laminate portion, which does not extend
beyond the ventral margin of the leaf lobe
(sect. Chonanthelia)                                                  9

2. Margins of leaves and underleaves strongly
undulate. Perianth 8-10-keeled                                F. arecae

2. Margins not undulate, plane. Perianth 4keeled
or 8-10-keeled                                                        3

3. Stylus large, more than 0.3 mm long,
suborbicular. Perianth 4-keeled                         F. confertiloba

3. Stylus small, less than 0.1 mm long, filiform                      4

4. Laminate portion of the lobule toothed.
Perianth 8-10-keeled                                         F. ringens

4. Laminate portion of the lobule entire                              5

5. Perianth 4-keeled. Lobule with a distinct
beak, formed by the sac extending narrowly
downwards along the free margin of the
laminate portion                                                      6

5. Perianth 8-14-keeled. Lobule with or
without beak                                                          7

6. Free margin of the laminate portion of
the lobule strongly folded below the beak.
Laminate portion of lobule wider than the
sac. Rare species                                         F. bogotensis

6. Free margin of the laminate portion of the
lobule plane, not folded below the beak.
Laminate portion of lobule not wider than
the sac. Very common species                          F. riojaneirensis

7. Lobule with a distinct beak                          F. tunguraguana

7. Lobule without beak                                                8

8. Female bracts and bracteoles densely
imbricate and incurved, forming a globose
head. Stylus minute, 1(-2) cells long.
Dioicous                                              F. sphaerocephala

8. Female bracts and bracteoles spreading,
not forming a globose head. Stylus 2-3 cells
long. Monoicous                                             F. ecklonii

9. Leaf apex acute. Described from Ecuador,
not yet recorded from Colombia                 [F. haematocysta Spruce]

9. Leaf apex broadly rounded                                         10

10. Leaves distinctly squarrose when moist.
Stylus large, rounded                                        F. gibbosa

10. Leaves plane when moist. Stylus filiform
or lanceolate, not rounded                                           11

11. Base of the laminate portion of the lobule
folded upwards                                               F. winteri

11. Base of the laminate portion of the lobule
plane                                                                12

12. Stylus 10-25 cells long and 4-8 cells wide           F. standaertii

12. Stylus less than 10 cells long                                   13

13. Underleaves on main stem maximally 3
times as wide as the stem, distant                                   14

13. Underleaves on main stem more than 3 times
as wide as the stem, distant or imbricate                            16

14. Underleaves more than 2 times wider
than the stem, bifid to 1/3. Lobule with a
beak. Dioicous. Montane forest species                    F. cuencensis

14. Underleaves smaller, 1-2 times wider than
the stem, bifid to 1/6-1/4. Lobule without
beak. Monoicous. Montane forest and
paramo                                                               15

15. Perianth 8-10-keeled. Montane forest               F. pluricarinata

15. Perianth 4-keeled. Paramo                             F. tetraptera

16. Laminate portion of the lobule large, to
ca. 0.85 mm long. Lobule with a distinct
beak, formed by the sac extending narrowly
downwards along the free margin of the
laminate portion                                                     17

16. Laminate portion of the lobule very small,
less than 0.4 mm long. Lobule not beaked                             18

17. Perianth 4-keeled. Underleaves with large
auricles, auricles approaching each other                  F. laxiflora

17. Perianth 8-10-keeled. Underleaves with
small auricles, auricles not approaching each
other. Known from Ecuador, Venezuela and
Costa Rica, not yet recorded from Colombia       [F. planifolia Steph.]

18. Underleaf apex undivided. Underleaf
bases with very large auricles, the auricles
[+ o -] touching each other. Described from             [F. holostipula
Venezuela, not yet recorded from Colombia              Hatt. & Griffin]

18. Underleaf apex short bifid. Underleaf
bases without or with small auricles                                 19

19. Underleaves [+ o -] rounded. Mouth of the
lobule narrow (ca. V the width of the
lobule). Lobule surface folded in the middle                 F. dusenii

19. Underleaves longer than wide. Mouth
of the lobule wide (almost as wide as the
lobule). Lobule surface smooth, not folded                   F. jelskii

Key 4. Frullania subgenus Meteoriopsis

1. Leaf margins toothed                                   F. dulimensis

1. Leaf margins entire                                                2

2. Leaf apex rounded to obtuse. Female bracts
entire (rarely toothed: F. grandifolia)                               3

2. Leaf apex acute to acuminate. Female bracts
toothed                                                               4

3. Underleaves ovate-oblong, 1.4-2 x longer
than wide, not overlapping. Female bracts
entire. Perianth distinctly exserted beyond
the bracts                                                 F. convoluta

3. Underleaves suborbicular, 1-1.3 x longer
than wide, overlapping. Female bracts
toothed. Perianth not exserted beyond the
bracts                                                   F. grandifolia

4. Underleaf base without auricles. Underleaves
narrow elongate, at least 2 x longer
than wide                                                  F. peruviana

4. Underleaf base with very large auricles
(more than 0.15 mm long). Underleaves
less than 2 x longer than wide. Known
from Ecuador and Venezuela; not yet
recorded from Colombia                        [F. phalangiflora Steph.]

Key 5. Frullania subgenus Thyopsiella

1. Leaf apex very long and finely acuminate,
the acuminate portion to 1 mm long. Known
from Ecuador; not yet recorded from
Colombia                                           [F. aculeata Taylor]

1. Leaf apex not very long and finely acuminate                       2

2. Lobules short, 1-1.5 x longer than wide.
Plants very small, 0.5-1 mm wide, tightly
prostrate. Cells in the upper part of the leaf
lobes isodiametrical, evenly thickened,
without trigones. Dorsal leaf base not
auriculate                                                    F. kunzei

2. Lobules 1.5-3 x longer than wide. Plants
larger. Cells in the upper part of the leaf
lobes longer than wide, with trigones.

Dorsal leaf base auriculate                                           3

3. Leaf apex apiculate to acuminate                                   4

3. Leaf apex rounded to subacute (never

apiculate to acuminate)                                              13

4. Underleaves reniform (much wider than
long), underleaf apex broadly truncate and
with 2 small, subulate teeth                          F. bicornistipula

4. Underleaves rounded, ovate to oblong,
not reniform; underleaf apex not broadly
truncate with 2 small teeth                                           5

5. Underleaf apices long acuminate. Lamina
of first branch underleaf undivided, long
acuminate                                                   F. setigera

5. Underleaf apices acute or obtuse. Lamina
of first branch underleaf bifid or undivided,
rounded to subacute                                                   6

6. Leaf apex distinctly recurved, apiculate                           7

6. Leaf apex plane                                                    8

7. Stylus sometimes with a small foliose
appendage at the base, ca. 5-10 cells long
and 3-7 cells wide (underleaves must
be carefully removed to examine this
character). Perianth sharply 3-keeled                    F. intumescens

7. Stylus without foliose appendage. Perianth
terete or with 3 weak, broad keels near the
apex                                                    F. brasiliensis

8. Leaf apex long mucronate to acuminate                    F. pittieri

8. Leaf apex obtuse to acute to short apiculate                       9

9. Underleaves almost as large as the leaf
lobes, 3-4 times as wide as the stem,
imbricate                                                            10

9. Underleaves much smaller than leaf lobes,
2-3 times as wide as the stem, distant or
imbricate                                                            11

10. Underleaves longer than wide, distant                   F. meridana

10. Underleaves rounded or wider than long,
imbricate                                                    F. formosa

11. Perianth terete. Female bracts entire               F. macrocephala

11. Perianth 3-keeled. Female bracts toothed
or entire                                                            12

12. Underleaf margins recurved                                F. atrata

12. Underleaf margins plane       F. beyrichiana (incl. F. breuteliana)

13. Ventral leaf base with 2 long and narrow
appendages, one hanging down and the
other standing up, parallel to the stem                    F. mirabilis

13. Ventral leaf base without long appendages                        14

14. Underleaves with long undulate auricles,
auricles V or more the length of the
underleaf (underleaf length measured from
the middle of the insertion to the apex)              F. lobato-hastata

14. Underleaves without or with short auricles                       15

15. Underleaves about as large as the leaf
lobe, more than 3 times as wide as the
stem, imbricate. Underleaf margins plane                 F. clandestina

15. Underleaves smaller, less than 3 times as
wide as the stem, distant to subimbricate.
Underleaf margins recurved or plane                                  16

16. Plants small, less than 1 mm wide when
moist. Lobules often diverging from the
stem at an oblique angle. Underleaves as
long as wide, deeply bifid (to V), underleaf
margins plane                                           F. ecuadorensis

16. Plants larger. Lobules standing upright,
[+ o -] parallel to the stem. Underleaves longer
than wide, less deeply bifid (to 1/5-1/3)                            17

17. Underleaves imbricate, margins strongly
recurved. Lamina of the first branch
underleaf undivided. Margins of female
bracts and bracteoles entire                               F. montagnei

17. Underleaves distant to subimbricate,
margins plane or slightly recurved. Lamina
of the first branch underleaf bifid. Margins
of female bracts and bracteoles toothed                     F. paradoxa

Alphabetical treatment of the Frullania
species of (Colombia

1. Frullania apiculata (Reinw. et al) Nees

On bark of trees in lowland and montane rain
forest, from sea level to 3000 m. Recorded
from Choco, Quindio and Santander. World
distribution: pantropical.

Frullania apiculata (subg. Diastaloba) is
recognized by the regularly (bi)pinnate
branching and the apiculate leaf apex. The
species is common throughout the tropics
and has been described and illustrated
in detail by Hattori (1973, 1980) for
Asia and by VandenBerghen (1976) for
Africa. Frullania exilis, described from the
Neotropics and recorded from Colombia,
has been considered a synonym of F.
apiculata (e.g. Gradstein & Ilkiu-Borges,
2009) but this synonymy was not based on
examination of type materials and needs
confirmation.

2. Frullania arecae (Spreng.) Gottsche                           Fig. 1

On bark and rock, 1600-3400 m. Recorded
from Boyaca, Casanare, Cauca, Cesar,
Cundinamarca, Huila, Magdalena, Quindio,
Risaralda, Santander and Tolima. World
distribution: pantropical.

Frullania arecae (subg. Chonanthelia) is a
very conspicuous, robust plant with undulate
margins of leaves and underleaves. The
perianth of F. arecae is 10-keeled and is
surrounded by a large and strongly toothed,
flower-like involucre. The species has been
confused with f. ecklonii (Yuzawa, 1991),
but the leaf and underleaf margins in the latter
speciers are not undulate.

3. Frullania atrata (Sw.) Nees                                   Fig. 2

Frullania crinoidea Spruce ex Steph. (syn.
fide Uribe & Gradstein, 2003)

On bark of trees in montane rain forests,
600-3000(-4000) m. Recorded from
Antioquia, Boyaca, Casanare, Cauca, Cesar,
Cundinamarca, Huila, Magdalena, Meta,
Santander and Tolima. World distribution:
tropical America.

Frullania atrata (subg. Thyopsiella) has
often been misidentified and confused with
members of the subgenus Meteoriopsis such
as F. convoluta and F. peruviana. However,
Uribe & Gradstein (2003) showed that F.
atrata is a member of the subgenus Thyopsiella
and possibly related to F. intumescens, which
differs by strongly recurved leaf apices (plane
in F. atrata). Frullania atrata is also close to
F. beyrichiana, which differs from F. atrata
by the plane underleaf margins (recurved in F.
atrata). The relationships of F. atrata to other
members of the subgenus Thyopsiella remain
poorly known and need more study.

4. Frullania beyrichiana (Lehm. & Lindenb.)
Lehm. & Lindenb.

On bark of trees, 1800-2100 m. Recorded
from Antioquia and Cundinamarca. World
distribution: tropical America.

Frullania beyrichiana (subg. Thyopsiella)
is very similar to F. atrata (see under the
latter species). Frullania breuteliana, which
has been reported from Colombia, may be
a synonym of F. beyrichiana and seems to
differ from F. beyrichiana only in the slightly
longer leaftip and the somewhat toothed
female bracts (entire in F. beyrichiana; see
Stotler, 1969).

5. Frullania bicornistipula Spruce

On bark of trees and soil, 1500-2300 m.
Recorded from Cauca, Huila, Norte de
Santander and Risaralda. World distribution:
northern Andes and Costa Rica.

Frullania bicornistipula (subg. Thyopsiella)
is a very characteristic species which is
immediately recognized by the reniform
underleaves with a broad truncate apex
bearing 2 short, subulate teeth. The two small
teeth are standing upwards and separated from
each other by some distance. The long filiform
stylus, 6-12 cells long, is another characteristic
feature of Frullania bicornistipula.

By the long mucronate leaf apex Frullania
bicornistipula is similar to F. pittieri but the
peculiar underleaves of F. bicornistipula
readily separate the two species.

6. Frullania bogotensis Steph.                                   Fig. 1

On logs, 700 m. Recorded from Boyaca,
Cundinamarca and Santander. World
distribution: only known from Colombia.

Frullania bogotensis (subg. Chonanthelia)
is closely related to F. riojaneirensis but
differs from the latter species by the laminate
portion of the lobule being broader than the
sac (narrower than the sac in f. riojaneirensis)
and having a folded margin (Yuzawa, 1991).

7. Frullania brasiliensis Raddi                                  Fig. 3

Frullania mucronata (Lehm. & Lindenb.)
Lehm. & Lindenb. (syn. fide Uribe &
Gradstein, 2003)

[FIGURE 1 OMITTED]

[FIGURE 2 OMITTED]

Common on bark of trees, logs, soil, or rock,
1200-3900 m. Recorded from Antioquia,
Boyaca, Caldas, Cauca, Cesar, Choco,
Cundinamarca, Huila, Magdalena, Meta,
Narino, Quindio, Risaralda, Santander,
Tolima and Valle del Cauca. World
distribution: common and widespread in
tropical America.

Frullania brasiliensis is the most common
neotropical species of subg. Thyopsiella and
is recognized by the recurved, apiculate leaf
apices and the terete perianth. Moreover,
the underleaf margins are always recurved.
According to Stotler (1969), F. brasiliensis
has deeply bifid underleaves (to 1/3),
however in material from Colombia and
Ecuador of F. brasiliensis the underleaf
incision varies considerably in depth (1/51/3).

8. Frullania caulisequa (Nees) Nees

Frullania gymnotis Mont.

Frullania obcordata (Lehm. & Lindenb.)
Lehm. & Lindenb.

On bark and rotten wood in lowland and
montane rainforest areas and woodlands,
from sealevel to about 3000 m. Recorded
from Amazonas, Boyaca, Casanare, Cesar,
Magdalena, Quindio, Risaralda, Santander
and Tolima. World distribution: tropical
America.

Frullania caulisequa (subg. Diastaloba) is
related to F. apiculata but is immediately
separated from the latter by the sparse,
irregular branching and rounded leaftips.
The species occurs on bark of trees at low
elevations. The species was treated by
Schuster (1992) under the names Frullania
obcordata (Lehm. & Lindenb.) Lehm. &
Lindenb. and Frullania gymnotis Mont.,
which are synonyms (see Gradstein &
Costa, 2003). Frullania gymnotis is a
form of F. caulisequa with toothed female
bracts.

9. Frullania clandestina (Nees & Mont.)
Nees

On bark of trees, 1500 m. Recorded from
Cauca. World distribution: northern Andes
(Colombia, Ecuador, Peru).

Frullania clandestina (subg. Thyopsiella) is
a rare species from lower montane elevation
in the northern Andes. The species is
recognized by the rounded leaf apex and the
large underleaves which are about as large
as the leaf lobes and imbricate. By its large
underleaves, F. clandestina resembles F.
formosa Spruce but in the latter species the
leaf apex is apiculate.

10. Frullania confertiloba Steph.                                Fig. 4

On bark of trees and logs, 700-1700 m.
Recorded from Boyaca and Santander.
World distribution: scattered in tropical
America.

Frullania confertiloba (subg. Chonanthelia)
is closely related to F. riojaneirensis but
differs by the large, rounded stylus (Yuzawa,
1991). The species is known from only few
locations in tropical America and may have
been overlooked.

11. Frullania convoluta Lindenb. & Hampe                         Fig. 5

On bark of trees, pendent from branches,
in upper montane cloud forest, to 3400 m.
Recorded from Boyaca, Cundinamarca,
Meta, Narino, Norte de Santander, Putumayo,
Risaralda, Santander and Tolima. World
distribution: Central America, Dominican
Republic, tropical Andes, Guayana
Highlands.

Frullania convoluta (subg. Meteoriopsis) is a
common species of the subgenus Meteriopsis
and is recognized by pendent growth, rounded
(to obtuse) leaf apex and the rather distant,
ovate-oblong underleaves (Uribe, 2008).

[FIGURE 3 OMITTED]

[FIGURE 4 OMITTED]

The species closely resembles Frullania
grandifolia, which is the only other member
of subgenus Meteriopsis with a rounded
leaf apex; for differences between the two
species see the key. Frullania convoluta has
sometimes been confused with F. peruviana,
which is also widespread in tropical America,
but in the latter species the leaf apices are
acute.

12. Frullania cuencensis Tayl.

On bark and logs, to 3000 m. Recorded
from Cundinamarca and Santander. World
distribution: tropical America.

Frullania cuencensis (subg. Chonanthelia) is
recognized by the beaked lobule with a large
sac and short laminate part, and rather deeply
bifid underleaves (to 1/3) which are 2-3 times
wider than the stem. The species has been
confused with F. tetraptera, but the latter
species has narrower and less deeply bifid
underleaves and occurs at higher elevation.

13. Frullania dulimensis Uribe                                   Fig. 6

On bark of isolated trees in open areas in
the upper montane belt, growing together
with other pendent species of the subgenus
Meteriopsis (F. peruviana, F. convoluta),
3200 m. Recorded from Tolima (Uribe, 2006).
World distribution: only known from the type
specimen from Colombia.

Frullania dulimensis (subg. Meteoriopsis)
is readily separated from the other members
of the subgenus Meteoriopsis by the toothed
leaves. The species is only known from a
single collection from Colombia.

14. Frullania dusenii Steph.

On bark and rock, ca. 1700 m. Recorded from
Cesar, Cundinamarca and Magdalena. World
distribution: tropical America.

Frullania dusenii (subg. Chonanthelia) is
recognized by the saccate lobule with a very
short lamina, a narrow mouth (ca. half the
width of the sac) and a plicate surface, and the
rather large, rounded, imbricate underleaves
which are 3-4 times wider than the stem.
According to Yuzawa (1991), the species is
largely restricted to elevations above 2000
m. Frullania dusenii may be confused with
F. gibbosa but the latter species has squarrose
leaves and a large, leaf-like stylus, and occurs
at lower elevations.

15. Frullania ecklonii (Spreng.) Gottsche
et al.

On bark and rock at high elevations,
3000-3500 m. Recorded from Boyaca,
Cundinamarca and Tolima. World
distribution: pantropical.

Frullania ecklonii (subg. Chonanthelia)
is recognized by the 10-keeled perianth,
the smooth margins of leaves, lobules and
underleaves, and the absence of a well-developed
lobule beak. The species has
sometimes been treated as a synonym of
Frullania arecae but in the latter species
the leaf and underleaf margins are undulate
and the lobule is distinctly beaked (Yuzawa,
1991).

16. Frullania ecuadorensis Steph.

On bark of trees and soil, 700-1200 m.
Recorded from Boyaca and Risaralda. World
distribution: scattered in tropical America.

Frullania ecuadorensis (subg. Thyopsiella) is
characterized by the small plant size (width
less than 1 mm), the rounded leaf apex and
the small, distant underleaves which are
hardly wider than the stem, deeply bifid (to
1/2), cuneate at the base and with flat margins.
The lobules in the species are often somewhat
diverging and positioned at an oblique angle
with the stem (Stotler, 1969). Frullania
ecuadorensis has been described based on
sterile material and its affinities remain little
known.

17. Frullania ericoides (Nees) Mont.

Common on bark and rock in rather dry,
open woodlands, from sea level to 2800 m.
Recorded from Antioquia, Boyaca, Casanare,
Choco, Cundinamarca, Quindio, Risaralda,
Santander and Tolima. World distribution:
pantropical.

Frullania ericoides (subg. Frullania) is
one of the most drought-tolerant epiphytic
liverworts and one of the few leafy liverwort
species that occur in the arid lowland belt of
the Galapagos Islands (Gradstein, 2008). The
species is readily recognized by the squarrose
and rather brittle leaves, which reproduce
asexually by leaf fragmentation. The lobules
are highly variable and may be saccate in some
populations and laminate, without formation
of watersac, in others. Frullania ericoides
may be confused with Frullania gibbosa
which also has squarrose leaves and occurs
in the same habitat. The two species can be
readily separated by their different sizes,
F. gibbosa being a larger plant with leafy
stems more than 2 mm wide (1-1.5 mm in F.
ericoides). Moreover, the leaves in F. gibbosa
are not brittle, the lobules are more broadly
attached to the lobe, the perianth surfaces are
smooth and the stylus is large, leaf-like (small
subulate in F. ericoides).

18. Frullania formosa Spruce

On branches of trees in lowland rain forest at
the foot of the Andes, ca. 500 m. Recorded
from Risaralda. World distribution: only
known from Colombia and Ecuador.

[FIGURE 5 OMITTED]

[FIGURE 6 OMITTED]

Frullania formosa (subg. Thyopsiella) is
recognized by the flat and short apiculate leaf
apex, and the large, imbricate underleaves,
which are more than 3 times as wide as the
stem (Stotler, 1969). By the characters of
the underleaves F. formosa is similar to F.
clandestina but in the latter species the leaf
apex is rounded, never apiculate.

19. Frullania gibbosa Nees

Common on bark and rock, usually in rather
dry, open places, 500-3200 m. Recorded from
Antioquia, Boyaca, Caldas, Casanare, Cauca,
Cesar, Cundinamarca, Magdalena, Risaralda,
Santander and Tolima. World distribution:
widespread in tropical America.

Frullania gibbosa (subg. Chonanthelia)
differs from all other members of the subgenus
Chonanthelia by the squarrose leaves and
the large, foliose stylus. The stylus is usually
covered the large and densely imbricate (and
shallowly bifid) underleaves, and can be
observed only by removing the underleaves.
Frullania gibbosa grows in rather open
locations at low elevations, often together with
F. ericoides with which it may be confused.
For differences between the two see under F.
ericoides.

20. Frullania grandifolia Steph.                                 Fig. 7

On bark in upper montane forest, 3400 m.
Recorded from Tolima (Uribe, 2008). World
distribution: rare species, known from a few
localities in Central America (Mexico, Costa
Rica) and the northern Andes (Colombia,
Ecuador).

Frullania grandifolia (subg. Meteoriopsis)
is recognized by the pendent growth, the
rounded leaf apex, the large, imbricate, [+ o -]
rounded underleaves, and the toothed female
bracts. The species is closely related to F.
convoluta but in the latter the underleaves are
distinctly longer than wide and not imbricate,
and the margins of the female bracts are entire
(Uribe, 2008).

21. Frullania intumescens (Lehm. &
Lindenb.) Lehm. & Lindenb.

On bark and rock in montane rain forest areas
and paramo, 1000-3800 m. Recorded from
Antioquia, Boyaca, Cauca, Cesar, Choco,
Cundinamarca, Huila, Magdalena, Quindio,
Risaralda, Santander, Tolima and Valle del
Cauca. World distribution: tropical America.
Frullania intumescens (subg. Thyopsiella)
is similar to F. brasiliensis but differs by
the perianth with 3 sharp keels (terete in F.
brasiliensis). Yuzawa & Koike (1989), in
addition, found that the two species differ
by the presence of a small foliose appendage
at the base of the stylus in F. intumescens
(Yuzawa & Koike, 1989). We have studied
the type material of F. intumescens (Jamaica,
Bancroft s.n., ex hb. Hooker 1833, lectotype
in hb. S [no. B27792]) and can confirm the
observations of Yuzawa & Koike (1989).
However, we have not observed this appendage
in any Colombian specimen identified as F.
intumescens.

Stotler (1969) treated F. intumescens in
a broader sense, including plants with
and without an appendaged stylus in F.
intumescens. According to Yuzawa & Koike
(1989), the plants without appendaged stylus
belong to F. closterantha which was described
from Ecuador by Spruce (1884) and was
treated as a synonym of F. intumescens s.l. by
Stotler (1969). It should be noted, however,
that, Stotler listed eight further species as
synonyms of F. intumescens that were not
studied by Yuzawa and Koike. Three of them
(F. baptistae Gottsche, F. cucullata Lindenb.
& Gottsche, F. longicollis Lindenb. &
Gottsche) are older names than F. closterantha
that might have priority and should therefore
be studied (types should be in W).

22. Frullania jelskii Loitlesb.

This rare species was recorded from
Colombia by Yuzawa (1991) without further
information on habitat and distribution.
World distribution: only known from Peru
(type) and Colombia.

Frullania jelskii (subg. Chonanthelia) is
characterized by the large saccate lobule with
a very short lamina, the broadly recurved leaf
apex, and the large underleaves (ca. 4 times
as wide as the stem). The species is closely
related to F. dusenii but the latter species
is readily separated from F. jelskii by the
narrow mouth of the sac, rounded underleaves
(slightly longer than wide in F jelskii), and the
plicate lobule surface (Yuzawa, 1991).

23. Frullania kunzei (Lehm. & Lindenb.)
Lehm. & Lindenb.

Frullania neesii Lindenb. (fide Gradstein &
Costa, 2003)

On bark in rather open lowland woodlands and
isolated trees, also on rock, 200 m. Recorded
from Amazonas. World distribution: tropical
America.

Frullania kunzei (subg. Thyopsiella) is
readily recognized by the very small plants,
less than 1 mm wide, which are creeping
on bark or rock, sometimes over larger
bryophytes or lichens, and always tighly
appressed to the subtrate. The rather short
lobules (less than 1.5 times longer than
wide), the isodiametrical, evenly thickened
upper leaf cells and the total absence of
auricles at leaf bases, separate F. kunzei
from other neotropical members of the
subgenus Thyopsiella (Schuster, 1992). The
species may be confused with members of
the subgenus Diastaloba (e.g. F. caulisequa)
which resemble F. kunzei by small size and
prostrate growth, but the very different lobule
in the latter species (positioned at some
distance from the stem, not close to the stem)
readily distinguishes them from F. kunzei.
Frullania kunzei is the only heteroicous
member of the genus Frullania in the
Neotropics; the dioicous phenotype of the
species has been described as F. neesii, which
is a synonym (Gradstein & Costa, 2003).

24. Frullania laxiflora Spruce

On bark in montane rain forest areas, 1500-3500
m. Recorded from Cauca and Tolima.
World distribution: scattered in the tropical
Andes (Bolivia to Venezuela) and the
Galapagos Islands.

Frullania laxiflora (subg. Chonanthelia) is
characterized by the strongly concave leaves,
the rather long laminate portion of the lobule,
which is about as long as the sac yet does not
extend beyond the ventral lobe margin, the
decurrent beak of the lobule, and the broad,
rounded underleaves with large basal auricles
(Yuzawa, 1991). The species may be confused
with F. riojaneirensis but in the latter species
the laminate portion of the lobule extends
beyond the ventral lobe margin and the beak
is not decurrent.

25. Frullania lobato-hastata Steph. Fig. 8

Frullania apollinarii Steph., syn. nov.

On bark, 1 800-3 400 m. Colombia:
Cundinamarca, Quindio and Santander. World
distribution: northern Andes (Colombia,
Ecuador).

Frullania lobato-hastata (subg. Thyopsiella)
is a robust plant from high elevation in the
northern Andes. The species is recognized by
the very long, undulate auricules at underleaf
bases, which are at least half as long as the
underleaf lamina (lamina length measured
from the middle of the underleaf insertion
line to apex). The leaf apex is broadly rounded
or, occasionally, short apiculate. The species
resembles F. brasiliensis in the almost terete
perianth but the leaf apex in F. brasiliensis is
always apiculate and the underleaf auricles
are small and not undulate.

Frullania apollinarii Steph., published by
Stephani in the same volume of the Species
Hepaticarum as F. lobato-hastata but 3 month
later, is a synonym of the latter species (see
Uribe, 2004a for description and illustration
of F. apollinarii).

[FIGURE 7 OMITTED]

[FIGURE 8 OMITTED]

26. Frullania macrocephala (Lehm. &
Lindenb.) Lehm. & Lindenb.

On bark of trees and rocks, 1400-2900 m.
Recorded from Boyaca, Cundinamarca,
Quindio, Risaralda and Santander. World
distribution: tropical America.

Frullania macrocephala (subg. Thyopsiella)
closely resembles F. brasiliensis but differs by
the [+ o -] flat leaf apex (recurved in F. brasiliensis;
see Stotler, 1969).

27. Frullania meridana Steph.                                    Fig. 9

On bark of trees, 1500-3200 m. Recorded from
Cundinamarca, Risaralda and Tolima. World
distribution: northern Andes (Venezuela
(type), Colombia).

Frullania meridana (subg. Thyopsiella) is
characterized by the characters given in the
key. The species is known from only few
collections, and only from male plants (Uribe,
2004b).

28. Frullania mirabilis Jack & Steph.

On bark of trees and soil, 1200-2400 m.
Recorded from Antioquia, Cauca, Choco,
Meta, Narino and Valle. World distribution:
northern Andes, Costa Rica.

Frullania mirabilis (subg. Thyopsiella) is
easily recognized by the two long appendices
at the ventral base of the leaf, one standing up
and one hanging down. The species has also
been described under the name F. pendulostyla
Steph. (Stotler, 1969)

29. Frullania montagnei Gottsche

On bark, 1500-2100 m. Recorded from
Magdalena, Santander and Tolima. World
distribution: tropical America.

Frullania montagnei (subg. Thyopsiella)
resembles F. paradoxa by the rounded leaf
apex and the rather narrow underleaves which
are less than 3 times as wide as the stem,
but differs by underleaves being imbricate
and having distinctly recurved margins, the
undivided lamina of the first branch underleaf,
and the entire margins of female bracts and
bracteoles (see Key).

30. Frullania nodulosa (Reinw. et al.)
Gottsche et al.

In crowns of trees in dense lowland rain forest,
and lower down on bark of trees and shrubs in
savannas and other open places, 200-250 m.
Recorded from Amazonas and Vaupes. World
distribution: pantropical.

The small, pendent lobules in Frullania
nodulosa (subg. Homotropantha) readily
separate this species from all other neotropical
members of the genus Frullania. The species
is furthermore characterized by its robust size,
the regularly pinnate to 2-pinnate branching,
the suborbicular leaves with a broadly rounded
apex and the large, imbricate underleaves
with a very shallow incision at the apex and
with large basal auricles (Gradstein & Ilkiu-Borges,
2009). The species is widespread
in the lowlands of Amazonia but has been
surprisingly little collected in Colombia.

31. Frullania paradoxa Lehm. & Lindenb.

On bark of trees, 3000-3700 m. Recorded
from Boyaca, Cauca, Cundinamarca, Quindio,
Risaralda and Valle. World distribution:
tropical Andes.

Frullania paradoxa (subg. Thyopsiella)
is characterized by the rounded leaf apex,
the longer than wide and rather narrow
underleaves which are hardly wider than
the stem, the shallowly bifid lamina (to V)
of the first branch underleaf, and the sharply
3-keeled perianth surrounded by bracts and
bracteoles with ciliate-laciniate margins
(Stotler, 1969). Frullania flexicaulis Spruce,
recorded from Colombia, is very similar to
F. paradoxa and may be a synonym of the
latter. The species differs from F. paradoxa
by the female bract lobes with entire margins
(usually toothed in F. paradoxa) and the
somewhat more deeply bifid lamina (to 1/2)
of the first branch underleaf.

32. Frullania peruviana Gottsche                                 Fig. 10

Frullania mathanii Steph. (syn. fide Uribe,
2008)

Frullania atrata sensu Spruce (non typus)
(fide Uribe, 2008)

On bark of tree branches in montane cloud
forest areas, 1800-3800 m. Recorded
from Antioquia, Boyaca, Caldas, Choco,
Cundinamarca, Magdalena, Narino,
Quindio, Risaralda, Tolima and Valle. World
distribution: Central America, tropical Andes,
Galapagos Islands.

Frullania peruviana (subg. Meteoriopsis) is
recognized by the pendent growth, acute leaf
apices and underleaves without basal auricles
(Uribe, 2008). The species is very common
in the Andean region and has been described
under several different names. Spruce (1884)
in his famous handbook on the liverworts of
the Amazon region and the Andes, described
the species under the name Frullania atrata,
which is a different species, however, and not
related to F. peruviana (see under F. atrata).

33. Frullania pittieri Steph.

Frullania mucronata auct. (non typus) (fide
Uribe & Gradstein, 2003)

On bark of trees, occasionally on shaded, wet
rock, from sea level to 2500 m. Recorded from
Boyaca, Cauca, Choco, Quindio and Risaralda.
World distribution: tropical America.

Frullania pittieri (subg. Thyopsiella) is
recognized by the flat and longly mucronate
to acuminate leaf apices (Stotler, 1969). The
species has sometimes been misidentified
as F. mucronata, which is a synonym of F.
brasiliensis (Uribe & Gradstein, 2003).

34. Frullania pluricarinata Gott.

On bark of trees, 1700-3000 m. Recorded
from Cundinamarca and Magdalena. World
distribution: scattered in the Andes and the
mountains of Central America, from Chile
to Mexico.

Frullania pluricarinata (subg. Chonanthelia)
is very similar to F. tetraptera but differs
by the 8-10-keeled perianth (4-keeled in F.
tetraptera). The two species seem to differ
somewhat in elevational range, F. tetraptera
being most common in paramo whereas F.
pluricarinata seems to occur mainly in the
upper montane belt.

35. Frullania ringens Spruce

On bark of trees, 3400 m. Recorded from
Cauca. World distribution: northern Andes
(Colombia, Ecuador).

Frullania ringens (subg. Chonanthelia) is
recognized by the large, 2-3-toothed laminate
portion of the lobule, the long, the decurrent
beak of the lobule, the large, rounded
underlaves with 2 very large auricles which
are usually overlapping each other, and 10-keeled
perianths (Yuzawa 1991). Frullania
ringens is most closely related to F. ecklonii
but differs from the latter by the lobule with
a long beak and toothed lamina (beak [+ o -]
lacking and lamina entire in F. ecklonii) and
the larger underleaf auricles, which are often
overlapping each other (not overlapping in
F. ecklonii).

36. Frullania riojaneirensis (Raddi)
Angstr.

On bark and rock, 500-3800 m. Recorded
from Antioquia, Boyaca, Caldas, Casanare,
Cauca, Cesar, Choco, Cundinamarca, Huila,
Magdalena, Quindio, Risaralda, Santander
and Tolima. World distribution: pantropical.
Frullania riojaneirensis (subg. Chonanthelia)
is a very common species of lower montane
elevations and is recognized by the large
laminate portion of the lobule (to be examined
on the leaves of the main stem) with a flat
margin and distinct beak, and the 4-keeled
perianth.

[FIGURE 9 OMITTED]

[FIGURE 10 OMITTED]

37. Frullania setigera Steph.                                    Fig. 11

On bark of trees, soil, rocks and logs, 900-3600
m. Recorded from Antioquia, Boyaca,
Cauca, Choco, Huila, Santander and Valle.
World distribution: scattered in tropical
America.

Frullania setigera (subg. Thyopsiella) is
a little-known species that is recognized
by the triangular-ovate underleaves with
narrowly acuminate apices and auriculate
bases, and by the long acuminate apex of
the lamina of the first branch underleaf
(Stotler, 1969). The leaf apex in F. setigera
is apiculate to short acuminate, plane or
recurved, and the perianth is 3-keeled.

38. Frullania sphaerocephala Spruce

On bark of trees in montane cloud forest areas
and paramo, 1700-3900 m. Recorded from
Cundinamarca, Meta, Quindio, Risaralda and
Tolima. World distribution: tropical Andes
from Bolivia to Colombia.

Frullania sphaerocephala (subg.
Chonanthelia) resembles F. ecklonii by the
pluriplicate perianth, the smooth, entire
leaves and underleaves, and the lobule with
a large appendage without distinct beak, but
is easily separated from F. ecklonii by the
large, inflated female bracts and bracteoles,
which are closely imbricate and incurved,
forming a globose head. Morover, F. ecklonii
is monoicous whereas F. sphaerocephala is
dioicous. By its peculiar, ball-shaped female
involucres F. sphaerocephala cannot be
confused with any other neotropical species
of Frullania. Further characteristics of F.
sphaerocephala are the minute, 1(-2)-celled
stylus, the large trigones of the leaf cells and
the large auricles of leaves and underleaves.
The species is characteristic of upper montane
cloud forests and shrubby paramo, above
2500 m.

39. Frullania standaertii Steph.

On bark of trees, 1500-4000 m. Recorded
from Cundinamarca and Santander. World
distribution: in paramos of the tropical Andes
from Colombia to Bolivia; also in East Africa
(Kenya).

Frullania standaertii (subg. Chonanthelia)
is recognized by the short laminate portion
of the lobule and the long, lanceolate stylus
(Yuzawa, 1991). A a long, lanceolate stylus
is also present in F. albertii Steph. and F.
arsenii Steph., which are very similar to F.
standaertii and have been recorded from
Colombia; probably they are synonyms of
Frullania standaertii. Frullania standaertii is
a characteristic species of the paramos of the
tropical Andes; in addition there is one record
from East Africa.

40. Frullania tetraptera Nees & Mont.

On bark of trees, 2000-3800 m. Recorded
from Cundinamarca, Meta and Risaralda.
World distribution: common at high elevations
in Central America and the Andes, from
Mexico to Chile and Argentina.

Frullania tetraptera (subg. Chonanthelia) is
a characteristic species of the paramo and is
recognized by its rather small size (plants ca.
1-1.4 mm wide), the short laminate portion of
the lobule, the very small stylus, the narrow
underleaves that are hardly wider than the
stem, and the monoicous sexuality (Yuzawa,
1991). The species has sometimes been
confused with F. cuencensis but the latter
species has larger underleaves and is dioicous.
Frullania tetraptera is also very similar to
F. pluricarinata, which is a poorly known
species from high elevation in the Andes, but
the latter species has a pluriplicate perianth
(see Key).

[FIGURE 11 OMITTED]

41. Frullania tunguraguana Clark & Frye

On soil, 1600 m. Recorded from Santander.
World distribution: scattered in the tropical
Andes (Venezuela to Argentina) and in
Mexico.

Frullania tunguraguana (subg. Chonanthelia)
is characterized by the pluriplicate perianth
and the distincly beaked lobule with a long
laminate portion (longer than the sac) whose
margins are entire and smooth, not folded
or undulate. Also, the margins of leaves
and underleaves in this species are never
undulate like in the closely related F. arecae.
Frullania tunguraguana somewhat resembles
F. riojaneirensis but in the latter species the
perianths have only 4 keels while those in
F. tunguraguana have more 12-14 keels
(Yuzawa, 1991). Frullania tunguraguana is
known from only very few collections in spite
of its wide neotropical distribution.

42. Frullania winteri Steph.

On bark of trees, 1700-2500 m. Recorded from
Boyaca, Quindio, Risaralda and Santander.
World distribution: northern Andes, Mexico.

Frullania winteri (subg. Chonanthelia) is
recognized by the lobule with a short laminate
portion whose lateral margin is folded and
projecting upwards towards the sac. Two
varieties have been recognized in this species:
var. winteri with a relatively large stylus
(10-13 cells long, 5-10 cells wide) and var.
vanderhammenii (Haarbrink) Yuzawa with a
smaller stylus (5-8 cells long, 1-2 cells wide).
The material from Colombia and Ecuador
belongs to var. vanderhammenii (Yuzawa,
1991). The species is known from only few
collections.


Doubtful taxa

Frullania armata Herz.--Recorded from Cesar and Magdalena

Frullania atropurpurea Steph.--Recorded from Cundinamarca

Frullania crenulifolia Jack & Steph.--Recorded from Antioquia

Frullania granatensis Gottsche--Recorded from Cundinamarca

Frullania guadalupensis Gottsche ex Steph. --Recorded from Magdalena

Frullania harpantha Herz.--Recorded from Cundinamarca

Frullania moritziana Lindenb. & Gott.--Recorded from Risaralda

Frullania occanniensis Steph.--Recorded from Antioquia

Frullania repanda Gottsche--Recorded from Cundinamarca

Frullania sabanetica Gottsche--Recorded from Cundinamarca

Frullania trianae Gottsche--Recorded from Nova Granata, S. L.

Frullania trollii Herz.--Recorded from Cundinamarca

ACKNOWLEDGMENTS

We thank Dr. Lars Hedenas, Stockholm, for the loan of the type material of Frullania intumescens and Mr. Denis Lamy for the permission to reproduce some figures from Cryptogamie, Bryologie.

Recibido: 07/03/2011

Aceptado: 19/08/2011

LITERATURE CITED

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ROBBERT GRADSTEIN

Museum National d'Histoire Naturelle, Dept. Systematique et Evolution, UMS 7205, Case Postale 39, 57 rue Cuvier, 75231 Paris cedex 05, France.

JAIME URIBE-M.

Instituto de Ciencias Naturales, Facultad de Ciencias, Universidad Nacional de Colombia, Apartado 7495, Bogota D.C., Colombia. juribem@bt.unal.edu.co
Table 1. Number off Frullania species recorded
from the Departments of Colombia.

Department           # Frullania species

Amazonas                       3
Antioquia                     10
Arauca                         0
Atlantico                      0
Bolivar                        0
Boyaca                        21
Caldas                         4
Caqueta                        0
Casanare                       6
Cauca                         11
Cesar                          8
Choco                          8
Cordoba                        0
Cundinamarca                  26
Guainia                        0
Guaviare                       0
Huila                          7
La Guajira                     0
Magdalena                     10
Meta                           5
Narino                         4
Norte de Santander             2
Putumayo                       2
Quindio                       13
Risaralda                     19
San Andres y                   0
Providencia
Santander                     17
Sucre                          0
Tolima                        15
Valle del Cauca                7
Vaupes                         0
Vichada                        0

Table 2. Altitudinal distribution of Colombian
Frullania.

Altitudinal belt   Number of species

     0-1000m            15
   1000-2000 m          30
   2000-3000 m          25
   3000-4000 m          20
  above 4000 m           8
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Title Annotation:Botanica-Sistematica
Author:Gradstein, Robbert; Uribe-M., Jaime
Publication:Caldasia
Date:Jul 1, 2011
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