Table Of ContentCaldasia 16(79): 429-438. 1991
A KEY TO THE COLOMBIAN SPECIES
OF HOLOSTIPOUS LEJEUNEACEAE (HEPATICAE)
S. ROB GRADSTEIN
Institute of Systematic Botany, Heidelberglaan 2, 3584 CS Utrecht, The Netherlands (Studies on Colombian
Cryptograms XLII)
Resumen
Se provee lasinopsis y laclave.con entasis en caracteres vegetativos, para 72 especies (en
34 generos) de lejeuneaceae con anfigastros enteros para Colombia y regiones aledanas,
Tarnblen se suministra una breve informaci6n sobre la ecologia y distribuci6n altitudinal de
las especies en Colombia. A la lista de Hepaticas de Colombia (Gradstein y Hekking 1979)
se agregan 10 especies de la flora de Colombia, as! rnismo, 36 nombres de especies
colombianas pasan a ser sin6nimas.
Abstract
A synopsis and key emphasizing vegetative characters are provided for 72 species (in 34
genera) of Lejeuneaceae with undivided underleaves, recorded from Colombia and
neighbouring areas. A brief statement on the altitudinal distribution and on the ecology ofthe
species inColombia is also provided. The checklist ofthe Hepaticae of Colombia (Gradstein
& Hekking 1979) is updated; 10 species are added to the Colombian flora and 36 species
names recorded from Colombia have fallen into synonymy.
Introduction groups: the "Holostipae", characterized by the
presence of undivided underleaves, the "Schi-
The Lejeuneaceae include hundreds of species zostipae", characterized by bifid underleaves,
inabout 80genera world-wide and arethelargest and the "Astipae", which lack underleaves. The
family of the Hepaticae. They are the dominant Holostipae comprise the more robust and often
group of bryophytes in tropical lowland rain brownish or blackish pigmented species, whe-
forest and usually grow on bark of trees or reas the Schizostipae and the Astipae contain
shrubs, or on living leaves. The species vary all the tiny, pale green taxa, including many
considerably in size: insome genera (AphanoJe- epiphylls. The taxonomy of the schizostipous
jeunea, CoJoJejeunea, MicroJejeunea) they are and astipous Lejeuneaceae ismuch more poorly
very tiny and only a few millimeters long, in known than that of the holostipous Lejeunea-
others (Btyoptetis, Thysananthus, Symbiezidium) ceae, and is in need of much work. A key to
they are rather robust and up to adecimeter long these groups would be urgently needed, yet
or more. The taxonomy of the Lejeuneaceae is would be very difficult to make in view of our
usually considered very difficult because of the poor knowledge of the species.
very large number of species and genera and
because of the extensive literature which is not The Key includes all 60 holostipous species (in
easily accessible to the nonspecialist. Although 32 genera) hitherto recorded from Colombia as
a number of revisions and monographs have well as 12 species recorded from neighbouring
appeared inrecent years, there isa lack of good areas. The latter species may be expected to
keys, descriptions, and illustrations. In particu- occur in Colombia as well. The Key has been
lar, there is a need for keys based on vegetative prepared inthecourse ofthe author's monograp-
characters as specimens are often collectec with. hic work onthe Lejeuneaceae for FLORA NEO-
out mature perianths or androecia. The present TROPICA (Gradstein 1987, 1990, in prep.).
paper is aimed at filling this gap inthe literature
by providing a key, based mainly on vegetative The Key emphasizes vegetative characters to
characters, for the Colombian species of Lejeu- facilitate the identification of sterile material
neaceae with entire underleaves. By their under- As much as possible information is included
leaves, Lejeuneaceae can be divided into three on the altitudinal distribution and ecology of the
429
Caldasia Vol. 16, 1991
species. The data on distribution and ecology Lindigianthus cipaconeus (Kruijt &Gradstein 1985).
are based on the literature, on label data of her- Lopholejeunea nigricans, L. subfusca (Schuster 1980,
barium specimens, as well as on the author's Gradstein in prep.).
Marchesinia brachiata, M. robusta (Spruce 1884).
unpublished field observations. The altitudinal
Mastigolejeunea auriculata (inclus. subsp. plicatif7o-
distribution is given according to the altitudinal
ra)(Schuster 1980, Gradstein in prep.).
belts of Prance (1989) and Frahm & Gradstein
Neurolejeunea breuteJii, N. seminervis (Spruce 1884,
(in press), as follows lowlands, up to 300-500
Schuster 1980).
m; submontane, up to 1000-1400 m; lower
Odontolejeunea decemdentata, O.lunulata, 0rhoma-
montane (= subandean), up to 1800-2400 m; lea (Teeuwen 1989).
upper montane (= andean), up to 3000-3400 Schiffneriolejeunea amaz6nica, S. polycarpa (Evans
m; subalpine (= high andean/subparamo), up 1908, Gradstein 1985a).
to the three line; alpine (= paramo). Species Spruceanthus theobromae (Spruce 1884, Gradstein
which occur throughout the lower and upper 1985a).
Stictolejeunea balfourii, S. squamata (Gradstein
montane belts are montane species.
1985b).
Symbiezidium barbif7orum, S. dentatum, S. transver-
Synopsis of Colombian species of holostipous
sale (Gradstein & van Beek 1985).
Lejeuneaceae
Thysananthus amaz6nica, T. pterobryoides (Ful-
The last overview of holostipous Lejeuneaceae ford 1941).
from Colombia was by Gradstein & Hekking Verdoornianthus griffinii (Gradstein 1978).
(1979). Their list included many names which LEJEUNEOIDEAE
are now considered synonyms (see list of Syno-
Amphilejeunea catinulifera, A. Patellifera, A. viri-
nyms at the end of this paper).
dissima (Schuster 1986).
Anoplolejeunea conterts (Evans 1908).
The species and genera of holostipous Lejeunea- Aureolejeunea aurifera, iulvu, A. paramicola, A.
ceae currently known from Colombia are listed quinquecarinata (Schuster 1986, 1987).
below in alphabetical sequence; they belong to Ceratolejeunea globuJifera, C. grandiloba (Grolle
the subfamilies Ptychanthoideae and Lejeuneoi- 1987).
deae. Species new to Colombia (not included Cheilolejeunea fragrantissima (Spruce 1884).
in the catalogue by Gradstein & Hekking 1979) Cyclolejeunea convexistipa (Evans 1904,Grolle 1984).
are in bold face. Species not yet recorded from Cyttolejeunes holostipa, C.saccatiloba, C. venezuelana
(Schuster 1970, 1978, Gradstein & Buskes 1985).-
Colombia but included in the Key are in roman.
Lejeunea ref7existipula (Spruce 1884),
Publications which provide full descriptions of
Lepidolejeunea eluta, spongia (Piippo 1986).
the species are referred to in brackets.
Leucolejeunea unciloba, L. xanthocarpa (Schuster
1980).
PTYCHANTHOIDEAE Luteolejeunea herzogii (Piippo 1986).
Omphalanthus filiformis, O. jackii, O. ovalis, O.
Acanthocoleus aberrans (Kruijt 1988).
platycoleus, wallisii (Gradstein etal. 1981, Gradstein
Acrolejeunea emergens, A. torulosa (Gradstein 1975).
& Buskes 1985).
Archilejeunea auberiana, A. crispistipula, A fusees-
Taxilejeunea sulphurea (Spruce 1884).
cens, A. parvif7ora, A. porelloides (Gradstein &Bus-
xes 1985). ' Key to Colombian species of holostipous
Blepharolejeunea incongrua, Bsaccata, O. securifoJia Lejeuneaceae
(van Siageren & Kruijt 1985).
Brachiolejeunea laxifoJia, B.leiboldiana, B phyllor- 1a. Leaf margin toothed, at least near apex 2
1b. Leaf margin entire 17
hiza (van Siageren 1985, Kruijt & Gradstein 1986).
2a. Stems thinand fragile, pale colored; ventral stem
Bryopteris diffusa, B. filicina (Stotler & Crandall-
surface only 2-3 epidermis cells wide (above or
Stotler 1974).
below the underleaf insertion area) 3
Caudalejeunea lehmanniana (Schuster 1980). 2b. Stems rather thickand rigid,usually brown;ventral
Dicranolejeunea axillaris (Kruijt 1988). stem surface 4 or more cells wide 12
FrulJanoides conicslis, F. densifolia, F. liebmanniana, 3a. Underleaves toothed. Odontolejeunea (Spruce)
F. tristis (van Slageren 1985). Schiffn 4
430
Gradstein: Lejeuneaceae
3b. Underleaves entire 5 12b. Underleaves entire .. 16
4a. Leaf margins revoluted when dry; leaf cells orien- 13a. Stems ± creeping, branching sparse or dichoto-
tated intransverse rows extending from dorsal to mous; underleaf apex recurved, rounded; line of
ventral leaf margin; lobule with (3-)4-6 teeth. On insertion of the underleaf very deeply arched
leaves or twigs in submontane and montane rain (depth of the insertion about half as long as the
forest (rarely lowlands) total underleaf); lobule with 3-4 small teeth; pe-
............................ O. rhomalea (Spruce) Steph. rianth flat, with 2 innovations. On bark or rock at
4b. Leaf margins plane, involuted orirregularly crisped submontane or lower montane elevations
when dry, not revoluted; leaf cells orientated in ................. Marchesinia robusta (Mitt.) Schiffn.
longitudinal rows (as usual in Leujeneaceae) ex- 13b. Stems erect or pendent, pinnately branched or
tending from leaf base to apex; lobule with 2-4 dichotomous; underleaf apex plane, rounded,
teeth. On leaves or bark in lowland and montane truncate or emarginate; underleaf insertion line
forest ...... O. lunulata (Web.) Schiffn. straight or slightly curved; lobule with one tooth
5a. Ocelli present inleaves: scattered, inashort row or with several very large teeth 14
or 1-2 near leaf base 6 14a. Underleaf apex emarginate; leaf cells isodiame-
5b. Ocelli lacking 8 tric, the cell walls uniformily thickened, trigones
6a. Plants pale-green; large rounded gemmae often lacking; lobule with along, curved tooth; perianth
present onleafmargins; perianth strongly toothed, with 2 innovations; plants densely pinnate, den-
without horns. On leaves, bark or rock inlowland droid. Along the Pacific coast (Choco) and locally
andmontane forest atthefootoftheAndes, insubmontane rainforest
....................... Cyclolejeunea convexistipa ..Thysananthus pterobryoides (Spruce) Schiffn.
(Lehm. & Lindenb.) Evans 14b. Underleaf apex rounded to truncate; leaf cells dis-
6b. Plants brown; gemmae lacking; periantn wiinoui tinctly longerthan wide, thewallswith smalltrigones
teeth, the apex with four horn-like extensions and intermediate thickenings; lobule with a
.......... Ceratolejeunea (Spruce) Schiffn. (subg. small tooth or with several very large teeth; pe-
Ceratophora 7 rianth on a short lateral branch without innova-
7a. Plants ca. 2 mm wide; underleaves 3-4x stem tions; plants pinnate or dichotomous. Common in
width. Epiphyte in montane and subalpine rain submontane and lower montane forests ..
forest C. grandi/oba Jack & Steph. ............... Bryopteris (Nees) Lindenb. 15
7b. Plants smaller, 1-1.5 mm wide; underleaves 2x 15a. Branching dichotomous; lobule with several very
stem width. Epiphy1einlower montane rainforest large teeth B.diffusa (Sw.) Nees
............................... C. globulifera Herz. 15b. Branching ± pinnate; lobule with 1small tooth ....
8a. Underleaves very large, 6-1Oxstem width, atapex ............................................B...filicina (Sw.) Nees
short bifid or notched; lobule minute; perianths 16a. Leafapex broadly recurved; cellwalls colorless; epi-
without keels, on short lateral, innovating bran- dermal cells thin-walled; lobule ca. '!3xleaf length.
ches. Widespread epiphyte of lower montane ele- Above 2000 m. on shrubs, tree ferns (8Ie-
vations Taxi/ejeunea sulphurea chnum) and over litter in humid, high montane
(Lehm. & Lindenb.) Schiffn. environments Lindigianthus cipaconeus
8b. Underleaves smaller, apex undivided 9 ............................................. (Gott.) Kruijt & Grads!.
9a. Leaves with (5-)7-25 teeth; underleaves often pro- 16b. Leaf apex plane, cell walls often with blackish pig-
ducing alarge, circular disc formed by coalesced mentation; epidermal cellsthickened; lobulesmaller;
rhizoids; plants often epiphyllous Odonto- usually below 2500 m 17
lejeunea (Spruce) Schiffn 10 17a. Lobule plane, with 2-4 teeth spaced regularly
9b. Leaves with 1-7 teeth; rhizoid disc lacking; not along the free margin; perianth smooth, on an
epiphyllous 11 elongated shoot and with 2 long innovations; epi
10a. Underleaf bases with sharp auricles; lobule with dermal cells scarcely larger than inner stem cells
2-4 teeth. On leaves or bark inlowland and mon- (stem crosssection). Onbarkorrockatsubmontane
tane forest O. lunulata (Web.) Schiffn. and lower montane elevations .
10b. Underleaf bases cuneate, without sharp auricles; ...................Marchesinia brachiata (Sw.) Schiffn.
lobule with 0-1(-2) teeth or reduced. On leaves or 17b. Lobule inflated-rounded, teeth lacking or obscure;
twigs inlowland and lower montane rainforest perianth ciliate-Iaciniate, on a very short branch
.................... O. decemdentata (Spruce) Steph. and with 1 short innovation; epidermal cells dis-
11a. Lobule at least '/2X leaf length; underleaf inser- tinctly larger than inner cells. Along the Pacific
tion straight; leaves strongly falcate. On bark at coast (Choco), on bark in lowland rain forest ........
lower montane elevations, notyet recorded from Symbiezidium dentatum Herz. (=S. transver-
Colombia ..Blepharolejeunea saccata (Steph.) salesubsp. dentatum(Herz.) Gradst, & van Seek)
van Slag. & Kruijt 18a. Leaves with ocelli 19
11b. Lobule smaller, up to 'hx leaf length, sometimes 18b. Ocelli lacking .... . 23
reduced; underleaf insertion arched; leaves notor 19a. Ocelli arranged in a row, forming a vitta (= nerve)
slightly falcate.............................. .. 36 of 7-22 cells long; lobule with a long, 3-5 cells long
12a. Underleaves toothed 13 tooth. Inthe canopy of lowland, Amazonian forest
431
Caldasia Vol. 16, 1991
and inbushes; not yet recorded from Colombia ,., surrounding athicker-walled medulla; lobule with
....... Neuro/ejeunea seminervis (Spruce) Schiffn. 3-10 teeth; perianth with 5-10 keels or plicae, with
19b, Ocelli scattered through the leaf; lobule with a or without innovations" """"""."." """. 26
short tooth ... .......... 20 26a. Plants usually black; innovations present; flagelli-
20a. Ventral stem surface 4 or more cells wide; Frulla- torm shoots lacking, Lowlands and montane "'"
nia-type branching present or lacking """."'.". Frullanoides Raddi 27
Sticto/ejeunea (Spruce) Schiffn. 20 26b, Plants yellowish-brown; innovations lacking; fla-
(Lepidolejeunea eluta(Nees) Schust. might alsokey gelliform shoots common, producing small cadu-
out here. This species has underleaves sha- cous leaves. Lowlands only, rather xerophytic, in
llowly bifid however, up to 0, 1-0,2 of their length; savanna, scrub, and inthe canopy and at the ed-
furthermore, lobules are rather large, ca, Voxlobe ges of rain forest """"""''''''''''''''''''''
length, the leaf apex isacute and thecells are very .""""", Acro/ejeunea (Spruce) Schiffn. 30
thin-walled). 27a. Leaf apex acute-apiculate orobtuse; underleaves
20b. Ventral stem surface 2 cells wide; Frullania-type distinctly auriculate (auricles 0.1-0.3 mm long,
branching lacking .. """" 22 appressed to the stem), the insertion line deeply
21a. Underleaves wider than long, (3)4-10 x stem arched. Common onbarkorrockatmontane eleva-
width; vegetative branches always ofthe Frullania- tions ..""". F. densifolia Raddi
type; plants 1,2-2,5 mmwide, creeping orpendent, 27b. Leaf apex broadly rounded; underleaves not or
when pendent regularly (bi-)pinnate, when cree- weakly auriculate (auricles less than 0.1 mm long),
ping irregularly branched. Common epiphyte in the insertion line straight or curved """ 28
lowland and lower montane rain forests 28a. Underleaves notauriculate; plants 1-1.5mm wide;
S. squamata (Willd. ex Web.) Schiffn. lobule teeth with 4-6 teeth, the small first tooth
21b. Underleaves suborbicular, 2-3 xstem width; vege- situated inthe sinus atthejunction with the ventral
tative branches ofthe Lejeunea-type (rarely aFru- leaf margin; dioicous; male bracts hypostatic. Lo-
IIania-type branch present); plants smaller, 1-1,3 wland epiphyte, along the coast """"""".
mm wide, creeping, irregularly branched. Rare F, corticalis(Lehm. & Lindenb.) van Slag.
species in the shaded understory of lowland rain 28b, Underleaves weakly auriculate; plants 1,5-3 mm
forest (on tree bases, roots, etc.), usually near wide; lobule with 3-9 teeth, no tooth in the sinus;
running water S. balfourii(Mitt.) EW. Jones malebracts epistatic. Submontane andmontane
22a. Stem fragile, the cell walls thin; lobule "normal" in . ",,,. ..."..29
position (opening oriented toward leaf apex), the 29a. Lobule with 3-5 rather large, triangular teeth; dioi-
keel forming an angle of 50-900 with the stem; cous F. liebmanniana (Gott. & Lindenb.) van
ocelli:;: equal insize toordinary leaf cells orsma- Slag.
ller, rather inconspicuous; trigones lacking. Inthe 29b. Lobulewith5-9narrow, sharpteeth; paroicous
canopy of montane rain forest or on shrubs F. tristis (Steph.) van Slag.
.". Lepidolejeunea spongia (Spruce) B, Thiers 30a. Lobule with 3-4 large triangular teeth, thefirst tooth
22b. Stem rigid, the, walls thickened; lobule curved erect """. Acrolejeunea emergens (Mitt.) Steph.
downwards, the keel forming an angle of 120-1400 30b. Lobule with 5-8teeth, thefirst tooth usually outwar-
with the stem; ocelli mostly larger than leaf cells; dly curved """""""" .."""""""".
trigones well-developed. Along the Pacific coast ".".""" .. A. toru/osa (Lehm. & Lindenb.) Schiffn.
(Choc6) in lowland and lower montane forest "" 31a. Ventral epidermal cells not larger than inner stem
Luteo/ejeunea herzogii (Buchloh) Piippo cells (cross section). Plants of lower elevations
23a. Leaf cetls more or less longer than wide, with cor- (below 1500 rn.) . """ .."".32
date trigones having 2 side convex and 1 side 31b. Ventral epidermis cells larger than inner cells. High
concave; leaves when dry convoluted (weakly so or low elevations """"""""""""" 35
in Caudalejeunea, couplet 38) ". """""". 23 32a. Lobule with (1-)2 teeth; cell walls colorless, plants
23b. Leaf cells isodiametric (elongated in Neurolejeu- turning brown; perianth with 4-5 keels, innovations
nea, couplet 41, and Spruceanthus, couplet 54), lacking ." .Schiffneriolejeunea Verd. 32
the trigones various but not cordate; leaves when 32b. Lobule with 0-1teeth; cellwalls ofolder leaves nor-
dry spreading, curved down or folded but not con- mally with dark pigmentation, plants turning black;
voluted. .41 perianth with 3(-8) keels, innovations pre-
24a. Lobule with 3-10 teeth (teeth sometimes inflexed sent "" " 34
and inconspicuous) " ...." 25 33a. Lobule ovate, '13-%x leaf length; perianth with 2
24b. Lobule with 1-2 teeth .... """"""""""""""".31 short, broadly rounded ventral keels; lobule ofinner
25a. Epidermis cells ±thick-walled, surrounding athin- female bract shorter than lobe. Insubmontane to-
walled medulla; lobule with 3-4 teeth; perianth with rest (canopy) and bushes, rare in Colombia
3 keels and 2 innovations; dorsal stem surface """""."""" S,polycarpa (Nees) Gradst.
with a1-4cells high lamellate paraphyllium. Com- 33b. Lobule narrow rectangular, 'I2-3/sxlobe length; pe-
mon at ~igh elevations (2000-3500 m.) on bark or rianth with 2 long, sharp ventral keels; lobule of
rock. Brachiolejeunea laxifolia (Tayl.) Schiffn. inner female bract as long as the lobe, In lowland
25b. Epidermis cells ±thin-walled (especially dorsally), savannas, bushes and tree crowns in Amazonia,
432
Gradstein: Lejeuneaceae
not yet recorded from Colombia . 39b. Leaves squarrose when moist, when dry strongly
........................................ S. amazonlca Gradst. convoluted; Frullania-typebranches usually pre-
34a. Leaf apex rounded orobtuse; underleaves trunca- sent; cell walls with 0-1 intermediate thickenings;
te; female bracts entire. Forming mats on bark in perianth with 3 keels and with innovations. Sub-
the canopy and at the edges of lowland and sub- montane and lower montane (300-2000 m.), on
montane forest and inbushes, up to 1500 m ...... bark or rock inexposed, rather dry habitats .........
.. Mastigolejeunea auriculata (Wils.) Schiffn. (A .. Brachiolejeunea(Spruce) Schiffn. 40
perianth with more than 3 keels is characteristic 40a. Stem dorsally with a 1cell high lamellate paraphy-
for M. auriculata subsp. plicatiflora (Spruce) Ilium; lateral perianth keels rounded, smooth; dioi-
Gradst.; this subspecies is common in Amazonia cous? (males unknown) B.leiboldiana
and should also occur inColombia) (Gott. & Lindenb.) Schiffn.
34b. Leaf apex sharply acute-apiculate; underleaf apex 40b. Paraphyllium lacking; lateral perianth keels sharp,
emarginate; female bracts toothed. Rather long, dentate-ciliate or smooth; autoicous, male bracts
often pendent plants in lowland Amazonian rain on short-specialized branches ..
forest, forming small festoons on branches and .................B.phyllorhiza (Nees) Kruijt & Gradst.
twigs Thysananthus amazonicus 41a. Ventral stem surface 4ormore cells wide (outside
(Spruce) Schiffn. underleaf insertion area) 42
35a. Lobule never reduced, the apex sharply truncate, 41b.Ventral stem surface 2 cells wide 66
with 2 dissimilar teeth: the second tooth large, 42a. Leaf cells small, ca. 10-20 urn, without or with
situated atlobule angle, sharp, pointing outwards, obscure trigones; lobule usually conspicuously
the first tooth smaller, incurved and blunt. High darker than lobe, bottle-shaped, with along, sharp
montane, 2000-4000 m., on bark or rock inexpo- tooth (usually incurved and invisible without dis-
sed habitats ........ Blepharolejeunea S. Am. 35 section); perianth keels atapex expanded into au-
35b. Lobule reduced or well-developed, the apex ricles; plants small, lessthan 1,5mm wide, usually
rounded or oblique, the teeth equal or the first blanckish. Lowlands to3000 m.,on bark orrock.
tooth larger than the second tooth. Lowland or ............. Neurolejeunea breutelii (Gott.) Evans
montane 37 42b. Leaf cells larger, usually with distinct trigones; lo-
36a. Lobule bordered by enlarged margin cells; pe- bule not conspicuously darker than lobe; perianth
rianth up to 1 mm long, the lateral keels sharp, keels not expanded into auricles 43
entire or ciliate; lobule of female bract small, less 43a. Lobule with 204 teeth 44
than 'lax lobe length . 43b. Lobule with only one tooth or tooth iacking 48
.........................B.incongrua (Iindben. & Gott.) 44a. Lobule well-developed, the apex truncate, with 2
van Slag. & Kruijt dissimilar teeth, the second tooth larger than the
36b. Lobule margin cells not enlarged; perianth over 1 first tooth: first tooth incurved and blunt, second
mm long, the lateral keels broadly rounded, entire; tooth situated at lobule angle, long and sharp. 45
lobule of female bract larger, V2-%X lobe length. 44b. Lobule reduced orwell-developed, the apex roun-
Above 3000 m. only B.securifolia ded oblique, the teeth identical or the first tooth
(Spruce) Schust. larger than the second tooth .. 46
37a. Ventral stem surface 2 epidermis cells wide (out- 45a. Plants whitish to pale grayish-green; lobule about
side leaf insertion area) 38 2x longer than wide; Frullania-typebranches lac-
37b. Ventral stem surface 4epidermis cells wide . 39 king. Lower montane, upto 2500 m
38a. Branches predominantly Frullania-type,plants of- ..Leucolejeunea unci/oba (Lindenb.) Evans 48
ten pendent when growing on bark; epidermis cells 45"!J.Plants darker, greenish-brown to dark brown; lo-
thinwalled, bulging outwards; female bracteole toot- bule 1-1.5x longer than wide; Frullania-type'bran-
hed. Common at submontane and montane ches present. Montane-alpine, 2000-4000 m .....
elevations, on bark, rock or soil inopen habitats . ............................ Blepharolejeunea S. Am. 35
....................................Dicranolejeunea axillaris 46a. Lobules sometimes reduced, when well-develo-
(Nees & Mont.) Schiffn. ped rectangular, with 2teeth atapex; underleaves
38b. Branches predominantly Lejeunea-type,plants al- 3-4,5 xstem stem width, the insertion linestraight;
ways creeping; epidermis cells ± thick-walled, not plants 1-2mm wide, greenish to yellowish-brown;
bulging outwards (Stem surface smooth); female perianth infalted, with 2 ventral keels. Lowland
bracteole entire. Submontane and lower montane, and submontane, notyet recorded from Colombia
on bark orrock, inopen habitats andtree crowns . (0-1000 m) .
.....................................Acanthocoleus aberrans ..Archilejeunea auberiana (Mont.) Evans (x A.
(Lindenb. & Gott.) Kruijt parviflora var. florentissima (Spruce) Gradst. &
39a. Leaves not squarrose when moist; Frullania-type- Buskes)
branches lacking; intermediate thickenings 1-3 per 46b. Lobules never reduced, ovate-trapezoidal, with 2-
cell wall; perianth with 4-5 keels, lacking innova- 4 teeth rsqularly spaced along the free margin;
tions. In lowlands on leaves or twigs in exposed underleaves 4-10 xstem, the line ofinsertion dee-
habitats Caudalejeunea lehmanniana plyarched; plants rather robust, usually more than
(Gott.) Evans 2mm wide, often blackish; perianth flattened, wit-
433
Caldasia Vol. 16, 1991
houtventral keels. Submontane andmontane, lac- often truncate and connate with the lobe surface
king in lowlands, on bark or rock . across 2-3 cells. Plants growing indense, appres-
.................................... Marchesinia S. Gray 47 sed mats in exposed, often rather dry habitats,
47a. Underleaf margin entire M. brachiata usually lowland-submontane .
(Sw.) Schiffn. .. L.subfusca (Nees) Schiffn.
47b. Underleaf margin dentate M. robusta 54a. Leaf cells with many intermediate thickenings (1-3
(Mitt.) Schiffn. percell), trigones radiate; hyaline papilla proximal
48a. Plants whitish or pale yellowish, growing appres- of the apical tooth of the lobule; male bracteoles
sed. Submontane-Iower montane, on bark or rock present throughout the male spike. Inlowland fo-
in rather exposed habitats . rests 55
.....................................Lecolejeunea.Evans 49 54b. Leaf cells without or with very few intermediate
48b. Plants darker, green, brown orblackish. Lowlands thickenings (0-1 per cell), trigones usually large
to alpine elavations, growing appressed or free and bulging, not radiate; hyaline papilla situated
from thesubstrate insheltered orexposed habitats in a small sinus (or on the inner lobule surface)
........................................................................ 50 distal of the apical tooth; male bracteoles limited
49a. Ventral leaf margin strongly involuted; lobule tooth to the base of the spike. Inmountain forests and
1-4 cells long L.xanthocarpa. paramos :........ 60
(Lehm. & Lindenb.) 55a. All or at least some lobules poorly developed; in-
Evans novations lejeuneoid (first leaf at the base of the
49b. Ventral leaf margin plane or slightly upcurved; lo- innovation isa lateral leaf) 56
bule tooth 3-7 cells long L. unciloba. 55b. All lobules well-developed, never tending to be-
(Lindenb.) Evans come reduced; innovations pycnolejeuneoid (first
50a. Epidermis cells larger than inner stem cells (cross leaf at the base of the innovation is an underleaf)
section) ;plants often blackish; perianth keels den- or lacking : 57
tate-Iaciniate 51 56a. Leaf cells distinctly elongated, about 2x longer
SOb.Epidermis cells not larger than inner cells; plants than wide; plants robust, ca. 3 mm wide; ventral
not blackish (except sometimes Archilej. Parviflo- stem surface ca. 12 cells wide. Endemic to the
ra, couplet 54); perianth keels smooth or denticu- Pacific coast of Ecuador, not yet recorded from
late 54 Colombia .
51a. Plants 2-5 mmwide; underleaves very large, 5-1Ox ......Spruceanthus theobromae(Spruce) Grads!.
stem width; lobule always small, lessthan 'l4xleaf 56b. Leaf cells isodiamemc: plants smaller, 1-2 mm
length, strongly infalted; perianth on a very short wide; ventral stem surface 4-6 cells wide. On bark
shoot (appearing lateral on the stem), with one or rock in moist lowland and submontane forests
short innovation. Lowlands to montane, up to ca. (0-1500 m.), always inshaded habitats .
2500 m : SymbiezidiuffJ Trevis 52 .. Archilejeunea parviflora (Nees) Gradst,
51b. Plants smaller, 1-2 mm wide; underleaves 2-5x 57a. Plants glossy brown; leaves when moist widely
stem width; lobule '/3-'/2X leaf length or reduced, spreading, notsquarrose; free margin ofthe lobule
inflated or flattenede; perianth on an elongated plane; innovations present; dioicous. Epiphytes in
shoot, usually without innovations. Lowlands to Amazonian lowland forests ..
lower montane, up to ca. 2000 m . ............................A.rchilejeunea (Spruce) Schiffn.
................. Lopholeujenea (Spruce) Schiffn. 53 (subgen. Archilej.) 57
52a. Plants 2-2.5(-3) mm wide; ventral surface of pe- 57b. Plants dull brown; leaves when moist suberect,
rianth rough due to spines or laciniae, which are not spreading widely, ± squarrose; lobule rectan-
distributed randomly aswell as (sometimes) in1-2 gular, the free margin partly incurved; innovations
rows. Sealevel to 2500 m., on bark in forests .... lacking; autoicous. Rare canopy epiphyte of Ama-
............................. S. barbi"orum (Gott.) Evans zonian lowland rain forest, not yet recorded from
52b. Plants 2.5-5 mm wide; ventral surface of perianth Colombia ........ Verdoornianthus griffinii Grads!.
entirely smooth or with afew spines and laciniae 58a. Underleaf margins undulate; ventral leaf margin
arranged in1-2rows (never randomly distributed). auriculate at the junction wit the keel. Epiphyte in
Distribution as S.barbiflorum but usually at lower periodically inundated Amazonian forest, not re
altitudes (not found above 1700 m.) .. corded from Colombia , , .
............................. S. transversale (Sw.) Trevis. .. A.crispistipula (Spruce) Steph.
53a. Leaf apex acute torounded; perianth long, distinc- 58b. Underleaf margins plane; ventral leaf margin not
tly exserted, weakly toothed-Iaciniate only; apex auriculate 59
of lobule acute, not connate with leaf surface. 59a. Plants robust, 2-3 mm wide; underleaves longer
Plants growing in loose mats in rather moist, sha- than wide to orbicular; lobule apex triangular, lac-
ded habitats, usually submontane-Iower montane king a distinct tooth. Epiphyte in Amazonian rain
........................... L.muelleriana (Gott.) Schiffn. forest andscrub. A. porelloides (Spruce) Schiffn.
53b. Leaf apex rounded; perianth short, hidden behind 59b. Plants smaller, 1-2 mm wide; underleaves wider
a very large bracteole, with very large laciniae than long or (in small plants) orbicular: lobule apex
emerging beyond the bracteole; apex of lobule with 1tooth. Epiphyte in lowland rain forest .
434
Gradstein: Leieuneaceae
.... A. fuscescens (Hampe) Fulf. (x A.juliformis 67b. Frullania-type branches entirely lacking; lobules
(Nees) Grads!. with 1 tooth only; male bracteoles limited to the
6Oa.Lobules quadrate-trapezioid, %-V3x lobe length; base of the spike; plants pale green, rarely brow-
plants usually pendent, long and little branched, nish 68
green or pale yellowish-brown in color; perianth 68a. Lobule large, ¥S-1f2xlobe length (or more) 69
with 0-5 keels Omphalanthus Nees 61 68b. Lobule small, less than '/3X lobe length 73
60b. Lobules oblong-rectangular, %-V2X lobe length; 69a. Underleaves large, 4-5x stem width, imbricate;
plants creeping or ± ascending, with short, irregu- plants rather robust, ca. 1-2 mm wide. Mostly
larly branched stems, brownish in color (golden above 2000 m 69
green to light brown in A. fulva, couplet 82); pe- 69b. Underleaves smaller, 1-2.5x stem width; plants
rianth with 3-10 keels. Aureolejeunea Schus!. 63 tiny and fragile, 0,5-1 mm wide. Submontane-
61a. Leaf apex acute to acuminate' underleaves usua- montane Cyrtolejeunea Evans 71
lly longer than wide' perianth with 5 keels and a 70a. Plants brownish; leaf cells with large, bulging trigo-
long (8 cells) beak 62 nes, intermediate thickenings lacking; stems rigid,
61b. Leafapex broader, rounded tosubobtuse; underlea- cells strongly thickened ....Aureolejeunea p.p. 65
ves wider than long (rarely longer than wide); pe- 70b. Plants pale greenish; leaf cells with small trigones
rianth with orwithout keels, beak shorter orlacking. and intermediate thickenings; stem cells ± thin-
Very common inthe Andes O. filiformis walled. Epiphyte in montane rain forest (canopy)
(Sw.) Nees (the perianth of 0. filiformis has no and bushes Amphilejeunea viridissima
keels; there are two species which are very similar Schus!. (twofurther species, A.catinulffera (Spruce)
to 0. filiformis but with different perianths: 0. pla- Schus!. and A. patellifera (Spruce) Schust.
tycoleus Herz., which has a 3-4-keeled perianth, from the Andes of Ecuador, have been placed in
and 0. wallisii (Steph.) Gradst., which has a5-kee- Amphilejeunea (Schuster 1986). The differences
led perianth. The status of the latter two species among the three species of Amphilejeunea are
is not clear). notclear, however, and need further study).
62a. Leaf apex (sub)acute; underleaves undivided. Lo- 71a. Lobule with a long acuminate tooth; leaves sube
wer montane O. ovalis (Lindenb. & Gott.) -rect; plants very small, Microlejeunea-Iike, ca. 0,5
Grads!. mm wide. On bark in rain forest ....... C. holostipa
62b. Leaf apex acuminate; underleaves short bifid. (Spruce) Evans
Rare, not yet recorded from Colombia ...O.jackii 71b. Lobule tooth short or lacking; leaves spreading;
(Steph.) Grads!. plants larger 72
63a. Leafcellsstrongly mamillose-papillose; leafmargins 72a. Lobule tooth short, blunt; underleaves short bifid;
crenulate. On bark inparamo . dioicous? Venezuela (Tachira), 500 m, not yet re-
............................ Aureolejeunea aurifera Schus!. corded from Colombia ... C. Venezuelana Schust.
63b. Leaf cells ± smooth; leaf margins entire 64 72b. Lobule tooth lacking; underleaves entire; autoi-
64a. Perianth pluriplicate, with 3-5 ventral keels; inno cous. Not yet recorded from Colombia ..
vations 2; plants rather robust, ascending from the .............................. C. saccatiloba(Steph.) Gradst.
substrate or pendent, Epiphyte, above 3000 m .... 73a. Plants firm, the cells of leaves and underleaves
..A. paramicola(Herz.) Schus!. (= Omphalanthus with large trigones; hyaline papilla of the lobule
paramfcola (Herz.) Grads!. distal of the apical tooth; underleaf apex undivided
64b. Perianth with only 1-2 ventral keels; innovations or notched. Epiphyte of submontane and lower
1-2; plants smaller, creeping 65 montane elevations ..
65a. Perianth with 5 long, sharp keels, the perianth ..Cheilolejeunea fragrantissima(Spruce) scbust.
apex rounded (beak not recessed); plants reddish- 73b. Plants delicate, the cells thin-walled with very
to dark-brown. On shrubs at the forest line and in small trigones; hyaline papilla proximal of the api-
the paramo A. quinquecarinata Schus!. cal tooth 74
65b. Perianth near apex with 304 short, broadly roun- 74a. Underleaf apex recurved. On bark inlowland and
ded keels, the perianth apex emarginate, the beak submontane rain forest ..
recessed; plants golden green to light brown. On ...... Lejeunea refltJxistipula(Lehm. & Lindenb.)
canopy branches in montane rainforest .. Gott. et al.
............~......... A. fulva. Schus!. 74b. Underleaf apex plane 75
66a. Lobules uniformily swollen and strongly involute, 75a. Underleaf apex short bifid or notched (underlea-
the free margin imolled 2-3 times. Plants small, ves very large); plants pale green; leaf apex roun-
pale whitish-green. Common epiphyte inforest ca- ded or pointed; ocelli lacking ....... Taxilejeunea
nopy and inscrub on twigs, ca. 500-3500 m ........ sulphurea 8
....... Anoplolejeunea conferta (Meissn.) Schiffn. 75b. Underleaf apex undivided; plants olive-green to
66b. Lobule not oronly weakly involute 67 brownish; leaf apex rounded; ocelli present inlea-
67a. Frullania-type branches present (though sometimes ves and underleaves, often rather similar to the
only few); lobule when well-developed with green cells insize and shape and therefore obs-
2 teeth; male bracteoles present throughout the cure in herbarium material ..
male spike; plants ± brownish pigmented ....... 38 .............................. Lepidolejeunea spongia 22
435
Caldasia Vol. 16, 1991
Synonyms M. plieatiflora (Spruce) Steph. =M. aurieulata
(Wils.) Schiffn. subsp. plieatiflora (Spruce) Gradst.
This list includes names of holostipous Lejeuneaceae
Odontolejeunea calcarea (Spruce) Steph. =0. lunu-
inthe checklist of Colombian liverworts (Gradstein &
lata (Web.) Schiffn.
Hekking 1979), which arenowconsidered synonyms.
O. longispiea Evans =0. deeemdentata (Spruce)
Steph.
Archilejeunea julifonnis (Nees) Gradst. =A. fusees-
O. obversiloba Herz. =O.rhomalea (Spruce) Steph.
eens (Hampe) Fulf.
O. sieberiana (Gott.) Schiffn. =0. lunulata (Web.)
A. leprieurii (Mont.) Schiffn. =A. parviflora (Nees)
Schiffn.
Gradst.
Omphalanthus grandistipulus Steph. =0. filifonnis
Braehiolejeuneaeolumbiea Steph. =Frullanoidesdensifo-
(Sw.) Nees.
liaRaddi. Pyenolejeunea granatensis Steph. =Lepidolejeunea
B. densifo/ia (Raddi) Evans =Frullanoides densifolia
eluta (Nees) GroUe.
Raddi.
Stietolejeunea kunzeana (Gott.) Schiffn. =S. squa-
B.nitidiuseula(Gott.)Schiffn. =Blepharolejeuneaincon-
mata (Willd. ex Web.) Schiffn.
grua(Lindenb. & Gott.) van Slag. &Kruijt.
Symbiezidium granulatum (Nees) Steph. =S. trans-
B. paramieola Herz. =Aureolejeunea paramicola
versale (Sw.) Trevis.
(Herz.) Sehust.
S. subrotundum (Kunth) Trevis. =S. transversale
B. rupestris (Gott.) Steph. =Frul/anoides desinfolia
(Sw.) Trevis.
Raddi B. sueeisa Steph. =Frul/anoides desinfolia
Raddi
B. wrightii Steph. =Frul/anoides tristis (Steph.) van
Literature cited
Slag.
Bryopteris trinitensis (Lehm. & Lindenb.) Lehm. &
EVANS,A.W.1904. Hepaticae of Puerto Rico IV.Odontole-
Lindenb. =B. filieina (Sw.) Nees. jeunea, Cyclolejeuneaand PrionoJejeunea.Bull.To-
B. ttuticuloss Tayl. =B. filicina (Sw) Nees. "ey Bot.Club31: 183-226.
Dieranolejeunea eipaeonea (Gott.) Steph. = Lindi- -- 1908. Hepaticae ofPuerto RicoIX.Brachiolejeunea,
gianthus eipaeoneus (Gott.) Kruijt & Gradst. , Ptychocoleus, ArchiJejeunea, Leucolejeunea and
D. gigantea Steph. =Frul/anoides liebmanniana (Lin- Anoplolejeunea.Bull.TorreyBot.Club35:155-179.
denbo & Gott.) van Slag. FRAHMJ,-P.&GRADSTEINS,.R.inpress:Analtitudinalzona-
b. grossiloba Steph. =Odontelejeunea lunulata tionofthetropical rainforest using bryophytes. Jour-
nalofBiogeography.
(Web.) Schiffn.
FULFORDM, .H. 1941. Studies on American Hepaticae I.
D. ineongrua (Lindenb. &Gott.) Steph. =Blepharo-
Revision of the genus Thysananthus. Bull. Twey
lejeunea ineongrua (Lindenb. & Gott.) van Slag. & Bot.Club68:32-42.
'Kor.uijt. GRADSTEINS,.R. 1975. A taxonomic monograph of the
loxensis (Gott.) Steph. =Aeanthoeoleus aberrans genus Acrolejeunea. Bryophytorum BibJiotheca4:
(Gatt.) Kruijt. 1-162.
D. phyl/orhiza (Nees) Schiffn. =Braehiolejeunea -- 1978. Verdoornianthus, a new genus from Ama-
phyl/orhiza (Nees) Kruijt & Gradst. zonas, Brasil. The Bryologist 80: 606-611.
D. tridentata S. Winkler =D. axillaris (Nees & --1985a. Aguide tothe holostipous Lejeuneaceae.
Beiheft Nova Hedwigia 80: 13-29.
Mont.) Schiffn.
-- 1985b. A revision of the genus Stictolejeunea
Hygrolejeunea patel/if era(Spruce) Steph. =Amphile-
(Spruce) Schiffn. Beiheft zur Nova Hedwigia 80:
jeunea patel/if era (Spruce) Schust.
195-220.
H. reflexistipula (Lehm. & Lindenb.) Steph. =L. --1987. The Ptychanthoideae of Latin America: an
reflexistipula Lehm. & Lindenb. overview. The Bryologist 90: 337-343.
Lopholejeunea muelleriana (Gott.) Schiffn. =L. ni- -- 1989. A key to the Hepaticae and Anthocerotae
grieans (Lindenb.) Schiffn. of Puerto Rico and the Virgin islands. TheBryo-
L. saxatilis (Gott. ex Aongstr.) Steph. =L. nigrieans logist 92: 329-348.
(Lindenb.) Schiffn. --1990. A Key to the New World species of holos-
tipous Lejeuneaceae. Trop. Bryol. 3: 45-57.
Marehesinia aguatiea Herz. =M. braehiata (Sw.)
--In prep.: Lejeuneaceae subfamily Ptychanthoi-
Schiffn.
deae. Flora Neotr6pica Monograph.
M. extensa (Steph.) =M. braehiata (Sw.) Schiffn.
-- MATSUDA,S.R. & A SAKAWA,Y. 1981. Oil bodies
M. longirostris Herz. =M. braehiata (Sw.) Schiffn. and terpenoids inLejeuneaceae and other selec-
Mastigolejeunea pittieri Steph. =M. aurieulata ted Hepaticae. Journal of the Hattori Botanical
rwus.: Schiffn. Laboratory 50: 231-248.
436
Gradstein: Lejeuneaceae
-- &V~ SEEK,J.1985. Arevision ofthe genus Sym- SlAGEREN,M.W.S.J.M. VAN1985. A taxonomic mono-
biezidium Trevis. Beiheft zur Nova Hedwigia 80: graph of the genera. Brachiolejeunea and Frulta-
221-248. noides. Medellingen van het Botanisch Museum
-- & SUSKES,C.M.C. 1985. A revision of neotropical en Herbarium Utrecht 544: 1-309.
Archilejeunea (Spruce) Schiffn. Beiheft zur Nova -- &KRUIJT,R.Ch. 1985. Areview ofthegenus Blep-
Hedwigia 80: 89-112. harolejeunea S. Am. Beiheft zur Nova Hedwigia
GROllE, R. 1984: Zur Kenntnis der Lejeuneoideae in 80: 113-145.
Cuba (1): Cyclolejeunea. Wissenschafteliche ScHUSTERR,.M. 1970. New Lejeuneaceae from Dominica
Zeitschrift der Friedrich-Schiller-Universitat Jena, andJamaica. Bull. Torrey Bot. Club 97:336-352.
Matematisch-Naturwissenschaftliche Reihe 33: -- 1978. Studies onVenezuelan Hepaticae I,II.Phy-
759-764. tologia 39: 239-251.
--1987.Ceratolejeunea subg. Ceratophora Schust. -- 1980. The Hepaticae and Anthocerotae of North
In: W. Schultze-Motel & M. Menzel, Die Leber- America, Vol. IV.Columbia University Press, New
moose im Bryotrop-Transekt von Peru. Beiheft York.
zur Nova Hedwigia 80: 91-92. --1986. Studies on Venezuelan Hepaticae III. On
KRUIJT,S.R. 1988. Amonograph ofthegenera Dicrsno- Amphilejeunea Schust. and Aureolejeunea
lejeunea and Acanthocoleus. Bryophytorum Bi- Schust. Nova Hedwigia 44: 1-23.
bliotheca 36: 1-135. --1987. OnAureolejeunea Schust. and Brachiolejeu-
-- &GRADSTEINS,.R.1986. On Brachiolejeunea phy- nea paramicola Herzog. Phytologia 61: 445-447.
Itorhiza (Nees) Kruijt & Gradstein comb. nov. SPRUCE,R. 1884. Hepaticae Amazonicae et Andinae.
Nova Hedwigia 43: 299-309. Jubuleae. Transactions and Proceedings of the
PIIPPO,S. 1986 A monograph of the genera Lepidole- Botanical Society of Edinburgh 15: 1-308.
jeunea andLuteolejeunea. Acta Botanica Fennica STOTLER,R. & GRANDAll-STOTlER, S. 1974. A mono-
132: 1-69. graph ofthe genus Bryopteris. Bryophytorum Bi-
PRANCE,G.T. 1989. Tropical American Forests. In: H. bliotheca 3: 1-159.
Lieth &M.J.A. Werger (eds.), Tropical Rain Forest TEEuwEN,M. 1989. Arevision ofthegenus Odontolejeu-
Ecosystems. Elsevier, Amsterdam. nea. Nova Hedwigia 48: 1-32.
437
