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Botanical References
to Páramo

      Richard Spruce, the great 19th century explorer of the Andes and author of the classic Hepaticae Amazonicae et Andinae (Spruce 1884-85), was one of the first to collect hepatics in páramos.  Surprisingly, he considered them quite poor in species (Spruce, 1886: 77): "La zone alpine des Andes est aussi pauvre en hépatiques que celle des Pyrenées...."  Spruce's negative impression of hepatic diversity in the páramos was probably due to the fact that he only visited the rather dry, species-poor páramos of central Ecuador (Pichincha and Chimborazo provinces).  His appreciation of the hepatic flora would undoubtedly have been different if he had visited the species-rich, humid páramos of northern Ecuador, Colombia, or Venezuela. 

      The most important hepatic collections made in páramos are those made by Antoine M. Cleef in Colombia during 1970-72 (Gradstein, 1982, 1983, 1990; Gradstein et al., 1977; Gradstein & Hekking, 1979) and by Rudolf M. Schuster in Venezuela in 1976 (Schuster, 1978, 1978b, 1985, 1986, 1987, 1991, 1995).  Other important collectors include G. Wallis, H. Bischler, J. Cuatrecasas, J. Aguirre, K. Mägdefrau, S. Winkler (Colombia), M. Allioni, E. Asplund, G. Harling, W. Jameson, H. Meyer (Ecuador), D. Griffin, J. Steyermark (Venezuela), G. Dauphin, N. Salazar Allen (Panama), A. M. Cleef et al., and G. Dauphin (Costa Rica).  The author has collected hepatics in páramos in Costa Rica (years 1993, 1994), Colombia (years 1980, 1982 1984, 1992), Ecuador (years 1976, 1988), and Peru (year 1982). 

      At present 291 species of hepatics, in 88 genera and 34 families, are known from the neotropical páramos.  Accepting about 1250 hepatic species in tropical America, in 189 genera (Salazar Allen et al., 1996), it appears that páramos harbour more than one-fifth of the neotropical species and almost half of the genera.  Taking into account the relatively small area occupied by the páramos, these figures indicate a very high diversity.  Another striking feature of the hepatic flora of the páramos is the high degree of endemism.  Seven genera and subgenera and about 70 species are exclusively known from páramos, accounting for 26.5% of the hepatic flora (Gradstein, 1998). 

      The principal hepatic groups of the páramos are listed in Table IV (see Introduction).  It appears that almost four-fifths of the neotropical hepatic families are represented in the páramos;  those lacking are usually small, specialized groups.  Many families have only few taxa in páramos, however. 

      It appears that the largest number of species recorded per country by far have been those from Colombian páramos (230 spp.);  the smallest number are known from Panama (7 spp.).  The relatively large area covered by páramo and the very moist climate may explain why Colombian páramos have so many species.  On the other hand, Colombian páramos have also been studied more intensively than those of other countries.  Therefore, the principal conclusion that can be drawn is that our knowledge of the páramo flora is very uneven and that much more collecting needs to be done to reach a reasonable level of understanding of hepatic diversity in the páramos of the different countries. 

      All taxa recorded from 3200 m and upwards in the Andes and above 3000 m in Costa Rica have been included in the list of species.  Species occurring at lower elevations are excluded from this list unless it is stated that the record was from páramo.  Elevational ranges of species are the total range of the taxon in the Neotropics. 

      The information is based on the references cited at the end of the list and on herbarium specimens in SJU (records from Costa Rica), PMA (records from Panama), and U (all countries).  The latter herbarium holds the largest collection of páramo hepatics worldwide and was a particularly rich source of information.  In some instances it was uncertain whether the record was actually from páramo.  This concerns particularly old references or old herbarium specimens, lacking details on locality or habitat.  Additional fieldwork on the habitat preferences of the species is needed to help resolve these uncertainties. 

ANTHOCEROTAE 

ANTHOCEROTACEAE 

Phaeoceros Prosk. 
P. pichinchensis (Spruce) Haessel;  EC;  ?-3600 m 
 

DENDROCEROTACEAE 

Megaceros Campb. 
M. sp.;  CO;  ?-4000 m 
 

HEPATICAE 

ACROBOLBACEAE 

Acrobolbus Nees 
A. laceratus R.M.Schust.;  VE;  3140 m 

Lethocolea Mitt. 
L. glossophylla (Spruce) Grolle;  CR, CO, VE, EC, PE;  2400-3750 m 

Marsupidium Mitt. 
M. latifolium R.M.Schust.;  VE;  3100 m 

Tylimanthus Mitt. 
T. setaceo-ciliatus Steph.;  CR?, CO, EC;  2800-4000 m 
 

ADELANTHACEAE 

Adelanthus Mitt. 
 Ref.:  Grolle, 1972;  Schuster, 1978. 
A. aureomarginatus R.M.Schust.;  CO, VE;  2000-3500 m 
A. crossii Spruce;  CR, CO, VE;  3000-4250 m 
A. decipiens (Hook.) Mitt. ssp. aureocinctus R.M.Schust.; VE; 3140 m  [Note:
   A. crossii and A. decipiens subsp. aureocinctus may be untoothed high-altitude forms
   of A. decipiens (Hook.) Mitt.] 
A. lindenbergianus (F.Lehm.) Mitt.;  CR, PA, CO, VE, EC, PE;  1800-4200 m 
A. pittieri (Steph.) Grolle;  CR, CO, VE;  2500-3500 m 
 

ANEURACEAE 

Cryptothallus Malmb. 
 Ref.:  Crum & Bruce, 1997. 
C. hirsutus Crum;  CR;  3100 m 

Riccardia Gray 
 [Note:  Riccardia is one of the most common hepatic genera in the neotropical páramos;  the taxonomy and distribution of the species are still very incompletely known.  The present list is largely based on Meenks (1987).  The lack of records from Costa Rican páramo is an artifact;  several (unidentified) species have been collected by the author and others.] 
 Ref.:  Meenks, 1987. 
R. aberrans (Steph.) Gradst.;  CO, PE;  3000-3700 m 
R. algoides (Taylor) Meenks [syn.: R. squarrosa (Steph.) Gradst.]; CO, EC; 3000-3900
   m 
R. capillacea (Steph.) Meenks & DeJong;  CO, VE, EC?, PE;  2600-3900 m 
R. ciliolata (Spruce) Gradst.;  CO, VE, EC, PE;  2650-3750 m 
R. columbica (Steph.) Hässel ex Gradst.;  CO;  1900-3700 m 
R. crassicaulis (Steph.) Meenks & DeJong;  CO, PE;  1900-3600 m 
R. foliacea Meenks & DeJong;  EC;  4000 m 
R. hansmeyeri (Steph.) Meenks & DeJong;  CO, VE, EC;  3200-4300 m 
R. herzogiana (Steph.) Meenks & DeJong;  CO, VE, PE;  1400-4000 m 
R. judithae Meenks & DeJong;  CO, PE;  3000-3400 m 
R. leptophylla (Spruce) Herzog;  CO, VE, EC, PE;  1000-3500 m 
R. pallida (Spruce) Meenks & DeJong;  CO, EC;  1500-4100 m 
R. papillata (Gottsche) Hässel ex Gradst.; CO, VE; 1950-3450 m [Note: The record of R.
   insignis Schiffn. from Colombian páramo (Winkler, 1976) is erroneous and refers to R.
   papillata (Meenks, 1987).] 
R. paramorum Meenks;  CO;  3400-3800 m 
R. parasitans (Steph.) Meenks & DeJong;  CO, VE, EC, PE;  3000-4500 m 
R. plumaeformis (Spruce) Meenks;  CO, EC, PE;  1500-3700 m 
R. poeppigiana (L.Lehm. & Lindenb.) Hässel ex Meenks & DeJong; CO, EC, PE;
   2900?-4200 m 
R. smaragdina Meenks & DeJong;  CO, VE;  2350-3600 m 
R. trichomanoides (Spruce) Meenks;  CO, EC, PE;  2900-3900 m 
R. wallisii (Steph.) Gradst.;  CO;  3000?-4000 m 
 

ARNELLIACEAE 

Gongylanthus Lindb. 
G. granatensis (Gottsche) Steph.;  CO, PE;  2500-4050 m 
G. liebmannianus (Lindenb. & Gottsche) Steph.; CR, PA, CO, VE, EC, PE; 2600-4300
   m 
G. limbatus (Herzog) Grolle & Vána (syn.: G. innovans S.Winkl., n.v.); CO, VE;
   3100-4300 m 
 

AYTONIACEAE 

Asterella P.Beauv. 
A. macropoda (Spruce) A.Evans;  CR, CO, EC, PE;  2300-3700 m 
 

BALANTIOPSIDACEAE 

Isotachis Mitt. 
 Ref.:  Fulford, 1963. 
I. lacustris Herzog (syn.: Triandrophyllum maegdefraui S.Winkl.); CO, EC; 2500-4700
   m 
I. lopezii (R.M.Schust.) Gradst. (syn.: Ruizanthus lopezii R.M.Schust.); CR, CO, VE, EC,
   PE; 3000-3800 m [Note: Ruizanthus lopezii (fide Gradstein 9747, Costa Rica, Cerro de
   la Muerte, on bank of Panamerican Highway together with R. venezuelanus, elev. 3000
   m, 29 Sep 1995, GOET) has spiralled instead of straight capsule valves. The 2-lobed
   leaves of this species also indicate a closer relationship to Isotachis than to Ruizanthus,
   which has 4-lobed leaves. Therefore, it was transferred to Isotachis, as I. lopezii
   (R.M.Schust.) Gradst.(basionym: Ruizanthus lopezii R.M.Schust., Phytologia 39: 241.
   1978).] 
I. lindigiana Gottsche;  CO;  2300-3500 m 
I. multiceps Gottsche;  CR, CO, VE, EC, PE;  1250-4100 m 
I. obtusa Steph.; VE, EC; 3200-3400 m [Note: Isotachis lacustris and I. obtusa are
   characteristic species of shallow lakes and rivulets in neotropical páramos. As shown by
   Fulford (1963), the type specimens have very different leaf shapes (ovate in I. lacustris,
   orbicular in I. obtusa) and leaf incisions. However, additional collections made by the
   author and others show considerable variation and overlap in these characters, suggesting
   that the two may be conspecific. In the latter case, I. obtusa Steph. would be the older,
   hence correct name.] 
I. serrulata (Sw.) Gottsche [syn.: I. haematodes (Lindenb. & Gottsche) Gottsche, I.
   madida (Hook. & Taylor) Mitt.]; PA, CO, VE, EC, PE; 700-4450 m [Note: Isotachis
   haematodes (Lindenb. & Gottsche) Gottsche and I. madida (Hook. & Taylor) Mitt. are
   treated as synonyms of the common, highly variable I. serrulata following Gradstein et al.
   (1977).] 

Ruizanthus R.M.Schust.  [Note:  See note under Isotachis lopezii.] 
R. venezuelanus R.M.Schust.;  CR, CO, VE;  3000-4050 m 
 

CALYPOGEIACEAE 

Calypogeia Raddi 
 Ref.:  Fulford, 1968. 
C. andicola Bischler;  CO, EC;  800-4100 m 
C. cyclostipa (Spruce) Steph.;  CO, PE;  1700-3600 m 
C. peruviana Nees & Mont.;  CO, EC, PE;  500-3800 m 
 

CEPHALOZIACEAE 

Cephalozia (Dumort.) Dumort. 
 Ref.:  Vána, 1988. 
C. bicuspidata (L.) Dumort.;  CO;  3500-3800 m 
C. crossii Spruce (syn.:  C. dussii Fulford);  CR, CO, EC, PE;  500-4100 m 
C. pleniceps (Austin) Lindb.;  CO;  3500-3700 m

Odontoschisma (Dumort.) Dumort. 
O. atropurpureum Steph.;  CO, VE, EC;  250?-4000 m 
O. denudatum (Nees) Dumort.;  CO, EC;  600?-3500 m 
 

CEPHALOZIELLACEAE 

Cephaloziella (Spruce) Schiffn. 
 Ref.:  Fulford, 1976;  Schuster, 1978b. 
C. divaricata (Sm.) Schiffn. (syn.:  C. andina Herzog);  CO, VE, EC;  3400 m 
C. fragillima (Spruce) Fulford;  CO, VE;  3000-3330 m 
C. granatensis (Jack) Fulford;  CR, PA, CO;  2500-4000 m 
C. grisea R.M.Schust.;  VE;  4150 m 
C. pungens Steph. ex Fulford;  VE;  3600 m 
C. stolonifera R.M.Schust.;  VE;  3650 m 
 

FOSSOMBRONIACEAE 

Austrofossombronia R.M.Schust. 
 Ref.:  Schuster, 1994. 
A. peruviana (Gottsche) R.M.Schust. (syn.: Fossombronia ptychophylla Spruce, n.v.);
   CO, EC, PE;  3600-4000 m 

Fossombronia  Raddi 
[Note: Neotropical Fossombronia has not been studied critically. The present list therefore likely contains errors in identification.] 
F. brasiliensis Steph.;  CO, EC;  1800-4200 m 
F. paranapanemae Schiffn.;  CO;  3000-4250 m 
F. sp.;  CR;  3000 m 
 

GEOCALYCACEAE 
 Ref.:  Fulford, 1976. 

Campanocolea R.M.Schust. 
 Ref.:  Schuster, 1997. 
C. fragmentissima (R.M.Schust.) R.M.Schust. (syn.: Lophocolea fragmentissima
   R.M.Schust.);  CO, VE, EC, PE;  3200-4200 m 

Clasmatocolea Spruce 
 Ref.:  Engel, 1980. 
C. vermicularis (F.Lehm.) Grolle;  CR, CO, VE, EC, PE;  1500-4500 m 

Heteroscyphus Schiffn. 
H. marginatus (Steph.) Fulford;  CO, VE, EC;  1000-3300? m 
H. polyblepharis (Spruce) Schiffn.;  CO, VE, EC, PE;  2300-3500 m 

Leptoscyphus Mitt. 
L. amphibolius (Nees) Grolle;  PA;  1000-3150 m 
L. cleefii Fulford;  CO;  3200-4100 m 
L. cuneifolius (Hook.) Mitt.;  CR, PA, CO, VE, EC, PE;  2600-4400 m 
L. jackii (Steph.) Grolle;  CO, EC;  3000-3900 m 
L. obcordatus (Spruce) Grolle;  CO, EC;  3300-3500? m 
L. physocalyx (Hampe & Gottsche) Gottsche;  CR, CO, VE, EC, PE;  2000-3850 m 
L. porphyrius (Nees) Grolle;  PA, CO, VE, EC, PE;  100-4300 m 

Lophocolea (Dumort.) Dumort. 
L. bidentata L. (syn.: L. coadunata (Sw.) Nees);  PA, CO, VE, EC, PE; 100-3700 m 
L. erosa Gradst.;  CR, EC;  3200-3700 m 
L. granatensis Gottsche;  CO, PE;  1000-3700 m 
L. muricata (F.Lehm.) Nees;  CO, VE;  1300-3400 m 
L. trapezoidea Mont.;  CR, CO, EC, PE;  500-4000 m 

Platycaulis R.M.Schust. 
 Ref.: Schuster, 1995. 
P. renifolia R.M.Schust.;  VE;  ? m 

Pseudocephaloziella R.M.Schust. 
 Ref.: Schuster, 1991. 
P. epiphytica R.M.Schust.;  VE;  3140 m 
 

GYMNOMITRIACEAE 

Gymnomitrion Corda 
 Ref.:  Schuster, 1996a;  Vána, 1976. 
G. andinum (Herzog) Herzog;  CO, PE;  3450-4200 m 
G. atrofilum Vána;  CO;  3400-4000 m 
G. setaceum Grolle & Vána;  CR, CO;  3000-4300 m 
G. truncato-apiculatum Herzog;  CR, CO;  3400-4300 m 

Marsupella Dumort. 
 Ref.:  Schuster, 1996a. 
M. emarginata (Ehrh.) Dumort.;  CO;  3150 m 
M. involuta Vána;  CO;  3600-4300 m 
M. microphylla R.M.Schust.;  VE;  4160 m 
M. revoluta (Nees) Dumort.;  VE;  4400-4500 m 
M. trollii Herzog;  CR, CO;  3500-4400 m 
M. xenophylla R.M.Schust. [syn.: Nanomarsupella xenophylla (R.M.Schust.)
   R.M.Schust.];  VE;  4160 m 

Paramomitrion R.M.Schust. 
 Ref.:  Schuster, 1996a. 
P. paradoxum R.M.Schust.;  VE;  4160 m 

Stephaniella Jack 
 Ref.:  Schmitt & Winkler, 1969. 
S. paraphyllina Jack;  CR, CO, VE, EC, PE;  2700-4500 m 
S. rostrata U.Schmitt;  CR, CO;  3000-4300 m 

Stephaniellidium S.Winkl. ex Grolle 
S. sleumeri (Müll.Frib.) S.Winkl. ex Grolle;  CO;  3100-? m 
 

HAPLOMITRIACEAE 

Haplomitrium Nees 
 Ref.:  Bartholomew-Began, 1992. 
H. blumei Nees [syn.: H. andinum (Spruce) R.M.Schust.];  EC;  3900 m 
 

HERBERTACEAE 

Herbertus S. Gray 
 Ref.:  van Reenen, 1982. 
H. acanthelius Spruce (syn.: H. limbatus Steph.);  CR, CO, EC, PE;  2400-4350 m 
H. colombianus Reenen;  CO;  3250-3750 m 
H. juniperoides (Sw.) Grolle;  CR, PA, CO, EC, PE;  2000-3600 m 
H. oblongifolius (Steph.) Gradst. & Cleef;  CO;  4060 m 
H. subdentatus (Steph.) Fulford (syn.: H. subnivalis S.Winkl.); CR, PA, CO, VE, EC,
   PE; 2400-4350 m 

Olgantha R.M.Schust. 
 Ref.:  Schuster, 1996b. 
O. eophylla R.M.Schust.;  EC;  4100 m 

Triandrophyllum Fulford & Hatcher 
[Note:  Triandrophyllum maegdefraui S.Winkl., described from the Sierra Nevada de Santa Marta, Colombia, seems to be a synonym of Isotachis lacustris judging from the original description and illustration.] 
T. subtrifidum (Hook. & Taylor) Fulford & Hatcher; CR, PA, CO, VE, EC, PE;
   1800-4500 m 
 

JUBULACEAE 

Frullania Raddi 
 Ref.:  Stotler, 1969;  Yuzawa, 1991. 
F. albertii Steph.; CO, EC; 3400-3950 m [Note: F. albertii is probably a robust,
   high-altitude form of F. standaertii Steph. Both taxa are characterized by the rather large,
   lanceolate stylus (Yuzawa, 1991).] 
F. arecae (Spreng.) Gottsche; CO, EC; 800-3800 m [Note: The páramo records of F.
   arecae are in need of revision. As shown by Yuzawa (1991), the species has been
   confused with F. ecklonii (Spreng.) Spreng. and with F. viminicola Spruce.] 
F. brasiliensis Raddi;  CO, EC;  200-3800 m 
F. cuencensis Taylor;  CO, EC;  2100-3400 m 
F. convoluta Lindenb. & Hampe;  PA, CO, VE, EC, PE;   1900-4000 m 
F. dusenii Steph.;  CO, EC;  1600-3400 m 
F. holostipula Hatt. & D.G.Griffin;  VE;  2600-3300 m 
F. lobato-hastata Steph.;  CO;  1800-3500 m 
F. peruviana Gottsche; CO, VE, EC, PE; 1500-3700 m [Note: Two species of Frullania
   sect. Meteoriopsis are recognized in neotropical páramos: F. convoluta (with rounded to
   obtuse leaf apex) and F. peruviana (with acute to short acuminate leaf apex). Both
   species have deeply cordate leaf bases with two large auricles.] 
F. planifolia Steph.;  CR;  3000 m 
F. pluricarinata Gottsche;  CO, EC;  1800-3350 m 
F. sphaerocephala Spruce;  CR, CO, EC, PE;  2700-4000 m 
F. tetraptera Nees & Mont.;  CO, VE, EC, PE;  3100-3850 m 
 

JUNGERMANNIACEAE 

Anastrophyllum (Spruce) Steph. 
 Ref.:  Vána, 1980, 1984;  Gradstein & Vána, 1987. 
A. auritum (F.Lehm.) Steph.;  CR, CO, VE, EC, PE;  2000-4500 m 
A. austroamericanum Vána (syn.: Marsupella austroamericana Vána, nom. inval.);
   CO,VE;   3750-4300 m 
A. leucocephalum (Taylor) Steph.;  CO, EC, PE;  2300-4250 m 
A. minutum (Schreb.) R.M.Schust. (syn.: A. gemmiferum S.Winkl.); CO, VE;
   3700-4500 m [Note: Anastrophyllum gemmiferum S.Winkl. (Type: Colombia, Sierra
   Nevada de Santa Marta, leg. S. Winkler, ULM) is treated as a synomym of A. minutum
   (Schreb.) R.M.Schust. at the advice of Dr. J. Vána (in litt.).] 
A. nigrescens (Mitt.) Steph.;  CR, CO, VE, EC, PE;  2000-4300 m 
A. pearcei (Steph.) R.M.Schust.;  CO, VE, PE;  ? m 
A. stellatum R.M.Schust. (syn.: A. gradsteinii Vána, nom. inval.); CR, CO, VE, EC, PE;
   3140-4000 m 
A. tubulosum (Nees) Grolle;  CO, VE, EC, PE;  ? m 

Andrewsianthus R.M.Schust. 
A. jamesonii (Mont.) Vána [syn.: A. kilimajaricus (S.W.Arnell) Grolle & Vána; Lophozia
   incisa (Schrad.) Dumort. ssp. austrigena R.M.Schust., n.v.); CO, VE, EC, PE;
   2000-4750 m 

Cryptochila R.M.Schust. 
 Ref.:  Grolle, 1971. 
C. grandiflora (Lindenb. & Gottsche) Grolle;  PA, CO, VE, EC, PE;  1300-4630 m 

Gymnocoleopsis (R.M.Schust.) R.M.Schust. 
G. multiflora (Steph.) R.M.Schust.;  CO, VE, PE;  3500-4400 m 

Jamesoniella (Spruce) Carring 
 Ref.:  Grolle, 1971. 
J. autumnalis (DC.) Steph.;  CO, VE;  2000-3500 m 
J. rubricaulis (Nees) Grolle;  CR, PA, CO, VE, EC, PE;  1350-4300 m 
J. undata (Mont.) Steph.;  CO, PE;  1900-4300 m 

Jungermannia L. 
 Ref.:  Vána, 1973, 1974. 
J. decolor Schiffn.;  CO, PE;  500-3500 m 
J. hyalina Lyell;  CO;  3150 m 
J. linguifolia Gottsche;  CO, EC, PE;  2500-3600 m 
J. ovato-trigona (Steph.) Grolle;  CO, EC, PE;  2800-4200 m 
J. sphaerocarpa Hook.;  CR, CO, VE, EC, PE;  2000-4100 m 

Lophonardia R.M.Schust. 
[Note:  According to Dr. J. Vána (in litt.), Lophonardia is a doubtful taxon, probably not meriting recognition as a separate genus.] 
L. caespitosa R.M.Schust.;  VE;  4150 m 

Lophozia (Dumort.) Dumort. 
L. incisa (Schrad.) Dumort.;  CR, CO, VE, EC, PE;  2500-4500 m 
L. laxifolia (Mont.) Grolle (syn.: L. subinflata Steph.); CR, CO, VE, EC; 3000-4100 m 
L. stolonifera R.M.Schust.;  VE;  ? m 
L. verruculosa R.M.Schust.;  VE;  ca. 3150 m 

Nardia Gray 
 Ref.:  Engel, 1988. 
N. succulenta (Rich. ex F.Lehm.) Spruce;  CO, EC;  800-3300 m 

Rhodoplagiochila R.M.Schust. 
[Note:  Following Inoue (1984), Rhodoplagiochila is placed in the Jungermanniaceae instead of Plagiochilaceae.] 
 Ref.:  Schuster, 1978;  Inoue, 1984. 
R. rosea R.M.Schust.;  VE;  3700-3750 m 

Syzygiella Spruce 
 Ref.:  Inoue, 1966. 
S. anomala (Lindenb. & Gottsche) Steph.;  CR, CO, EC, PE;  2000-3600 m 
S. campanulata Herzog;  CR, CO, VE;  2000-3750 m 
S. integerrima Steph.;  CO;  1000-3550 m 
S. liberata Inoue;  CR, CO;  3000-3900 m 
S. manca (Mont.) Steph.;  CO, EC, PE;  3000?-3500 m 
 

LEJEUNEACEAE 

Amphilejeunea R.M.Schust. 
 Ref.:  Schuster, 1986. 
A. patellifera (Spruce) R.M.Schust.;  CR;  2500-3200 m 
A. viridissima R.M.Schust.;  CO, VE;  3000-4150 m 

Anoplolejeunea (Spruce) Schiffn. 
A. conferta (Meissn.) A.Evans;  CR, CO, VE, EC, PE;  600-3700 m 

Aureolejeunea R.M.Schust. 
 Ref.:  Schuster, 1986, 1987. 
A. aurifera R.M.Schust.;  CO, VE;  3500-3700 m 
A. paramicola (Herzog) R.M.Schust. [syn.: A. paramoensis R.M.Schust.; Omphalanthus
   paramicola (Herzog) Gradst.]; CO, VE, EC;  2000-3700 m 
A. quinquecarinata R.M.Schust.;  CO, VE;  3000-4000 m 

Blepharolejeunea R.M.Schust. 
 Ref.:  Gradstein, 1994. 
B. incongrua (Lindenb. & Gottsche) Van Slageren & Kruijt; CR, CO, VE, EC, PE;
   1800-4100 m 
B. securifolia (Spruce) R.M.Schust.;  CR, CO, VE, EC, PE;  3000-4600 m 

Brachiolejeunea (Spruce) Schiffn. 
 Ref.:  Van Slageren, 1985;  Gradstein, 1994 
B. laxifolia (Taylor) Schiffn.;  CR, CO, VE, EC, PE;  1500-3800 m 

Cheilolejeunea (Spruce) Schiffn. 
[Note:  Two additional species of Cheilolejeunea from the páramos of Colombia were described by Schuster (1992).  Since Latin descriptions were not provided, the names are invalid and have been omitted from the present list.] 
C. choachina (Gottsche) Gradst., [syn.: Lejeunea choachina Gottsche, Ann. Sci. Nat.
   Bot., Sér. 5, 1:156. 1864; Strepsilejeunea choachina (Gottsche) Steph.]; CO;
   3000-4300 m 
C. erostrata R.M.Schust.;  VE;  3140 m 
C. laevicalyx (Jack & Steph.) Grolle;  CO;  3300 m 

Colura Nees 
C. calyptrifolia (Hook.) Dumort.;  VE;  3200+ m 
C. naumannii Steph. (syn.:  C. patagonica Ast);  CO, EC;  3200-3800 m 
C. ornithocephala Herzog;  CO, VE, EC;  3300-3800 m 
C. tenuicornis (A.Evans) Steph.;  VE;  2300-3450 m 

Diplasiolejeunea (Spruce) Schiffn. 
D. alata Ast.;  VE;  2900-3300 m 
D. involuta S.Winkl. ssp. andicola Pócs;  VE;  3300 m 
D. papilionacea R.M.Schust.;  VE;  2900-3450 m 
D. pauckertii (Nees) Steph.;  CO, VE, PE;  2900-3340 m 
 

Drepanolejeunea (Spruce) Schiffn. 
 Ref.:  Bischler, 1964. 
D. andina Herzog;  CO;  3500-4100 m 
D. araucariae Steph.;  VE;  3050-4000 m 
D. aurita Bischler;  EC;  3350-3450 m 
D. granatensis (Jack & Steph.) Bischler;  CO, VE;  2000-3700 m 
D. navicularis Steph.;  CO, VE;  3000-3700 m 

Frullanoides Raddi 
 Ref.:  Van Slageren, 1985;  Gradstein, 1994. 
F. densifolia Raddi;  CO, EC, PE;  500-3650 m 

Harpalejeunea (Spruce) Schiffn.
?H. ancistrodes (Spruce) Schiffn.;  CO, EC;  3000-3800 m 
H. cinchonae (Nees) Schiffn.;  CO, VE, PE;  700-3400 m 
H. grandis Grolle;  CO;  3750 m 
H. verrucosa Herzog;  VE;  3200 m 
H. sp. nov.;  CO;  3200-3680 m 

Leucolejeunea A.Evans 
L. xanthocarpa (F.Lehm. & Lindenb.) A.Evans;  CO, VE, EC;  500-3500 m 

Lindigianthus Kruijt & Gradst. 
 Ref.:  Gradstein, 1994. 
L. cipaconeus (Gottsche) Kruijt & Gradst. [syn.: Dicranolejeunea cipaconea (Gottsche)
   Steph.];  CR, CO, VE, EC, PE;  1700-3500 m 

Macrolejeunea (Spruce) Schiffn. 
M. pallescens (Mitt.) Schiffn.;  CR, CO, VE, EC, PE;  2000-3800 m 

Microlejeunea (Steph.) Schiffn. 
M. bullata (Taylor) Steph.;  CR, CO, VE, EC;  (0-)1000-3600 m 
M. colombiana Bischl.;  VE;  3090-4070 m 

Omphalanthus Nees 
O. filiformis (Sw.) Nees;  CR, CO, EC, PE;  1200-3900 m 
O. platycoleus Herzog;  CO;  3000?-3750 m 
 

LEPICOLEACEAE 

Lepicolea Dumort. 
 Ref.: Fulford, 1963. 
L. ochroleuca (Spreng.) Spruce;  CR;  1600-3100 m 
L. pruinosa (Taylor) Spruce;  CR, CO, VE, EC, PE;  1700-3800 m 
L. ramentifissa Herzog;  VE;  2900-3300 m 
 

LEPIDOZIACEAE 

Bazzania S. Gray 
[Note:  The neotropical species of Bazzania and Lepidozia are poorly known and in need of revision;  the present list likely contains misidentifications and some species names may prove to be synonyms.] 
 Ref.: Fulford, 1963. 
B. arcuata (Lindenb. & Gottsche) Trevis.;  CR, CO, EC;  200-3600 m 
B. canelensis (Steph.) Fulford;  CO;  ?-3750 m 
B. chilensis (Steph.) Fulford;  CO, EC;  2500-3600 m 
B. crassidentata Fulford;  CO;  3500-3600 m 
B. diversicuspis Spruce;  CO;  100-3500 m 
B. falcata (Lindenb.) Trevis.;  CO;  1000-3600 m 
B. hookeri (Lindenb.) Trevis. (syn.:  B. robusta Spruce);  CO, VE, PE;  100-3700 m 
B. jamaicensis (F.Lehm. & Lindenb.) Trevis.;  CR, CO;  1500-3500 m 
B. latidens (Gottsche) Fulford;  CO;  3750 m 
B. longistipula (Lindenb.) Trevis.;  CO, EC;  750-3500 m 
B. placophylla (Taylor) Grolle;  CO, EC, PE;  ?-3750 m 
B. roraimensis (Steph.) Fulford;  CO, PE;  1500-3600 m 
B. taleana (Gottsche) Fulford;  VE;  3200+ m 

Kurzia G.Martens 
 Ref.:  Pócs, 1984. 
K. capillaris (Sw.) Grolle (syn.: K. verrucosa Steph.); CR, CO, VE, EC, PE; 500-3800 m 
K. flagellifera (Steph.) Grolle;  CO;  1500-3700 m 

Lepidozia (Dumort.) Dumort. 
[Note:  The neotropical species of Bazzania and Lepidozia are poorly known and in need of revision;  the present list likely contains misidentifications and some species names may prove to be synonyms.] 
 Ref.:  Fulford, 1966. 
L. alstoni Fulford;  CO, EC;  3200-4500 m 
L. andicola Beauverd;  CO, VE, EC;  2650-3600 m 
L. auriculata Mitt. ex Steph.;  CO, EC, PE;  3300?-4000 m 
L. caespitosa Spruce;  CO, VE, EC, PE;  1200-3800 m 
L. cupressina (Sw.) Lindenb.;  PA, CO, VE, PE;  1000-3600 m 
L. macrocolea Spruce;  CO, VE, EC, PE;  1000-4000 m 
L. peruviensis Steph.;  CO, VE, EC, PE;  2500-3650 m 
L. wallisiana Steph.;  CO, VE, EC;  1000-3400 m 

Paracromastigum Fulford & J.Tayl. 
P. bifidum (Steph.) R.M.Schust.;  VE;  2000-3400 m 
P. granatense (Gottsche) R.M.Schust. [syn.: Bonneria granatensis (Gottsche) Fulford &
   J.Tayl.];  CR, CO;  ?-3200 m 

Pseudocephalozia R.M.Schust. 
 Ref.:  Schuster & Engel, 1974. 
P. quadriloba (Steph.) R.M.Schust.;  CR, CO, EC, PE;  3200-4100 m 

Telaranea Spruce ex Schiffn. 
 Ref.:  Schuster, 1978. 
T. microstipulata R.M.Schust.;  VE;  ? m 
T. nematodes (Gottsche ex Austin) Howe;  CR, PA, CO, VE, EC, PE;   200-4250 m 
T. quadrifida R.M.Schust.;  VE;  ? m 
 

MARCHANTIACEAE 

Dumortiera Nees 
D. hirsuta L.;  CO, EC;  150-3500 m 

Marchantia L. 
 Ref.: Bischler, 1984. 
M. berteroana F.Lehm. & Lindenb.;  CR, CO;  1800-3600 
M. plicata Nees & Mont.;  CR, CO, EC, PE;  1700-3800 
M. polymorpha L.;  CO;  3000-3800 
 

METZGERIACEAE 

Metzgeria Raddi 
[Note:  The neotropical species of Metzgeria are still poorly known, notwithstanding the work by Kuwahara (1986);  the present list should be considered preliminary.] 
 Ref.:  Kuwahara, 1986. 
M. albinea Spruce;  CO, EC;  500-3800 m 
M. atramentaria Kuwah.;  CO;  3450-4000 m 
M. attenuata Steph.;  CO;  3350-3600 m 
M. bischlerae Kuwah.;  CO;  3500-3600 m 
M. cleefii Kuwah.;  CO;  3300 m 
M. consanguinea Schiffn.;  CO, VE, PE;  2100-3500 m 
M. decipiens (C.Massal.) Schiffn.;  CO, PE;  500-3900 m 
M. dorsipara (Herzog) Kuwah.;  CO, EC;  3350-3900 m 
M. filicina Mitt.;  CO, EC, PE;  2000-3300 m 
M. gigantea Steph.;  CR, CO, PE;  1500-4350 m 
M. lechleri Steph.;  CO, PE;  3000-3800 m 
M. leptoneura Spruce;  CO, VE, EC, PE;  500-4200 m 
M. liebmanniana Lindenb. & Gottsche;  CO, VE, EC, PE;  1500-4000 m 
M. metaensis Kuwah.;  CO;  4000-4200 m 
M. neotropica Kuwah.;  CO, EC, PE;  3100-4000 m 
M. papulosa Steph.;  CO, VE;  700-3600 m 
M. parviinvolucrata Kuwah.;  PE;  3400 m 
M. polytricha Spruce;  CO, PE;  1000-4000 m 
M. procera Mitt.;  CO;  200-3600 m 
M. sandei Schiffn.;  CO;  650-3950 m 
 

MONOCLEACEAE 

Monoclea Hook. 
 Ref.:  Gradstein et al., 1993. 
M. gottschei Lindb. ssp. elongata Gradst. & Mues; CR, CO, EC; 150-3500 m 
 

PALLAVICINIACEAE 

Jensenia Lindb. 
 Ref.:  van der Gronde, 1980. 
J. erythropus (Gottsche) Grolle;  CR, PA, CO, VE, EC, PE;  2200-4100 m 
J. florschuetzii van der Gronde;  CO, EC;  4000-4100 m 

Symphyogyna Nees & Mont. 
 Ref.:  Uribe & Aguirre, 1995. 
S. brasiliensis Nees & Mont.;  CR, CO, EC;  1000-3300 m 
S. brogniartii Mont.;  CO, VE, EC;  700-3600 m 
S. marginata Steph.;  CO;  800-4000 m 
S. podophylla (Thunb.) Mont. & Nees;  CO;  1800-3850 m 
S. rubescens Steph.;  CO;  ?-3600 m 
 

PELLIACEAE 

Noteroclada Taylor ex Hook. & Taylor 
N. confluens Taylor ex Hook. & Wilson; CR, CO, VE, EC, PE; 1800-4000 m 
 

PLAGIOCHILACEAE 

Plagiochila (Dumort.) Dumort. 
[Note:  The neotropical species of Plagiochila are very poorly known.  The present list is largely based on identifications of the late Dr. H. Inoue, whose revision of the group remained unfinished atby his untimely death in 1989.] 
 Ref.:  Inoue, 1987, 1989;  Robinson, 1967. 
P. cleefii Inoue;  CO;  4050 m 
P. cuatrecasii H.Rob.;  CO;  3600-4200 m 
P. dependula Taylor [syn.: Jamesoniella dependula (Taylor) Steph.]; CO, EC;
   3500-4200 m 
P. echinella Gottsche;  CO, VE?, EC, PE;  2400-3750 m 
P. fuscolutea Taylor (syn.: P. jelskii Loitl., P. scopulosa Steph.); CR, CO, VE, EC, PE;
   2600-4000 m 
P. guevarii H.Rob. (syn.: P. dana-griffinii Inoue, n.v.);  CO, VE;  3300-3700 m 
P. jamesonii Taylor;  CO, EC;  2000-3600 m 
P. jaramilloi H.Rob.;  CR, CO;  3000-3520 m 
P. longispina Lindenb. & Gottsche;  CR, CO, VE, EC;  2800-3700 m 
P. oblita Steph.;  CO;  3750 m 
P. ovata Lindenb. & Gottsche;  CR, CO;  2500-3450 m 
P. pachyloma Taylor;  CO, EC, PE;  2000-3600 m 
P. paraphyllina Herzog;  CO;  3200 m 
P. revolvens Mitt.;  CO, EC;  4000-4300 m 
P. stolonifera Lindenb. & Gottsche;  CR, CO;  1900-4200 m 
P. triangulifolia Steph.;  CO;  4050 m 
P. umbrosa Steph.;  CO;  3000-3750 m 
P. verruculosa R.M.Schust.;  CO;  1500-3300 m 

Steereochila Inoue 
 Ref.:  Inoue, 1988;  Gradstein et al., 1994. 
S. ecuadorica Inoue;  CR, EC;  2500?-3200 m 
 

PLEUROZIACEAE 

Pleurozia Dumort. 
 Ref.:  Thiers, 1993. 
P. paradoxa Jack [syn.: Eopleurozia paradoxa (Jack) R.M.Schust.]; CO, VE, EC;
   2500-3800 m 
 

PORELLACEAE 

Porella L. 
 Ref.:  Swails, 1970. 
P. leiboldii (F.Lehm. & Lindenb.) Trevis.;  CR;  1360-4000 m 
P. squamulifera (Taylor) Trevis.;  CO, VE, EC, PE;  2200-3900 m 
 

PSEUDOLEPICOLEACEAE 
 Ref.:  Schuster, 1985. 

Blepharostoma (Dumort.) Dumort. 
B. trichophyllum (L.) Dumort.;  CR, CO, VE, PE;  2700-4000 m 

Temnoma Mitt. 
T. chaetophylla R.M.Schust.;  CO, VE;  3000-3700 m 
 

RADULACEAE 

Radula Dumort. 
 Ref.:  Jans, 1980;  Yamada, 1988;  Reiner-Drehwald, 1994. 
R. episcia Spruce (syn.: R. cornucopiae Spruce);  CO, EC;  1000-3600 m 
R. frondescens Steph.;  CO, VE;   ?-3800 m 
R. jamesonii Taylor;  CO, EC;  ?-3700 m 
R. nudicaulis Steph.;  CO;  1500-4000 m 
R. sonsonensis Taylor;  CO;  3000-4200 m 
R. voluta Taylor (syn.: R. ramulina Taylor);  CR, CO, EC, PE;  1500-4200 m 
 

RICCIACEAE 

Riccia L. 
R. lamellosa Raddi (syn.: R. indusiata S.Winkl.); CO; 4200 m [Note: Riccia indusiata
   S.Winkl., described from the Sierra Nevada de Santa Marta, Colombia, is a synonym of
   R. lamellosa Raddi (fide holotype in ULM;  identity confirmed by S. Jovet-Ast, Paris).] 
R. sorocarpa Bisch.;  PE;  1500-4550 m 
 

SCAPANIACEAE 
 Ref.:  Gradstein & Vá_a, 1987. 

Diplophyllum (Dumort.) Dumort. 
D. andicolum R.M.Schust.;  VE;  4200 m 
D. obtusatum (R.M.Schust.) R.M.Schust.;  CO, VE, EC, PE;  2500-4200 m 

Scapania (Dumort.) Dumort. 
S. cuspiduligera (Nees) Müll.Frib.;  CO;  4300 m 
S. portoricensis Hampe & Gottsche;  CR, CO, VE, EC, PE;  1500-3750 m 
 

TRICHOCOLEACEAE 

Trichocolea Dumort. 
[Note: The taxonomy of neotropical Trichocolea is still incompletely known;  the present list likely contains misidentifications and some species names may prove to be synonyms.  Most species listed are forest taxa occurring only accidentally in páramo.] 
 Ref.:  Fulford, 1963. 
T. elliotii Steph.;  CO;  1000-3650 m 
T. filicaulis Steph.;  CO;  800-3500 m 
T. robusta Steph.;  CO;  2800-360 m 
T. sprucei Steph.;  CO, PE;  1800-3600 m 
T. tomentosa (Sw.) Gottsche;  CR, PA, CO, VE, EC, PE;  100-3600 m 
 

Acknowledgments 

 This paper was drafted while the author was an Andrew W. Mellon Visiting Scientist at the Smithsonian Institution, Washington.  I am grateful to the Andrew W. Mellon Foundation for making my visit possible (grant to Dr. James L. Luteyn) and to the Director and staff of the Botany Department, Smithsonian Institution for support and use of their facilities.  Thanks are also due to Gregorio Dauphin (San José) for providing additional records from Costa Rica and Panama, to Tamás Pócs (Eger) for additional records from Venezuela, to Andrea Lücking (Ulm) for loan of the type of Riccia indusiata, and to Antoine Cleef, Jim Luteyn, and Orlando Rangel for valuable suggestions. 

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