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Las Briofitas. Bolivia Ecologica 59
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Bryophytes of the Amazon

Amazon forest with palms. Photograph by A. Araujo M.

The Amazon is located in the north and northeast of Bolivia. This region, referred to as the Southwest Amazon by Ibisch et al. (2003, 2004), contains ca. 22% of the land surface of Bolivia. Elevation ranges from 100 to 500 meters. Three subregions are recognized: Pando Amazon Forest (at elevations from 100-500 m), Beni-Santa Cruz Amazon Forest (overlying the Brazilian Shield, at elevations from 150-400 m), and Flooded Amazon Forest (várzea and igapó, at elevations from 100-500 m). The Floodplain Savannas, are also included here under the Amazon region, occupying 12% of the Bolivian land surface. The Moxos Floodplain Savannas, the larger of the two subregions, is characetized by extensive grasslands and gallery forests, the other, the Pantanal Flooded Savannas are found along and extending into Brazil in the extreme eastern portion of the country. The total land surface area of the Amazon region, including the flooded savannas, occupies about half of the Bolivian eastern lowlands (Oriente).

The Amazon basin forest, the largest vegetation represented in the Neotropics, reaches its southwestern range in Bolivia. Amazonian forest (Pando, Beni-Santa Cruz, and Floodplain) contains the core of bryophytes that are widespread through the basin. The savanna ecoregions are likely to exhibit greater affinities with the Amazon bryophyte flora but there is insufficient data at present. The Moxos Floodplain Savannas, particularly the gallery forest, and the Pantanal Flooded Savannas are associated with the Amazon. The Cerrado ecoregion of both La Paz and Beni may have an affinity with the Amazon but neither have been explored.

Amazon is a highly diverse region, particularly with regard to trees. Knowledge of vascular plant diversity in the Amazon, however, is still very incomplete. As with bryophytes, greater inventory efforts are required. The important vascular plant families (mostly trees) include: Annonaceae, Arecaceae, Burseraceae, Combretaceae, Eurphorbiaceae, Lauraceae, Lecythidaceae, Leguminosae, Meliaceae, Moraceae, Sapotaceae, and Vochysiaceae. Tree diversity ranges from 650 to more than 1000 species. Vascular epiphyte diversity is estimated at 150 to 200 species represented primarily by the Orchidaceae and Araceae.

Map of the Amazon ecoregion. Courtesy of FAN (Fundacion Amigos de la Naturaleza), Santa Cruz, Bolivia. Graphics modified by E. Calzadilla.

Bryophyte Diversity

There are presently recorded 102 species of mosses distributed among 55 genera and 23 families for the Bolivian Amazon region. This accounts for approximately 30% of the species known from the Amazon basin (Churchill 1998). It is estimated that the Amazon ecoregion contains ca. 10% of the total species diversity for Bolivia. Species rich families include: Fissidentaceae (13 spp.), Calymperaceae (12 spp.), Sematophyllaceae (12 spp.), Pilotrichaceae (9 spp.), and coequal at 7 spp. Hypnaceae and Macromitriaceae. Two genera are notably diverse, Fissidens (13 spp.) and Syrrhopodon (8 spp.). Knowledge of hepatics is still very incomplete. At present 14 species, 11 genera and 7 families are recorded. A minimum of 50 species can be expected for the Bolivian Amazon. This estimation is extrapolated from a bryophyte inventory by Pinheiro da Costa (2003) for the state of Acre in Brazil (on the border with Bolivia) where 46 species of hepatics have been recorded.

The first notable reports of bryophytes from the Bolivian Amazon were made by Williams (1903, 1910); see Dorr (1991) for site localities visited by R. S. Williams, in particular Río Mapiri and Tumupasa. Reese (1979) provided several additional records for the Bolivian lowlands.


Epiphytes are almost exclusively found on trees, less frequent or rare on lianas and shrubs. On trees epiphytes are found most frequently at the base, buttress planks, and less often on trunks and exposed roots. In general the canopy of the Amazonian forest have not been sampled.

Hepatics: Bazzania phyllobola, Bryopteris filicina, Lejeunea sp., Leucolejeunea sp., Plagiochila subplana, Xylolejeunea crenata. Mosses: Calymperes afzelii, Calymperes palisotii, Crossomitrium patrisiae (epiphyllous), Crossomitrium saprophilum, Donnellia commutata, Groutiella obtusa, Groutiella tomentosa, Groutiella tumidula, Henicodium geniculatum, Holomitrium arboreum, Lepidopilum surinamense, Leucobryum martianum, Leucobryum subobtusifolium (rare as an epiphyte), Macromitrium stellulatum, Meteorium nigrescens (rare), Neckeropsis disticha, Neckeropsis undulata, Octoblepharum albidum, Octoblepharum cocuiense, Octoblepharum pulvinatum, Pelekium scabrosulum, Pilosium chlorophyllum, Pilotrichum evanescens, Pireella pohlii, Porotrichum substriatum, Potamium lonchophyllum (exposed tree roots in stream), Pseudocryphaea domingensis, Rhacopilopsis trinitensis, Schlotheimia jamesonii, Schlotheimia torquata, Sematophyllum subsimplex, Syrrhopodon cryptocarpus, Syrrhopodon hornschuchii, Syrrhopodon incompletus var. incompletus, Syrrhopodon parasiticus, Taxithelium planum, Zelometeorium patulum.


Logs, including standing dead trees, are the second most diverse substrate for bryophytes in the Amazon forest. Typically wood in the form of trunks and branches decompose very slowly, due to the hardness of the wood. Corticulous logs are usually more diverse than decorticulous logs.

Hepatics: Bryopteris filicina (rare on logs), Pallavicinia lyellii, Plagiochila disticha, Plagiochila montagnei, Riccardia cf. digitiloba, Telaranea nematodes. Mosses: Callicostella pallida, Calymperes afzelii, Chryso-hypnum diminutivum, Donnellia commutata, Ectropothecium leptochaeton, Fissidens guianensis var. guianensis, Fissidens lagenarius var. muriculatus, Fissidens pellucidens, Isopterygium tenerum, Leucobryum martianum, Leucobryum subobtusifolium, Leucomium strumosum, Neckeropsis disticha, Neckeropsis undulata, Pelekium involvens, Pelekium scabrosulum, Pelekium schistocalyx, Pterogonidium pulchellum, Sematophyllum subsimplex, Syrrhopodon cryptocarpus, Syrrhopodon incompletus var. incompletus, Taxithelium planum, Trichosteleum subdemissum, Vesicularia vesicularis.


Soil substrate supports few bryophytes, most notably and diverse is the genus Fissidens.

Mosses: Brachymenium coarctatum, Callicostella pallida, Dicranella hilariana, Fissidens allionii, Fissidens angustifolius, Fissidens pellucidens, Fissidens scariosus, Fissidens zollingeri, Microdus sp., Philonotis uncinata, Vesicularia vesicularis.

Selected Literature

Churchill, S. P. 1998. Catalog of Amazonian mosses. J. Hattori Bot. Lab. 85: 191-238.

Dorr, L. J. 1991. The vascular plant collections of R. S. Williams from Bolivia and Peru (1901-1902). Brittonia 43: 211-239.

Reese, W. D. 1979. New records of Bolivian mosses. Phytologia 43: 337-338.

Costa, D. 2003. Floristic composition and diversity of Amazonian rainforest bryophytes in the state of Acre, Brazil. Acta Amazonica 33: 399-414.

Williams, R. S. 1903. Bolivian mosses, I. Bull. New York Bot. Gard. 3: 104-134.

Williams, R. S. 1910. Bolivian mosses, II. Bull. New York Bot. Gard. 6: 227-261.

Checklist of the Amazon Bryophytes


Riccardia cf. digitiloba (Spruce ex Steph.) Pagán
Cephalozia crussifolia (Lindenb. & Gotts.) Fulford
Lophocolea bidentata (L.) Durmort.
Lophocolea sp.
Ceratolejeunea cornuta (Lindenb.) Schiffner
Lejeunea spp.
Leucolejeunea sp.
Xylolejeunea crenata (Mont.) X.-L. He & Grolle
Bazzania phyllobola Spruce
Telaranea nematodes (Austin) M. Howe
Pallavicinia lyellii (Hook.) Carruth.
Plagiochila disticha (Lehm. & Lindenb.) Lindenb.
Plagiochila montagnei Nees
Plagiochila subplana Lindenb.


Philonotis hastata (Duby) Wijk & Margad.
Philonotis uncinata (Schwägr.) Brid.
Zelometeorium patulum (Hedw.) Manuel
Brachymenium coarctatum Bosch & Sande Lac.
Bryum apiculatum Schwägr.
Calymperes afzelii Sw.
Calymperes erosum Müll. Hal.
Calymperes nicaraguense Renauld & Cardot (rare)
Calymperes palisotii Schwägr.
Syrrhopodon cryptocarpus Dozy & Molk.
Syrrhopodon gaudichaudii Mont.
Syrrhopodon hornschuchii Mart.
Syrrhopodon incompletus Schwägr. var. incompletus
Syrrhopodon ligulatus Mont.
Syrrhopodon parasiticus (Sw. ex Brid.) Paris
Syrrhopodon simmondsii Steere
Syrrhopodon xanthophyllus Mitt.
Schoenobryum concavifolium (Griff.) Gangulee (marginal)
Campylopus savannarum (Müll. Hal.) Mitt.
Campylopus surinamensis Müll. Hal.
Dicranella hilariana (Mont.) Mitt.
Holomitrium arboreum Mitt.
Leucoloma tortellum (Mitt.) A. Jaeger
Microdus cf. exiguus (Schwägr.) Besch.
Fissidens allionii Broth.
Fissidens angustifolius Sull.
Fissidens crispus Mont.
Fissidens elegans Brid.
Fissidens flaccidus Mitt.
Fissidens guianensis Mont. var. guianensis
Fissidens inaequalis Mitt.
Fissidens lagenarius var. muriculatus (Spruce ex Mitt.) Pursell
Fissidens pallidinervis Mitt.
Fissidens pellucidens Hornsch. var. pellucidens
Fissidens pellucidens Hornsch. var. papilliferus (Broth.) Pursell
Fissidens scariosus Mitt.
Fissidens submarginatus Bruch in C. Krauss
Fissidens zollingeri Mont.
Chryso-hypnum diminutivum (Hampe) W.R. Buck
Ectropothecium leptochaeton (Schwägr.) W.R. Buck
Isopterygium tenerifolium Mitt.
Isopterygium tenerum (Sw.) Mitt.
Pterogonidium pulchellum (Hook.) Müll. Hal.
Rhacopilopsis trinitensis (Müll. Hal.) E. Britton & Dixon
Vesicularia vesicularis (Schwägr.) Broth.
Orthostichella rigida (Müll. Hal.) B.H. Allen & Magill (marginal)
Pseudocryphaea domingensis (Spreng.) W. R. Buck
Leucobryum martianum (Hornsch.) Hampe
Leucobryum subobtusifolium (Broth.) B. H. Allen
Ochrobryum gardneri (Müll. Hal.) Lindb.
Ochrobryum subulatum Hampe in Besch.
Leucomium strumosum (Hornsch.) Mitt.
Groutiella obtusa (Mitt.) Florsch.
Groutiella tomentosa (Hornsch.) Wijk & Margad.
Groutiella tumidula (Mitt.) Vitt
Macromitrium stellulatum (Hornsch.) Brid.
Schlotheimia jamesonii (Arn.) Brid.
Schlotheimia rugifolia (Hook.) Schwagr.
Schlotheimia torquata (Hedw.) Brid.
Meteorium nigrescens (Hedw.) Mitt.
Neckeropsis disticha (Hedw.) Kindb.
Neckeropsis undulata (Hedw.) Reichardt
Porotrichum substriatum (Hampe) Mitt.
Octoblepharum albidum Hedw.
Octoblepharum cocuiense Mitt.
Octoblepharum pulvinatum (Dozy & Molk.) Mitt.
Callicostella depressa (Hedw.) A. Jaeger
Callicostella pallida (Hornsch.) ångstr.
Callicostella rivularis (Mitt.) A. Jaeger
Crossomitrium patrisiae (Brid.) Müll. Hal.
Crossomitrium saprophilum Broth.
Lepidopilum affine Müll. Hal.
Lepidopilum scabrisetum (Schwägr.) Steere
Lepidopilum surinamense Müll. Hal.
Pilotrichum evanescens (Müll. Hal.) Crosby
Barbula arcuata Griff.
Barbula indica var. gregaria (Mitt.) R.H. Zander
Hyophila involuta (Hook.) A. Jaeger
Henicodium geniculatum (Mitt.) W. R. Buck
Jaegerina scariosa (Lorentz) Arzeni
Orthostichidium quadrangulare (Schäwgr.) B.H. Allen & Magill (marginal)
Orthostichopsis praetermissa W. R. Buck (marginal)
Pireella pohlii (Schwägr.) Cardot
Racopilum tomentosum (Hedw.) Brid.
Acroporium pungens (Hedw.) Broth. (marginal)
Donnellia commutata (Müll. Hal.) W.R. Buck
Hydropogon fontinaloides (Hook.) Brid.
Hydropogonella gymnostoma (Schimp.) Cardot
Potamium lonchophyllum (Mont.) Mitt.
Sematophyllum galipense (Müll. Hal.) Mitt. (marginal)
Sematophyllum subpinnatum (Brid.) E. Britton
Sematophyllum subsimplex (Hedw.) Mitt.
Taxithelium planum (Brid.) Mitt.
Trichosteleum ambiguum (Schwägr.) Paris
Trichosteleum papillosum (Hornsch.) A. Jaeger
Trichosteleum subdemissum (Schimp. ex Besch.) A. Jaeger
Entodontopsis angustiretis (Broth.) W. R. Buck & Ireland
Entodontopsis leucostega (Brid.) W. R. Buck & Ireland
Eulacophyllum cultelliforme (Sull.) W. R. Buck & Ireland (marginal)
Pilosium chlorophyllum (Hornsch.) Müll. Hal.
Pelekium involvens (Hedw.) Touw (syn. Cyrto-hypnum)
Pelekium scabrosulum (Mitt.) Touw
Pelekium schistocalyx (Müll. Hal.) Touw

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