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Las Briofitas. Bolivia Ecologica 59
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Bryophytes of the Puna


Polylepis forest in humid puna. Photograph by C. Aldana.

The puna extends from northern Peu through Bolivia into portions of northwest Argentina and northern Chile. This is the major high elevational vegetation of the central Andes, including the area occupied by the vast Altiplano. The Bolivian puna is located in the southwestern portion of the country, occupying about 20% of the land surface. Elevation ranges from 3200 to more than 6000 meters, but much of the puna occur within a range of 3800 to 4200 meters.

Four broadly define puna zones in Bolivia can be recognized: humid, semihumid, dry and desert. A general climatic trend extends from humid and semihumid puna with higher precipitation (400-1600 mm) and fewer dry months (3-6) to dry and desert puna with higher precipitation (less than 50 to 400 mm) and a greater number of dry months (6-12). Likewise vegetation varies from dense grasslands and, at least originally, with open or closed forest of Polylepis in the humid and semihumid puna (but absent in the Altiplano plans), whereas the dry and desert vegetation is open or sparsely covered by grasses and shrubs, and what forest exist (or existed) are confined to protected valleys or canyon slopes.

Humid Puna. The humid puna occupies less than 1% of the land surface and ranging in elevation from 3800 to 4100 m. Annual average temperature ranges from 7-9C. Annual average rainfall 500-1600 mm, with approximately 3-5 dry months. This zone inclues the environs of Lake Titicaca and foothil.ls of the Cordillera Real. Vegetaation is dominated by shrub and grasslands. Important families include Asteraceae, Brassicaceae, Caryophyllaceae, Cyperaceae, Fabaceae, gentianaceae, Juncaceae, Poaceae and Rosaceae. Species include Ageratina azangaroense, Baccharis incarum, Buddleja coriacea, Chquiraga jussieui, Escallonia resinosa, Festuca dolichophylla, Geranium bangii, Lachemilla pinnata, Poa buchtienii, Polylepis besseri ssp. incarum, Satureja boliviana. Marsh species include Distichia muscoides and Oxychloe andina. This region is the most densely populated zone in the Altiplano, and has been under cultivation fro several thousand years; most of the original vegetation is no longer present.


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

Semihumid Puna. The semihumid puna occupies about 6% of the land surface, and ranges in elevation from 3200 to 4200 m. Annual average temperature ranges from 7-10C. Annual average rainfall 400-1100 mm, with approximately 4-6 dry months. Important families include: Asteraceae, Cactaceae, Caryophyllaceae, Fabaceae, Gentianaceae, Poaceae and Rosaceae. Characteristic species include Adesmia spinosissima, Baccharis boliviensis, Baccharis incarum, Buddleja coriacea, Deyeuxia hterophylla, Festuca dolichophylla, Polylepis besseri, Polylepis racemosa, Stipa leptostachya, and Tetraglochin cristatum. Also, as in the humid puna, the marsh species Distichia muscoides and Oxychloe andina are a commen element.

Dry Puna. The dry puna occupies 3% of the land surface, and ranges in elevation from 3500-4100 m. Annual average temperature ranges from 2-7C. Annual average rainfall 250-400 mm, with approximately 6 to 9 dry months. The vegetation is almost exclusively shrubland and thickets. Important families include Asteraceae, Cactaceae, Caryophyllaceae, Fabaceae, Poaceae, and Rosaceae. Marshy localities include the families Frankeriaceae and Junaceae. Important species include Adesmia spp., Baccharis boliviensis, B. incarum, Deyeuxia breviaristata, Fabiana densa, Festuca orthophylla, Parastrephia lepidophylla, Stipa curviseta, and Tetraglochin cristatum.

Desert Puna (and Cordillera Occidental). The desert puna, the largest puna zone, occupies about 9% of the land surface, and ranges in elevation from 3800 to over 6000 m. Annual average temperature ranges from less than 0 to 5C. Annual average rainfall less than 50 to 250 mm, with approximately 9-12 dry months. Important families include the Apiaceae, Asteraceae, Cactaceae, Caryophyllaceae, Chenopodiaceae, Frankeriaceae, Fabaceae, Poaceae, Rosaceae, and Verbenaceae. Important species include Baccharis boliviensis, Chaetanthera stuebelii, Deyeuxia crispa, Echinopsis spp., Festuca chrysophylla, Munroa decumbens, Oreocereus spp., Parastrephia lepidophylla, Polylepis tarapacana, Polylepis tomentella, and Stipa boumanii.

Prepuna. The prepuna occupies less than 1% of the land surface, and ranges in elevation from 2300-3400 m. Annual average temperature ranges from 15-17C with minimum at less than 0 to maximum at 30C. Annual average rainfall 300-400 mm, with appoximately 8-10 dry months. This vegetation zone extends into northwestern Argentina, and contains one of the highest levels of endemism with regard to the Cactaceae. Prepuna is found in western most Chuquisaca and Tarija, and eastern most Potosi. Vascular plant diversity is estimated at 324 species distributed among 200 genera and 61 families (Lopez & Beck 2002). The most diverse vascular plant families include the Asteraceae, Cactaceae, Poaceae, Bromeliaceae, and Leguminosae.


Subpuna-like vegetation west of Vallegrande, Department of Santa Cruz. Photograph by S. Churchill.

Bryophyte Diversity

The Bolivian puna contains about 338 species of mosses. Families of high diversity include the Pottiaceae (52 spp.), Bryaceae (39), Dicranaceae (34), Bartramiacae (29), Grimmiaceae (25), and Amblystegiaceae (16). Species rich genera include Campylopus (20 spp.), Bryum, Grimmia and Schizymenium (co-equal with 14 spp.), Leptodontium (14), Breutelia (10), Bartramia (9), and Syntrichia (8). The vast majority of mosses are terrestrial, occurring on soil, soil covered rocks or rocks. Very few epiphytes are present, mostly found on Polylepis (Rosaceae). The only family of note found in semi-aquatic or aquatic environs, i.e., bogs, marshes, margins of streams, ponds and lakes, is the Amblystegiaceae and Sphagnaceae. Hepatic diversity is estimated at 60 species.

Selected Literature

López, R.P. 2000. La prepuna boliviana. Ecologia en Bolivia 34: 45-70

López, R.P. & S. Beck. 2002. Phytogeographical affinities and life form composition of the Bolivian Prepuna. Candollea 57: 77-96.

Bryophyte Checklist of the Puna

Amblystegium tenax (Hedw.) C. E. O. Jensen
Amblystegium varium (Hedw.)Lindb.
Cratoneuron filicinum (Hedw.) Spruce
Drepanocladus aduncus (Hedw.) Warnst.
Drepanocladus longifolius (Mitt.) Broth. ex Paris
Drepanocladus polygamus (Schimp.) Hedenäs
Drepanocladus sordidus (Müll. Hal.) Hedenäs
Koponenia holoneuron (Herzog) Ochyra
Pseudocalliergon turgescens (T. Jensen) Loeske
Scorpidium cossonii (Schimp.) Hedenäs
Scorpidium scorpioides (Hedw.) Limpr.
Straminergon stramineum (Brid.) Hedenäs
Vittia elimbata Hedenäs, Vanderpoorten & Goffinet
Vittia pachyloma (Mont.) Ochyra
Warnstorfia exannulata (Schimp.) Loeske
Warnstorfia luipichensis (R. S. Williams) Hedenäs

Acroschisma wilsonii (Hook. f. & Wilson) A. Jaeger
Andreaea acutifolia Hook.f. & Wilson
Andreaea brevipes Spruce
Andreaea nitida Hook.f. & Wilson
Andreaea rupestris Hedw.
Andreaea subulata Harv. in Hook.

Aulacomnium palustre (Hedw.) Schwägr.

Anacolia laevisphaera (Taylor) Flowers
Bartramia ambigua Mont.
Bartramia brevifolia Brid.
Bartramia longifolia Hook.
Bartramia mathewsii Mitt.
Bartramia microstoma Mitt.
Bartramia pilicuspis Herzog
Bartramia polytrichoides Müll. Hal.
Bartramia potosica Mont.
Bartramia strumosa (Hampe) Mitt.
Breutilia austro-arcuta (Müll. Hal.) Paris
Breutilia brittoniae Renauld & Cardot
Breutilia harpophylla Herzog
Breutilia integrifolia (Taylor) A. Jaeger
Breutilia nigrescens Herzog
Breutilia polygastrica (Müll. Hal.) Broth.
Breutilia secundifolia (Müll. Hal.) Müll. Hal.
Breutilia subarcuata (Müll. Hal.) Schimp.
Breutilia tomentosa (Brid.) A. Jaeger
Breutilia trianae (Hampe) A. Jaeger
Conostomum clesitocarpum Herzog
Conostomum macrotheca Herzog
Conostomum pentastichum (Brid.) Lindb.
Flowersia setifolia (Hook. & Arn.) D.G. Griffin & W.R. Buck
Philonotis andina (Mitt.) A. Jaeger
Philonotis curvata (Hampe) A. Jaeger
Philonotis fontanella (Hampe) A. Jaegerr
Philonotis scabrifolia (Hook.f. & Wilson) Braithw.
Philonotis thwaitesii Mitt.

Brachythecium austrosalebrosum (Müll. Hal.) Paris
Brachythecium calliergonoides Broth.
Brachythecium cavifolium Herzog
Brachythecium occidentale (Hampe) A. Jaeger
Brachythecium praelongum Schimp. ex Müll. Hal.
Kindbergia praelonga (Hedw.) Ochyra
Platyhypnidium aquaticum (A. Jaeger) M. Fleisch.
Platyhypnidium validum (Herzog) Ochyra

Anomobryum humillimum (Müll. Hal. ex E. Britton) Broth.
Anomobryum julaceum (Schrad. ex P. Gaertn., B. Mey. & Scherb.) Schimp.
Anomobryum robustum Broth. in Herzog
Anomobryum steerei A. J. Shaw
Brachymenium acuminatum Harv. in Hook.
Brachymenium fabronioides (Müll. Hal.) Paris
Bryum algovicum Sendtn. ex Müll. Hal.
Bryum alpinum Huds. ex With.
Bryum argenteum Hedw.
Bryum capillare Hedw.
Bryum cellulare Hook.
Bryum challaense Broth.
Bryum crassinervum (Herzog) A. J. Shaw
Bryum ellipsifolium Müll. Hal.
Bryum imbricatum (Schwägr.) Bruch & Schimp.
Bryum laevigatum Hook.f. & Wilson
Bryum muehlenbeckii Bruch & Schimp.
Bryum pallescens Schleich. ex Schwägr.
Bryum paradoxum Schwägr.
Bryum pseudotriquetrum (Hedw.) P. Gaertn., B. Mey.& Scherb.
Leptobryum wilsonii (Mitt.) Broth.
Mielichhoferia cuspidata (Herzog) A. J. Shaw & H. A. Crum
Pohlia chilensis (Mont.) A. J. Shaw
Pohlia cruda (Hedw.)Lindb.
Pohlia wahlenbergii (F. Weber & Mohr) A. L. Andrews
Schizymenium ampullaceum (Broth.) A. J. Shaw
Schizymenium andinum (Sull.) A. J. Shaw
Schizymenium bogotense (Hampe) A. J. Shaw
Schizymenium bolivianum (Schimp. ex Müll. Hal.) A. J. Shaw
Schizymenium bryocarpum (Broth.) A. J. Shaw
Schizymenium campylocarpum (Hook. & Arn. in Hook.) A. J. Shaw
Schizymenium clavellatum (Mitt.) A. J. Shaw
Schizymenium cygnicollum (Schimp. ex. Müll. Hal.) A. J. Shaw
Schizymenium decurrens (Müll. Hal.) A. J. Shaw
Schizymenium latifolium (Herzog) A. J. Shaw
Schizymenium lindigii (Hampe) A. J. Shaw
Schizymenium nanum Taylor
Schizymenium pusillum (Hook.f. & Wilson) A. J. Shaw
Schizymenium subglobosum (R. S. Williams) A. J. Shaw

Catagonium brevicaudatum Müll. Hal.
Catagonium emarginatum S. H. Lin?

Amphidium tortuosum (Hornsch.) Cufod.
Aongstroemia julacea (Hook.) Mitt.
Camptodontium fallax (Herzog) Broth.
Camptodontium flexipes (Herzog) Broth.
Campylopodiella flagellacea (Müll. Hal.) J.-P. Frahm & Isoviita
Campylopus albidovirens Herzog
Campylopus amboroensis Thér.
Campylopus areodictyon (Müll. Hal.) Mitt.
Campylopus argyrocaulon (Müll. Hal.) Broth.
Campylopus cavifolius Mitt.
Campylopus dicnemoides (Müll. Hal.) Paris
Campylopus edithae Broth.
Campylopus flexuosus (Hedw.) Brid. var. incacorralis (Herzog) J. -P. Frahm
Campylopus griseus (Hornsch.) A. Jaeger var. ingeniensis (R. S. Williams) J.-P. Frahm
Campylopus huallagensis Broth.
Campylopus jugorum Herzog
Campylopus nivalis (Brid.) Brid. var. nivalis
Campylopus nivalis (Brid.) Brid. var. multicapsularis (Müll. Hal.) J.-P. Frahm
Campylopus perexilis (Müll. Hal.) Paris
Campylopus pilifer Brid. var. pilifer 
Campylopus pittieri R.S. Williams
Campylopus reflexisetus (Müll. Hal.) Broth.
Campylopus richardii Brid.
Campylopus subjugorum Broth.
Campylopus trichophylloides Thér. in Herzog
Campylopus trivialis Müll. Hal. ex E. Britton
Chorisodontium nigricans (Herzog) Broth.
Chorisodontium speciosum (Hook.f. & Wilson) Broth. var. speciosum
Dicranella campylophylla (Taylor) A. Jaeger
Dicranoweisia brunnea Herzog
Hygrodicranum bolivianum Herzog
Oreoweisia brasiliensis Hampe?
Oreoweisia erosa (Müll. Hal.) Kindb.
Pilopogon guadalupensis (Brid.) J.-P. Frahm
Pilopogon laevis (Taylor) Thér.
Pilopogon tiquipayae Herzog
Polymerodon andinus Herzog
Rhabdoweisia fugax (Hedw.) Bruch & Schimp.

Astomiopsis amblyocalyx Müll. Hal.
Astomiopsis subulata Müll. Hal.
Ceratodon stenocarpus Bruch & Schimp.
Chrysoblastella chilensis (Mont.) Reimers
Distichium capillaceum (Hedw.) Bruch & Schimp.
Ditrichum bogotense (Hampe) Broth.
Ditrichum crinale (Taylor) Kuntze
Ditrichum gracile (Mitt.) Kuntze
Ditrichum submersum Cardot & Herzog
Pleuridium andinum Herzog
Tristichium lorentzii Müll. Hal.
Tristichium mirabile (Müll. Hal.) Herzog

Encalypta asperifolia Mitt.
Encalypta ciliata Hedw.
Encalypta rhaptocarpa Schwägr.

Fissidens andicola (Herzog) Brugg.-Nann.
Fissidens crispus Mont.
Fissidens curvatus Hornsch.
Fissidens excurrentinervis R.S. Williams
Fissidens incisus Herzog
Fissidens rigidulus Hook.f. & Wilson
Fissidens wallisii Müll. Hal.

Entosthodon bonplandii (Hook.) Mitt.
Entosthodon jamesonii (Taylor) Mitt.
Entosthodon laevis (Mitt.) Fife
Entosthodon laxus (Wilson in Hook. f.) Mitt.
Entosthodon lindigii (Hampe) Mitt.
Entosthodon subaloma (Herzog) S.P. Churchill

Neosharpiella turgida (Mitt.) H. Rob. & Delgad.

Coscinodon bolivianus Broth.
Coscinodontella bryanii R.S. Williams
Grimmia anodon Bruch & Schimp.
Grimmia atrata Miel. ex Hornsch.
Grimmia austrofunalis Müll. Hal.
Grimmia bicolor Herzog
Grimmia donniana Sm.
Grimmia elongata Kaulf.
Grimmia fuscolutea Hook.
Grimmia longirostris Hook.
Grimmia navicularis Herzog
Grimmia pseudoanodon Deguchi
Grimmia reflexidens Müll. Hal.
Grimmia tergestina Tomm. ex Bruch & Schimp.
Grimmia trichophylla Grev.
Grimmia trinervis R.S. Williams
Jaffueliobryum wrightii (Sull.) Thér.
Ptychomitrium chimborazense (Spruce ex Mitt.) A. Jaeger
Racomitrium crispipilum (Taylor) A. Jaeger
Racomitrium cucullatifolium Hampe
Racomitrium lanuginosum (Hedw.) Bird.
Racomitrium subsecundum (Hook. & Grev.) Wilson
Schistidium andinum (Mitt.) Herzog
Schistidium apocarpum (Hedw.) Bruch & Schimp.
Schistidium falcatum (Hook.f. & Wilson) B. Bremer

Hedwigidium integrifolium (P.-Beauv.) Dixon in C. E. O. Jensen

Hypnum amabile (Mitt.) Hampe
Hypnum cupressiforme Hedw. var. lacunosum Brid.

Lepyrodon tomentosus (Hook.) Mitt.

Leptopterigynandrum austro-alpinum Müll. Hal.
Leptopterigynandrum subintegrum (Mitt.) Broth.
Leptopterigynandrum tenellum Broth.
Leptopterigynandrum tenuicaule (R.S. Williams) S. He

Orthotrichum cupulatum Brid.
Orthotrichum psychrophilum Mont.
Orthotrichum rupestre Schleisch. ex Schwägr.
Orthotrichum steerei Lewinsky
Zygodon pichinchensis (Taylor) Mitt.

Notoligotrichum trichodon (Hook.f. & Wilson) G. L. Sm.
Oligotrichum erosum (Hampe) Lindb.
Pogonatum campylocarpon (Müll. Hal.) Mitt.
Polytrichadelphus aristatus (Hampe) Mitt.
Polytrichastrum tenellum (Müll. Hal.) G. L. Sm.
Polytrichum juniperinum Hedw.
Psilopilum gymnostomulum (Müll. Hal.) Paris

Aloinella boliviana Broth.
Aloinella cucullifera (Mitt.) Steere
Anoectangium aestivum (Hedw.) Mitt.
Bryoerythrophyllum bolivianum (Müll. Hal.) R. H. Zander
Bryoerythrophyllum campylocarpum (Müll. Hal.) H. A. Crum
Bryoerythrophyllum fuscinervium (Mitt.) R.H. Zander
Bryoerythrophyllum jamesonii (Taylor) H. A. Crum
Didymodon australasiae (Hook. & Grev.) R.H. Zander
Didymodon challaense (Broth.) R.H. Zander
Didymodon herzogii R.H. Zander
Didymodon laevigatus (Mitt.) R. H. Zander
Didymodon pruinosus (Mitt.) R. H. Zander
Didymodon vinealis (Brid.) R.H. Zander
Erythrophyllastrum andinum (Sull.) R.H. Zander
Erythrophyllopsis fuscula (Müll. Hal.) Hilp.
Hennediella angustifolia (Herzog) R.H. Zander
Hennediella denticulata (Wilson) R.H. Zander
Hennediella limbata (Mitt.) R.H. Zander Puna.
Hennediella polyseta (Mitt.) R.H. Zander
Hymenostylium recurvirostre (Hedw.) Dixon
Leptodontium brachyphyllum Broth. & Thér. in Thér.
Leptodontium capituligerum Müll. Hal.
Leptodontium erythroneuron Herzog
Leptodontium flexifolium (Dicks. ex With.) Hampe in Lindb.
Leptodontium longicaule Mitt. var. longicaule
Leptodontium longicaule var. microruncinatum (Dusén) R.H. Zander
Leptodontium planifolium Herzog
Leptodontium proliferum Herzog
Leptodontium pungens (Mitt.) Kindb.
Leptodontium subintegrifolium Thér. & Herzog
Leptodontium tricolor (R.S. Williams) R.H. Zander
Leptodontium viticulosoides (P. Beauv.) Wijk & Margad. var. exasperatum (Cardot) R.H. Zander
Leptodontium wallisii (Müll. Hal.) Kindb.
Mironia ehrenbergiana (Müll. Hal.) R.H. Zander
Molendoa sendtneriana (Bruch & Schimp. in Bruch, Schimp. & W. Gümbel) Limpr.
Pseudocrossidium apiculatum R.S. Williams
Pseudocrossidium chilense R.S. Williams
Pseudocrossidium elatum (R.S. Williams) Delgad.
Pseudocrossidium pachygastrellum (Herzog) Broth.
Pseudocrossidium replicatum (Taylor) R.H. Zander
Rhexophyllum subnigrum (Mitt.) Thér. ex Hilp.
Sagenotortula quitoensis (Taylor in Hook.) R.H. Zander
Syntrichia aculeata (Wilson) R.H. Zander
Syntrichia andicola (Mont.) Ochyra
Syntrichia fragilis (Taylor) Ochyra
Syntrichia obtusissima (Müll. Hal.) R.H. Zander
Syntrichia percarnosa (Müll. Hal.) R.H. Zander
Syntrichia princeps (De Not.) Mitt.
Syntrichia ramosissima (Thér.) R.H. Zander
Syntrichia ruralis (Hedw.) F. Weber & D. Mohr var. ruralis
Syntrichia ruralis var. spiralis (Herzog) R.H. Zander
Trichostomum tenuirostre (Hook. & Taylor) Lindb.

Rhacocarpus apiculatus (Renauld & Cardot) Paris
Rhacocarpus excisus (Müll. Hal.) Paris
Rhacocarpus purpurascens (Brid.) Müll. Hal.

Rhytidium rugosum (Hedw.) Kindb.

Blindia magellanica Schimp.

Sphagnum cuspidatum Ehrh. ex Hoffm.
Sphagnum limbatum Mitt.
Sphagnum magellanicum Brid.
Sphagnum meridense (Hampe) Müll. Hal.
Sphagnum recurvum P. Beauv.

Thuidium peruvianum Mitt.

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