ࡱ> @ VEbjbj "xuuV=<<<<,0<e<<<<<<<<<_eaeaeaeaeaeae$gRnileM<<MMe<<eQ\Q\Q\M <<_eQ\M_eQ\Q\:Sa,a<< ~DwG<Za ade0eaRiQ\iaia <8BQ\EHS<<<ee Q\ 4Terrestrial habitat types 4.7. Montane grassland (pramo and puna) General description and geographic variation Montane grasslands are found on mountain summits with an elevation range approximately between 9,800 16,400 feet (3000 5000 m), above the tropical forest limit and up to the upper limit of permanent snow. One characteristic of montane tropical climate is the sharp difference between diurnal and nocturnal temperatures, up to 360F (200C) during the dry season. In the Andes, the climate is more seasonal towards the south and annual precipitation decreases as one moves south. The vegetation of the pramos develops in a humid environment due to frequent rain, fog and snow, from Venezuela to the far North of Peru. Puna vegetation is found on high plateaus of the dry Andes, from Peru southwards. Pramo: It is identified by a special type of vegetation bunchgrasses, rosette plants, evergreen bushes with coriaceous and sclerophyllic leaves and cushion plants. It has high biodiversity with a very high level of endemism. The pramo flora is the richest high mountain flora of the world (Luteyn, 1999). It is normally accepted that pramos extend from Costa Rica to Peru. By the end of this exercise we will include vegetation similar to that of the pramo, for example, montane grasslands of southeastern Mexico, Guatemala, Bolivia, north of Argentina, and Brazil. The diversity and biotic richness of pajonales and grasslands is higher in the zonal than in the azonal vegetation. Some variation in pramo composition is based on the origin and migration of species. In general, diversity is higher in the northern Andean zone. Both variety and diversity, as well as structural complexity of mountain shrublands varies from the Mexican volcanoes to the Peruvian puna. Northern Andean communities are richer and more diverse. On the mountain pajonales, continuous grazing and use of fire for grazing create a tendency towards uniformity of structural pattern. Puna: It is normally considered to be found throughout the Peruvian Andes to northern Argentina. The diversity and biotic richness of pajonales and grasslands is greater in zonal than azonal vegetation. Some variation in floristic composition of the puna is based on quantity and north-south/east-west precipitation gradient. In general, diversity increases to the north and east. Community types/zonation and major gradients within the system (patterns) Pramos: Chuscales are bamboo areas dominated by species of Chusquea (e.g., Chusquea tessellata y C. subtesellata.) y Neurolepis. Shrublands dominated by species of Asteraceae (Diplostephium sp., Chuquiraga sp.), Hypericum sp. and Ericaceae. Bosquecitos (little forests) of Polylepis or Comarostaphylos in the pramo. Pajonales dominated by bunchgrasses, such as Calamagrostis spp. (C. effusa, C. bogotensis), Festuca, and Stipa. Frailejonales dominated by species of the genera Espeletia, Espeletiopsis, Coespeletia, and Libanothamnus. Wetlands and peat bogs dominated by species of the genera Sphagnum, Juncus, Carex, Isoetes, and Plantago. Superpramo over rocky periglaciar areas dominated by lichens and Asteraceae. Puna: According to Luteyn and Churchill (2000), there are three types of puna vegetation:wet puna, dry puna and arid puna. Wet Puna: is found in areas with annual precipitation between 16 59 inches (400 1500 mm), with vegetation dominated by Calamagrostis, Cortaderia, Festuca and Stipa. In wet areas one can find peat bogs (turberas) dominated by Cyperaceae y Juncaceae. Dry Puna: is found in areas with annual precipitation between 4 16 inches (100 400 mm), with vegetation dominated by tola shrubs (Lepidophyllum quadrangulare, Parastrephia spp.) and bunchgrasses (Festuca orthophylla). Arid Puna: is found in areas with less than 4 inches (100 mm) of annual precipitation. Vegetation is sparse and xerophilous, dominated by cacti (Cereus, Oreocereus), thorn bushes, cushion plants and bunchgrasses. Ecological integrity factors for landscape context Table xxx. Ecological integrity factors for landscape context of montane grassland (pramo and puna) Key FactorsJustification for factor selectionMinimum Integrity Threshold(s)Justification for determining the threshold (e.g., Natural Range of Variation)Field-based monitoring indicatorsFactor PriorityClimate regimeDetermining the distribution of the dominant flora Annual precipitation: 0.4 >24 inches (10 - >600 mm). (Arroyo et al., 1988) There is seasonality (wet due to summer snow, the season when sun is high: dry in winter, the season when sun is low on the horizon.) Evapotranspiration Rate:??. [Experts, please provide an estimate or literature citation. Thanks.] Average annual temperature: 460F (80C). Annual range of average monthly temperature < 34-36 0F (1-20C) : Horn (1989) includes data for the Cerro Pramo meteorological station in the Buenavista pramo of Costa Rica that show annual range to be less than 340F (10C) at that site. Maximum daily temperature fluctuation: up to 720F (400C) [Experts, please provide literature citation. Thanks.] A main determinant of geographic distribution.Precipitation and temperature data. Daily tracking of cloud cover.Vegetation continuity across de altitudinal gradient. Maintain habitat for typical species.?It can be measured and characterized with land models.Measurement of vegetation cover combined with abiotic characteristics such as elevation and soils. Presence of forest fauna species.Pramo: Vegetation continuity and soil permeability level that prevent excessive drainage.Control water storage capacity. Maintain pramo structure, composition and function to sustain the systems hydraulic properties.When the pramo starts to drain, there is lack of functionality. Disappearance of streams and other bodies of water in the pramo.Pramo: Depth of the soils organic horizon.Pramo: To influence the quantity of carbon absorption.Pramo: <12 inches (<30 cm) (depends on the substrate) [Experts, please provide literature citation. Thanks.]Pramo: Natural range of variation: 12 inches ~ 8 feet (30 cm ~ 2.5 m). [Experts, please provide literature citation. Thanks.] Puna: Natural disturbance regime. Puna y Pramo: Many species require disturbance to regenerate. Slow recruitment and growth.Puna: Recruitment of new individuals through seeds or seedlings.Fire regime.Pramo and puna: Fires affect vegetation composition and dry organic matter. Puna: More frequent in the wet puna (more inflammable). Forage production.Fire frequency (every xx years): for Costa Rica Pramo: 6-30 years (Horn, 1991 & 1998). Size of burned area (xxx hectares) Fire intensity (amount of energy by unit of time) (Reference - Analysis of a single year by Grau 2001). [Experts, please complete the citation in the References section.]Pramo: There is a Colombian group working with fire regime in the Chingaza pramo and also a study by Univ. Catlica and others in Ecuador. [Experts, please complete the citation in the References section.] Puna: There are no studies to determine if fires occurred before human colonization.Depth of fire on soils. Quantity of dry organic matter. Burned surface. Period between two consecutive fires.Aeolian and fluvial erosion. Natural regulation of vegetation composition. For its effect combined with human activities, like mechanized agriculture and overgrazing (more open soil and more impact from human activities).Highly diminished capacity for sustainability of agriculture and cattle raising activities.Lieberman about use of tractors. [Experts, please complete the citation in the References section. Thanks.]Erosion rate observations. Ecological integrity factors for condition Table xxx. Ecological integrity factors for condition of montane grassland (pramo and puna) Key FactorsJustification for Factor SelectionEcological Thresholds: Min. Dynamic Area Desired Future Condition (Increase in MDA to Rate Good or Very Good) Justifications or Recommendations for Calculating Minimum Dynamic Area (MDA) and Desired Sizes above MDAIndicators for Field-Based MonitoringFactor PriorityPresence of key plant communities.To maintain pollinator populations. Important to maintain environmental heterogeneity.Minimum pollinator density? Measures of environmental heterogeneity.Use methodology published by Arroyo et al., (1985) to quantify pollination. Natural herbivory by native fauna. (Pramos: deer, rodents, Andean bear, mountain tapir, and agouti. Puna: camelids.)It affects structure and functions (nutrient cycles) and composition (natural regulator).Optimum density can be defined for each animal species. [Experts, if you know of any published studies, please cite. Thanks.]Studies describing key species diet to clarify the effect on composition and structure. [Experts, if you know of any published studies, please cite. Thanks.]Presence of trampling (particularly on resistant species), browsing, animal scat, direct observations, and evidence of hunting.Herbivory by introduced species (Pramos: cattle, Puna: burro and sheep cause most of the damage).It affects structure and functions (nutrient cycles) and composition (natural regulator). Leads to destruction of affected areas of pramo in a short time. Fire regime changes with presence of cattle, hunting of predators along with cattle presence, introduction of exotic plants and soil compactation.% of altered surface. Load capacity calculation.Presence of trampling (with preference to resistant species), browsing, animal scat, direct observations. Perennial herbs are very susceptible to cattle over-grazing. Puna: increase of thorn bushes.Key Factors Justification for factor selectionMinimum Integrity Threshold(s)Justification for determining the threshold (e.g., Natural Range of Variation)Field-based monitoring indicators Factor PriorityPuna: Presence of spring-fed cushion bogs. Water source for animals and habitat for migrant birds. Stream flow regulation. Over-use by mining industry.There are studies about water load in the system and its extraction by mining industry. [Experts, please provide literature citation. Thanks.]Size of the spring-fed cushion bog, carcavas (trenches). Peat extraction. Use by animals, such as lamas and alpacas. Bog indicators: Juncacea, Distichlia sp. Recommended priorities for conservation-driven research agenda and next steps Sally Horn: 1) Further research on the long-term history of climate, vegetation, and human impacts in pramo and puna environments. Were there fires before the arrival of people? How did post-glacial changes in vegetation affect fire regimes? 2) Additional research on modern and historic treeline dynamics, and on the dynamics of Polylepis and Comarostaphylis islands within pramos. These bosquecitos are often regarded as indicating a formerly much higher treeline, but little detailed field evidence is available to support this supposition. 3) Research on changes in species distributions and treeline in response to global warming and to increased carbon dioxide concentrations directly. Literature Cited Arroyo, M.T.K., J.J. Armesto and R.B. Primack. 1985. Community studies in pollination ecology in the high temperature Andes of central Chile. II. Effect of temperature and visitation rates and pollination possibilities. Plant Systematics and Evolution 149: 187-203. Arroyo, M.T.K., F.A. Squeo, J.J. Armesto and C. Villagrn. 1988. Effects of aridity on plant diversity in the northern Chile Andes: results of a natural experiment. Annals of the Missouri Botanical Garden 75: 55-78. Grau 2001, from a single year. Horn, S.P. 1998. Fire Management and Natural Landscapes in the Chirrip Pramo, Chirrip National Park, Costa Rica. Pp. 125146. In Zimmerer, K. and Young, K. (Eds.), From Nature's Geography: Biogeographical Landscapes and Conservation in Developing Countries. Madison, Wisconsin: University of Wisconsin Press. Horn, S.P. 1991 Fire History and Fire Ecology in the Costa Rican pramos. pp 289-296. In: S.C. Nodvin and T.A. Waldrop (eds.), Fire and the environment: Ecological and cultural perspectives. Proceedings of an International Symposium, March 20-24, 1990, Knoxville, TN. U.S.D.A. Forest Service Gen. Tech. Rep. SE-69. Horn, S.P. 1989. Postfire vegetation development in the Costa Rican Pramos. Madroo 36(2): 93114. League, B. L. and S. P. Horn. A 10000 years [no s] record of Pramo fires in Costa Rica. Journal of Tropical Ecology (2000) 16: 747-752. Lieberman Luteyn, J.L. 1999. Pramos: a checklist of plant diversity, geographical distribution, and botanical literature. Mem. New York Bot. Gard. V. 84. 278 pp. New York, USA. Luteyn, J. L. and S. P. Churchill. 2000. Vegetation of the tropical Andes. Pp. 281-310. In D. L. Lentz (ed.) Imperfect balance: landscape transformations in the Precolumbian Americas. Columbia University Press, New York. Recommended resources Balslev, H. and J.L. Luteyn. (eds.) 1992. Pramo: An Andean ecosystem under human influence. Academic Press. New York. Hofstede, R. 1995. Effects of burning and grazing on a Colombian pramo ecosystem. Ph.D. Thesis University of Amsterdam: 190 pp. Amsterdam. Horn, S.P. 1998. Fire Management and Natural Landscapes in the Chirrip Pramo, Chirrip National Park, Costa Rica. Pp. 125146. In Zimmerer, K. and Young, K. (Eds.), From Nature's Geography: Biogeographical Landscapes and Conservation in Developing Countries. Madison, Wisconsin: University of Wisconsin Press. Horn, S.P. 1991 Fire History and Fire Ecology in the Costa Rican pramos. pp 289-296. In: S.C. Nodvin and T.A. Waldrop (eds.), Fire and the environment: Ecological and cultural perspectives. Proceedings of an International Symposium, March 20-24, 1990, Knoxville, TN. U.S.D.A. Forest Service Gen. Tech. Rep. SE-69. League, B. L. and S. P. Horn. A 10000 years [no s] record of Paramo fires in Costa Rica. Journal of Tropical Ecology (2000) 16: 747-752 Luteyn, J.L. 1999. Pramos: a checklist of plant diversity, geographical distribution, and botanical literature. Mem. New York Bot. Gard. V. 84. 278 pp. New York, USA. Molinillo, M. F. and M. Monasterio. 1997. Pastoralism in pramo environments: practices, forage, and impact on vegetation in the cordillera de Mrida, Venezuela. Mountain Res. Developm. 17: 197-211. Mora-O., L.E. & H. Sturm (eds). 1994. Estudios ecolgicos del pramo y del bosque altoandino cordillera Oriental de Colombia. Coleccin Jorge Alvarez Lleras. Acad. Colomb. Cienc. Exact. Bogot. Rangel-Ch., J.O. 2000. Clima De la Regin Paramuna en Colombia. Colombia Diversidad Bitica III:85-125. Rangel-Ch., J.O. (ed.) 2000. Colombia diversidad bitica III. La regin de vida paramuna. Instituto de Ciencias Naturales-Instituto Alexander von Humboldt. 902 pp. Bogot, Colombia. Sturm, H. 1998. The ecology of the paramo region in tropical high mountains. Verlag Franzbecker, 286pp. Hildesheim, Berlin. Sturm, H. & J.O. Rangel-Ch. 1985. Ecologa de los pramos andinos: Una visin preliminar integrada. Biblioteca J.J. Triana. No. 9: 292 pp. Instituto de Ciencias Naturales. Bogot. Troll, C. (ed.) 1968. Colloquium geographicum, Band 9: Geoecology of the mountainous regions of the Tropical Americas. UNESCO. Verweij, P.A. 1995. Spatial and temporal modeling of vegetation patterns: Burning and grazing in the pramo of Los Nevados National Park, Colombia. Ph.D. Thesis, University of Amsterdam: 223pp. Amsterdam. Vuillemier, F and M. Monasterio (eds.) 1986. High Altitide Tropical Biogeography. Oxford University Press. New York. (this is a classic on mountain ecosystems). 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