ࡱ> |~{@ 4bjbj "`uu,0000$(0-F<00000000EEEEEEE$iGRIlE*=00*=*=E00ExBxBxB*=00ExB*=ExBxBxB00 P#cG00AHxBBE0-FxB'JxB'JxB'JxB 04xB7L9D000EExB4Terrestrial habitat types 4.3. Pine/pine-oak/oak forest General description and geographic variation In the Neotropics, pine-oak forests are found in Mexico, Guatemala, Honduras and Northern Nicaragua, and some Caribbean islands. Pine trees never crossed the Nicaraguan depression during migration from Holarctic regions into the Neotropics. Oak forests, that is, forests without pines, are found from North America down to Central America and Northern South America. There have been reports of oak in Colombia, but not in Ecuador. Its southern most limit is the border of these two countries. Oak forests sometimes occur in lowlands, e.g., lowland Pacific dry oak (Quercus oleoides) forest in Costa Rica, but are generally more common in mountain regions. Some 12 species are known in Costa Rica (most important species are lower montane Q. seemannii and upper montane Q. copeyensis and Q. costaricensis), while only one species, Quercus humboldtii, is known in Colombia. In Mexico, the total number of oak species is about 10 times higher than in Costa Rica. Oak forests are a common and dominant element of the interior slopes of Colombia cordilleras between the elevations of 1000 and 3400 m. Quercus humboldtii immigrated from the north 250,000 years ago when they were widely distributed. Between the elevations of 1000 and 2000 m Q. humboldtii are associated with species of Nectandra and Ocotea (Lauraceae) and above 2000 m with species of Weinmania and Clusia. The stands in Colombia are usually established in the sites with annual precipitation between 1200 and 1500 mm and in very humid areas with annual precipitation exceeding 3000 mm. The structure and floristic composition of oak forests change across the Americas. Mexican oak forests are much drier, with a large number of holarctic species and shrubby or grassy understoreys, while Costa Rican oak forests are much taller, with trees over 35 m, and sometimes over 50 m. Costa Rican oak forests are generally mono- or di-specific at the canopy level and harbor bamboos of the genus Chusquea in the understorey. Costa Rican oak forests are extremely mixed in the sense that they are made up of both temperate and tropical elements-- holarctic oaks, alders, and rabbits occur together with tropical palms, melastomes, and tapirs. Wood volume (m3 per ha) and biomass (ton per ha) are highest in these Costa Rican oak forests and are among the highest values known. In Colombia, oak forests contain numerous typical Andean species that are local endemics. Neotropical and pantropical species are other important components in these forests. There is also an Austral-Antarctic group of genera, including Weinmannia, Drymis, Gunnera, Gaultheria, Escallonia, which are also known in Costa Rica, but in lower numbers. Community types/zonation and major gradients within the system (patterns) Oak forest communities form distinguishable zonation along elevation gradients. In Costa Rica, from lower elevations upward, oak starts to dominate at about 2000 m. Above 2000 m, in the upper parts of lower montane community, oak is found as a codominant genus, together with lauraceaous species of genera Ocotea and Nectandra. Above c. 2300 m, up to c. 3000 m, in the lower parts of upper montane community, oak dominates the canopy layer (35 to 45 m) and is associated with lower montane genera (e.g., Tovomitopsis, Hyeronima, Guarea, Sapium, Billia, Alfaroa and Phoebe) in the subcanopy layer (up to 20 m) and with dwarf palms (e.g., Chamaedorea and Geonoma) and gesneriads in the understorey. The oak forests are often dominated by species of Quercus, Podocarpus (a conifer) and Magnolia in the canopy, and Weinmannia, Vaccinium, Viburnum, Ocotea, Prunus, Styrax, Symplocos, Cornus, Ilex, Miconia, and the strangler Clusia in the subcanopy. Here, the Chusquea bamboo characterizes the understorey. Above c. 3000 m, up to a maximum of 3400 m, the upper montane and subalpine communities, oak (max. height 15 to 20 m) is accompanied by the ericad Comarostaphylis and genera such as Schefflera, Gaiadendron, Drymis, Weinmannia, Vaccinium, Brunellia, Buddleja, Escallonia and Miconia. Epiphytes abound in all montane oak forest communities. However, epiphytic orchids and bromeliads dominate between c. 2000 and c. 3000 m, while bryophytes and lichens are mainly found between c. 2500 and 3400 m. Ecological integrity factors for landscape context Table xxx. Ecological integrity factors for landscape context of pine/pine-oak/oak forest Key FactorsJustification for Factor SelectionMin. Integrity Threshold(s) Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityClimate Regime (vertical and horizontal precipitation, temperature.)Determines the occurrence and distribution of dominant flora and sets the boundaries for adjacent vegetation type.Less than 2,000 mm annual precipitation with significant dry season(s). [Experts, please provide a range of the length of dry seasons or literature citation. Thanks.]Along with fire and soil conditions, a primary determinant of geographic distribution. Litter accumulation Length of dry season Slope stability and landslide regimeSlope stability determines the frequency of landslides triggered by earthquakes and high rainfalls. Landslide regime in turn determines landslide disturbance patterns and creates landscape heterogeneity. Monitoring landslide factors: date of occurrence, causes, size of affected area, direction of landslide.Fire regimeAffects vegetation structure. Affects vegetation or faunal composition. With altered fire regime, Increase pines susceptibility to diseases and pests.Guess estimates: Surface fire frequency: maybe 20 years. Crown fire frequency: maybe 150-250 years. Needs to investigate abundance of oak versus pine Amount of litter Canopy structure Presence and size of seedlings and saplings Fire-scarred pines Sprouting following fire Soil charcoal, or presence of charred wood.Succession after disturbance (storms, hurricanes, landslides)Increases heterogeneity of vegetation structure and between-habitat (beta) diversity, important for patch dynamics.For example, one year after disturbance, check the growth of native plants (in meter) and the presence or absence of exotic plants. It is the time to check whether all the pioneer species are native.Hydrological regime and fluvial dynamicsAlong with the cover and structure of the vegetation, hydrological regime and fluvial dynamics determine the water yield and runoff rates. Fluvial dynamics associated with large rainfall events provides mineral seedbeds for pine regeneration (Caribbean). [Experts, if you know of any published studies, please cite. Thanks.]Water yield: annual streamflow totals = XX% of annual precipitation Run-off rate: XXIndicators of change in hydrologic regime such as water quantity, discharge, sediment load, and period of inundation.  Ecological integrity factors for condition Table xxx. Ecological integrity factors for condition of pine/pine-oak/oak forest Key FactorsJustification for Factor SelectionMin. Integrity Threshold(s) Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor Priority Viable populations of frugivorous and granivorous species. Regulate seed dispersal of pioneer, secondary and primary forest tree and shrub species. Acorn specific and pine seed specific in pine/oak forest.Minimum population sizes of frugivorous and granivorous species. [Experts, please provide an estimate of minimum population sizes of a few species as examples, or literature citation. Thanks.]Observational censuses at regular time intervals. Viable populations of mycorrhiza and fungal decomposers. Maintain decomposition and symbiotic relations with key tree species such as oak. Maintain the nutrient availability to species at ground level in closed forest.Minimum population sizes of mycorrhiza [Experts, if you know of any published studies (e.g., any papers by Jean Lodge?), please cite. Thanks.]Recording of mycorrhiza presence at regular time intervals. Presence of hojarascaVegetation structure(size, age class, strata)Provide diversity in micro-habitats and niches.Measure # of strata, abundance of life forms or guilds, types of functional groups and size/age classes. Ecological integrity factors for size Table xxx. Ecological integrity factors for size of pine/pine-oak/oak forest Key Factors Justification for Factor SelectionMin. Integrity Threshold(s) Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityMean and Maximum Fire Disturbance Area Fire is the principal disturbance regime and occurs with regularity.One example from DR of moderate intensity fire - Over 10,000 ha burned, crowned in 5-10% of the area (Horn et al 2000)Reference Fule in north-central Mexico. [Experts, please complete the citation in the References section. Thanks.]Presence of sensitive species [Experts, please give a few examples with species names. Thanks.] Species play various functions in pine / pine-oak/ oak [Experts, please specify what kinds of function. Thanks.}Need studies? (references?)Biotropica reference. Compendia on migratory songbirds and habitat needs. [Experts, please complete the citation in the References section. Thanks.]Presencia y abundancia de Loxia [Experts, please indicate the species name of Loxia. Thanks.]  Literature Cited Horn, S.P., K.H. Orvis, L.M. Kennedy and G.M. Clark. 2000. Prehistoric fires in the highlands of the Dominican Republic: evidence from charcoal in soils and sediments. Caribbean Journal of Science 36 (1-2):10-18. Jean Lodge Fule Biotropica articles Recommended resources Darrow, W.K. and T.A. Zanoni 1993. El pino de La Espaola (Pinus occidentalis Swartz): un pino subtropical poco conocido de potencial econmico. Moscosoa 7: 15-37. Horn, S.P., K.H. Orvis, L.M. Kennedy and G.M. Clark. 2000. Prehistoric fires in the highlands of the Dominican Republic: evidence from charcoal in soils and sediments. Caribbean Journal of Science 36 (1-2):10-18. Kappelle, M. 1996. Los Bosques de Roble (Quercus) de la Cordillera de Talamanca, Costa Rica: Biodiversidad, Ecologa, Conservacin y Desarrollo. Universidad de Amsterdam y Instituto Nacional de Biodiversidad (INBio). Amsterdam - Santo Domingo de Heredia, Costa Rica. 336 pp. Kappelle, M. and N. Zamora. 1995. Changes in woody species richness along an altitudinal gradient in Talamancan montane Quercus forest, Costa Rica. Pp. 135-148. In Churchill, S.P., H. Balslev, E. Forero and J.L. Luteyn, (eds). Biodiversity and Conservation of Neotropical Montane Forests. The New York Botanical Garden. Bronx, New York. 702 pp. Richardson, D.M. (ed.), 1998. Ecology ann biogeography of Pinus. Cambridge University Press, Cambridge, U.K. Rzedowski, J. 1994. Vegetacin de Mxico (sexta reimpresin]. Editorial Limusa, S.A. de C.V. Incomplete citations: Rzedowski 1978 Vegetacion de Mexico Limusa - Editor Biogeography of Pines Velazquez, Alejandro and Isolde Luna (Several papers in different journals) Breedlove, Dennis Barbour and Billings Gerald Islebe (several papers in several journals) Sally Horn et al Tim Fahey (not yet) in press =j M Y l y ~ ? 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