ࡱ>    o@ 2.bjbj p p #FooZDDDDX 4"X omt̨̨Z&&&HlJlJlJlJlJlJl$oR5rnlQ[ڳڳ[[nlDD&& l\[cD&D&Hl[Hl$ϋ)DLh & |e!IT4n amTomvIs!nIsX X DDDDIsr)Qf{|Ay3,ܶnlnlX X ĹdX X The Nature Conservancy Insert logo here.Measuring Ecological Integrity in the Neotropics A Resource Guide for Conservation Practitioners Draft, March 9, 2004 Preface Subtitle to be determined. To be determined. Acknowledgements To be determined. Thanks to the workshop participants, Shirley Keel, workshop organizers, expert reviewers, Jeff Parrish, Angela Martin, Maarten Kappelle. Executive Summary To be determined. Chapter1Introduction Ecological integrity in the neotropics Background To be determined. Purpose of resource guide To be determined. Methodology for developing resource guide To be determined. Chapter2Overview of ecological integrity What is it? Key ecological factors To be determined. Natural ranges of variation To be determined Monitoring indicators To be determined Ecological integrity and The Nature Conservancy Include: Ecoregional planning overview Conservation area planning overview Viability analysis within conservation area planning Chapter3Aggregations of ecological system targets To be determined. Chapter4Terrestrial habitat types 4.1. Lowland moist non-flooded broadleaf forest General description and geographic variation Lowland tropical moist forest is a closed, high forest, characterized by the large number of tree species occurring together. Gregarious dominants are uncommon. The forest in general does not have conspicuous stratification, but is conventionally regarded as having three tree layers, emergent trees, main stratum of 25-35m, and smaller shade-tolerant trees. Understory vegetation, especially herbaceous plants, is often sparse. Cylindrical bole, pinnate leaf, large leaf blade, buttress, liana, and cauliflory (flower borne on the trunk) are common. It occurs in a climate where water stress is absent with no regular annual dry season and an average of monthly rainfall >=100 mm, or where water stress is intermittent with short dry season of monthly rainfall <=60 mm or with particular soil conditions. This factor is coupled with high temperature (mean temperature >=180C of the coldest month of the year) and a strong evapotranspiration derived from the low-latitude and low-altitude rain forests and wetlands. Tropical moist forest was coined to cover both rain and seasonal forests (Whitmore, 1990). The Lowland Neotropical moist forest occurs in five regions (Prance, 1989), determined by biogeographic history: Mexico, Central America, and Pacific coasts of Colombia and Ecuador: The moist forest extends from the southern part of Veracruz (190N) in discontinuous patches to Panama and merges into the Choc rainforest, from northern Colombia to northern Ecuador. Choc rainforest is the wettest region in the world with annual precipitation >=9000 mm, and is an important center of rainforest endemism. In Central America, more moist forests exist along the wetter Caribbean sea coast. Transandean South America: along foothills of the Andes with elevation below 1,000 m. Amazonia, Orinoco and the Guianas host the largest area of continuous moist forest in the world, which includes part of the territory of Brazil, Bolivia, Peru, Ecuador, Colombia, Venezuela, Guyana, Suriname, and French Guiana. In most regions of northern South America the lower limit of annual precipitation3 for humid forests is approximately 1,700 mm. The Atlantic coastal forest of Brazil, a narrow belt 120-160 km wide stretching from southern Bahia to northern Rio Grande do Sul about 300S along the River Taquari, has a high degree of endemism, but the forest landscape is highly fragmented and only about 4% of original forest remain (Por, 1992). The Atlantic forest extends beyond the tropics. Moist forest in the subtropical zone is restricted to the wettest sites and the deepest, most fertile soils. In South America, subtropical moist forest, including Alto Parana forest, extends from southern Brazil and easternmost Paraguay to northeastern Argentina. Caribbean islands: The moist forest occurs mainly in areas where the presence of mountains increases rainfall, e.g., northern part of eastern Cuba, northeastern Jamaica, eastern Hispaola, northeastern Puerto Rico, and small patches in the Lesser Antilles. Community types/zonation and major gradients within the system (patterns) Major factors that determine variation in community types within lowland tropical moist forest: Precipitation: The amount of rainfall and length of dry season often determine the occurrences of evergreen forest or seasonally dry forest. Temperature: Yearly extreme temperature fluctuations result in cold-front stressed forests in SW Amazonia and the southern Atlantic region and non-cold-front stressed forests in Mexico and Central America. Topography: Zonation may occur depending on whether the forest is on a plain, or rolling hills, or foothills of a mountain range. Edaphic conditions (soil quality or fertility) can create special community types. Forests on white sand soil, on clay soil, or over limestone/ ultrabasic rock differ considerably in species composition. Natural disturbance includes earthquakes, hurricanes, landslides, extreme droughts, fire outbreaks and downbursts. Earthquakes or hurricanes are the most frequent causes of landslides. Earthquake prone areas cover Central America and eastern Andes, while the hurricane belt stretches from Mexico to Central America, and from the Caribbean islands to Yucatan. Fire outbreaks can occur in bamboo forests of western Amazonia during extreme drought years. Forest fires occur at an interval of approximately 600 years. Downbursts occur only in Amazonia. The frequency is high in western Amazonia, low in eastern Amazonia, and local in other areas. In summary, lowland tropical moist forests here include (Prance, 1989): Lowland evergreen rain forest Semi-evergreen rain forest (seasonally dry and with slight annual shortage of soil water) Forests over limestone/ ultrabasic rock, white-sand soil, or clay soil. Amazonian transitional forests (open evergreen forest, liana forest, and bamboo forest). Ecological integrity factors for landscape context Table xxx. Ecological integrity factors for landscape context of Lowland Moist Non-Flooded Broadleaf Forest Key FactorJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold) Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor priorityDisturbance regimes from catastrophic natural causes, e.g. hurricanes (hurricanes occur between 10-20 degree north and south of Equator), rare catastrophic floods, or multiple landslides, or volcanism, or earthquakes, rare extreme cold fronts, rare extreme droughts. These are extremely rare events that can be very important for ecological dynamics. Create canopy gaps of great size allowing pioneer species, e.g., Cercropia sp. to colonize and initiate successional processes, e.g., hurricanes play a major role in landscape-scale dynamics of forests on Caribbean islands.Stand turnover time (the number of years to replace a particular stand of forest) for heterogeneous primary forests: 200+ years. Catastrophic disturbance events in different scales: landslides >10,000 ha, river meanders >10,000 ha, tropical storms >1,000,000 ha, Cold fronts >1,000,000 ha. Successional processes: catastrophic landslides and river overflows lead to primary succession. Due to lack of topsoil and seed banks, the successional processes will therefore take longer than catastrophes derived from tropical storms, cold fronts, and droughts. Resilience (the rate to return to pre-disturbance condition): highly resilient to disturbance if physical structure (e.g. biomass) is maintained. An estimate of fresh biomass of a forest in central Amazonia: 730.7 metric tons/ha and 255 for below-ground root systems, for a total of 985.7 metric tons/ha (cited by Lovejoy in Four neotropical rainforests).The thresholds are intuitive extrapolations made by experienced field biologists who have occasionally observed large-scale events.Identify large areas at same successional stage. Monitor the presence of species that require very large-scale disturbance e.g. extensive stands of mahogany, native bamboo or vine-dominated forest stands and associated fauna (e.g., Goeldis monkey, bamboo rat, bamboo specialized birds, insects, & frogs.)[Experts, please give a few speices names of bamboo specialized birds. Thanks.]Fire due to dry spell or prolonged dry seasons or human activities.Certain species might be maintained because of this big, very rare catastrophic event. For example, mahogany thrives on fire outbreaks.Natural fires are rare in undisturbed tropical moist forest today (Ewel, 1983), but during years of exceptionally dry season, catastrophic fires may occur. El Nio in 1998 caused an exceptionally long dry season that increased the vulnerability of forests and the intensity and spread of fires in moist forests of southern Mexico. Large-scale, drought related fire return interval in Amazonia: 400-700 years (Nepstad et al., 1996).Presence of pure stands of balsa (Ochroma lagopus), whose seeds germinate in high temperature (Whitmore, 1990) Presence of fire-sustained grasses such as: Imperata brasiliensis, Hyparrhenis rufa, Panicum sp., and Saccharum sp. Presence of palm species that are resistant to fire, e.g., thousands of hectares of monospecific stands of Attalea speciosa (=Orbignya martiana) in once speciesrich seasonal forests of southeastern Amazonia (Ewel and Bigelow, 1996). Generations of mahogany of very distinctive size categories are found in large areas between south of the Rio Amazonas and west of the Rio Tocantins. Could this be a result of fire disturbance?Background disturbances- Small gaps, small landslides, downbursts, normal cold fronts, and normal seasonal precipitation variability. Important for creating and maintaining habitat heterogeneity and species and structural diversity, preventing competitive exclusion. Drives regeneration. Stand turnover times (the number of years to replace a particular stand of forest) for heterogeneous primary forests: <200 years. Disturbance events in different scales (Denslow, 1996); The value has been pushed up a lot to be almost catastrophic. What were previously catastrophic are now more background disturbances: landslides up to 100 ha, river meanders < 100 ha, tropical storms <1,000,000 ha, cold fronts = < 1,000,000 ha. Successional processes (Ewel, 1983): -within a year after disturbance, patches of weedy herbaceous plants and vines invade the area; -from one to a few years, vines and woody pioneer species (e.g. Cecropia sp.) dominate; -from a few to 25 years, dominating pioneer species form an even-aged, nearly closed canopy; -after 25 years, tree and animal diversity increase. Growth rates of trees are highest but their wood density is lower relative to later stages. The high net primary productivity of successional forests supports a large animal population, but not the same species as a mature forest. Known forest turnover rates provide a useful baseline to assess possible increments in the occurrence of background disturbances. Identify the presence of successional vegetation dominated by high light-demanding pioneer species (site specific). [Experts, please give a few species names at specific sites as examples. Thanks.] Satellite images can be used to quantify area, distribution, and frequency of background disturbances. (Note: theres concern that these background disturbances will be taking place at a higher than normal rate.) Spatial integration and coverage (e.g., connectivity by riparian habitats) allowing migration of animals and plants outside of lowland forest. Allow to define at landscape level integrity of ecosystem. Allow to assess the extent of potential for species extinction. Spatial integration important for species to maintain contact with all habitats required for life cycles. Thresholds of size and connectivity very difficult to say,. They are function of productivity and vary depending on type of forest and what species are there. Fragmented forests with sufficient connectivity to maintain ecologically functioning system: 100,000- 1,000,000 ha. (minimum dynamic area) Landscape mosaics necessary for individual species vary: -male jaguar in Pantanal needs 90 km2, -wetland peccaries spontaneous disappearance of groups. We dont know why, but caution dictates that areas of at least 10,000 km2 are necessary for long-term survival. -Corcovado in Costa Rica, large mammals have been seen in huge herds such as jaguars, etc. It seems to have dense mammal populations for fairly small area. Corcovado forests extending to the coast with exceptionally rich resource availability, can support sizable faunal populations. Many jaguars feed on sea turtles on the beach. Those inland have largely different food habits. Montes Azules in Lacandon with 360,000 ha of forest has good representation of harpe eagles, tapirs, peccaries, and jaguars. Studies in central Amazonia show that small fragments (< 100 ha) have lower plant biomass and carbon sequestration than continuous forest (Laurance et al. 1998).Monitor wide-ranging top predators and herbivores (e.g., Heliconia butterfly occurs only on large landscapes. Monitor plants that require rare disturbance and large intact landscape. [Experts, please give a few species names as examples. Thanks.] Remote sensing like satellite imagery with time series observations. Biogeochemical dynamics (referring to regional and global processes such as global warming, ozone depletion, CO2 concentration, atmospheric and soil pollution, etc.)Affects basic ecosystem functioning at both global and local levels. Thresholds: hard to come up with numbers. Small changes are likely to be significant because expected variation is not large.Small variation from baseline can be indicative of large changes in ecosystem functioning. (It is impossible to get baseline data that doesnt reflect human intervention for these large-scale processes).Need baseline studies at global scale of mortality rates of trees, long-term measurements of growth, frequency of disease outbreaks etc.Soil type or fertility. Affects forest primary productivity and species richness. It is one of the major factors determining forest types in Amazonia. Soil type is also relevant to tree mortality rate, treefall frequency, forest regeneration mode, and stand turnover time (Hartshorn, 1990).Treefalls appear to be more frequent in the forests on clay soils (oxisols & ultisols) than on white sand soils (podzols). Stand turnover time is shorter and tree mortality rates are higher in forests on fertile soils which usually support structurally more complex and floristically more diverse forests (Hartshorn, 1990). Nutrient-poor soils with low primary productivity such as white sand soils tend to have a lower species richness of carnivorous vertebrates, insects, and scavengers (Janzen, 1983).Requires much active management. Ecological integrity factors for condition Table xxx. Ecological integrity factors for condition of Lowland Moist Non-Flooded Broadleaf Forest Key FactorJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold) Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor priorityDiversity of above-ground plant functional groups (species that share morphological, chemical, structural or life history characteristics) Determines the role of biodiversity in ecosystem functioning such as nutrient cycling, forest regeneration and successional patterns. Threshold: significant change in diversity of functional groups; significant change is defined relative to site- specific baseline. Life forms: life histories arranged in order of possible range of importance- criticality for to biodiversity and ecosystem functioning and dynamics. Canopy and emergent trees 4,7 Climax tree species or shade-tolerant species16 Pioneer tree species or light-demander or shade-intolerant species16 Lianas and vines Epiphytes and hemiepiphytes4,7 Understory treelets or shrubs 4,7 Some life forms may be more susceptible to natural and human-induced perturbations (e.g., Palm vulnerability to cyclones in Puerto Rico [SK comments: Species of Roystonea show adaptation to Caribbean habitat by losing their leaves early in hurricanes and thus capable of withstanding strong winds without being blown down.-Henderson et al., 1995) ,vines to fire in selectively-logged forest in Amazonia, emergent trees more susceptible to high winds). This may affect other organisms (e.g. animals) and ecological processes (e.g. biotic interactions, gap regeneration). (Dirzo, 2001, in press, Orians et al. 1996) Longitudinal transects can easily be conducted to assess presence and relative abundance of keystone and representative species of each functional group. ( RRepresentative species of functional groups and life forms will be site specific. See Denslows paper for review and examples. Experts, if his paper is not the one cited in the References section Denslow, 1996, please provide the literature citation. Thanks.) Monitor presence and abundance of life forms in seed banks. Soil samples can be obtained and sieved to check seed diversity. For example, small seeds usually correspond to pioneer species and large seeds typically correspond to mature forest species. Diversity of animal functional groups Such diversity determines a number of key ecological processes such as trophic structure, nutrient cycling, systems resilience to disturbance.Threshold: significant change in diversity of functional groups; significant change is defined relative to site- specific baseline.Suggests the level of unexpected difference; in some groups major disruptions of trophic change occur (if take out top predators in some cases). Monitor the abundance of Keystone species of each functional group such as top predators and important pollinators (see pollination indicators below). Interview local communities to monitor the status of the functional groups.Community composition/ Diversity /structureAffects species diversity and several ecosystem-level processes. Several site-specific studies have calculated the relative importance value (IV) of individual species. [Experts, please provide site names and literature citation. Thanks.] Some evidence suggests that a set of species with the greatest IV may define a threshold for several aspects of ecosystem functioning (Orians et al. 1996).Species composition and diversity is the most distinguishable attribute of tropical lowland forests. Direct and indirect value of species for humans, particularly non-timber forest products (NTFP). Diversity of species and functional groups may be the most critical component of tropical forest resilience against environmental change. Presence of mosaic microhabitats in which different species co-occur, ranging from pioneer species (Cecropia spp., Balsa trees, some Ficus spp.figs) to mature forest species (Dipteryx spp, Brosimum spp., Oenocarpus bataua). Censuses on standing and fallen dead trees. These provide important resources and habitat for animals. Standing dead Palms are important nesting sites for macaws. Fallen dead trees provide habitat for beetle larvae, fungi, and many other decomposers. Censuses on diversity and abundance of canopy mixed speciesp flocks of birds. Monitor the presence and abundance of alien invasive speciesp away from human settlements. In order to monitor, develop a list of locally present exotic speciesp.Gap dynamicsProvides light, the major environmental limiting factor to plant growth in the close-canopy tropical forest, and maintains the forest in shifting mosaic steady state.16  Rate of recovery6: more rapid, if mycorrhizal fungus populations and seed sources of mature forest species are present. Annual tree mortality9: 1-2% (3% tree mortality on Barro Colorado Island in the 1983 dry season associated with El Nio.) Median tree (>10 cm dbh) longevity9: 30-50 years. Turnover Turnover rate 60-100 years in Mexico and Central America forests. Average gap size9: about 100 m2, ranging from a few m2 (e.g., branchfalls) to a few hectares. Blowdowns by downburst winds in Amazonia can be larger than 1000 ha.  Size, distribution, and timing of gaps determines the spectrum of life forms and speciesp diversity in these gapsecosystems. (site specific).- Establish line transects to check time distribution and size distribution of gaps on a periodic basis. Larger Gaps can be identified using remotely sensed images. Time series images can be used to detect changes in rates. Biotic interactions: Pollination (bees, butterflies, beetles, moths, bats, and hummingbirds) Important for reproductive success: pollinators influence the frequency and distribution pattern of plant species. Difficult to define the threshold. Necessary to study population dynamics of pollinators. Variation in species diversity of these functional groups: need to have plenty of diversity in all these groups. Is the diversity of pollinators proportional to the diversity of plant species? Is there a minimum number of pollinators? Necessary to study the complex relationships between pollinators and flowers they pollinate. Pollinators: bees are the most important group of pollinators. Bats, birds, large moths and bees can move pollen over distances of 0.1 to 1 km or more (Janzen, 1983). You need the baseline as a reference. Indicator of fruit set: Decline in fruit production (palms, figs, Brazil nuts, legumes, orchids). Fruit set in Bombacaceae is indicative of pollination by bats. Consultations with locals on fruit set. Census on bees with fragrances. Biotic interactions: seed dispersal executed by fruit-eating birds (e.g. toucans and cotingas), mammals (e.g., tapirs, peccaries, monkeys, bats, etc.) and ants.Important for reproductive success: seed dispersal agents affect food webs in tropical forests by making available reproductive resources to other consumers and Influence the frequency and distribution pattern of plant species, especially woody species. The composite seed shadow generated by the entire forest is the base of the food chain for a large number of animals (Janzen,1983).More than 60% of woody species in wet tropical forests are adapted for seed dispersal by vertebrates (Rejmnek, 1996). Difficult to define the threshold: Start monitoring on the basis of reliable knowledge. Compare with baseline to define thresholds. Check whether thresholds differ significantly from baseline. Necessary to study population dynamics of these animals. Variation in species diversity of these functional groups: need to have plenty of diversity in all these groups. Animal-generated seed shadows tend to be skewed at resting places, sleeping places or other high concentrations of animals. You need the baseline as a reference. Indicator: Censuses on charismatic frugivores (toucans, monkeys, tapirs, cotingas, kinkajous) Rate of removal of indicative fruits such as Piper spp. (dispersal by bats) Consultations with locals on abundance of frugivores. Censuses on recruitment of animal-dispersed species. Biotic interactions: seed predation (parrots, cracids, beetles, agoutis, etc). Important for reproductive success: seed predation affects population recruitment and establishment of diverse plant species (e.g. palms and legumes). Seed predators occasionally act as dispersers. Seed predation is a specialized form of herbivory. Vertebrates involved are often objects of hunting by humans.Necessary to study population dynamics of these animals. Variation in species diversity and abundance of this functional group is site specific. Threshold is unexpected change in species diversity relative to site-specific variation. Seed dispersal agents can also be seed predators. Examples include: fig wasps (Agaontidae), acting as fig florets pollinators and fig seed predators; Bairds tapir (Tapirus bairdii) a pure disperser of fig seeds, but a pure predator of Sideroxylon capiri (Sapotaceae) seeds; agouti (Dasyprocta), an important disperser and predator of Attalea palm seeds; peccaries (Tayassu) and forest-floor rodents acting as major predators of dispersed tree seeds (Janzen, 1983). You need the baseline as a reference. Consultations with locals or censuses on abundance of vertebrate seed predators (parrots & macaws, curassows, monkeys, agoutis, pacas, peccaries). Consultations with locals or censuses on abundance of insect seed predators (e.g. Bruchid Beetles are collected for food by many local populations).Biotic interactions: herbivores including insects (e.g. caterpillars, leaf-cutter ants), parasitic fungi, and vertebrates (e.g., peccaries, tapirs, deer, some monkeys).Herbivory affects vigor and mortality of plants of all sizes, especially understory seedlings, and Influences food chain and species composition of understory. Threshold: Observations that go beyond the known baseline levels of herbivory should be a warning to look at alterations in the components involved. The 10% can be considered baseline against which to assess possible outbreaks. Folivorous insects are much more important than vertebrate herbivores, and their species diversity may be dependent on plant species richness due to host-specificity. Studies (de la Cruz and Dirzo, 1987) in several forests indicate that leaf area consumed by herbivores is roughly 10%. There is considerable variation in time and among species and life forms. Studies by Coley (1982) show that insect herbivory on mature leaves of saplings of 21 canopy tree species in Barro Colorado Island is at an average annual rate of 21%. Tropical forest trees often occur at low densities or in patchy distributions. Pathogens or defoliating insects have localized rather than standwide effects. Dominant plant species however are vulnerable to standwide defoliation by specialized herbivores and pathogens. You need the baseline as a reference.Surveys of standing leaf area eaten. Elevated populations of herbivorous insects, e.g. in light traps, frass, nests, or insect activity in the forest. In the case of vertebrates: transects of sightings, quantification of animal tracks, and interviews with local communities. Biotic interactions: presence of top predators.Controls the populations of small mammals and herbivores. Army ants form an important ecological system as predators of insects and species complex that follow them (e.g., parasitic flies, birds etc.) Army ants are also critical for maintenance of vegetation structure. [Experts, please provide literature citations. Thanks.]Minimum viable size of population: need to determine population growth rate. If its >= 1 (lambda) Jaguar Population density in fairly undisturbed areas: 1/15 km2 ~ 1/64 km2 (home range of a male jaguar: 28 -168 km2) (Emmons, 1991).Preliminary data by Terborgh suggest that maintenance of top predators natural range of variation may be critical for forest structure and diversity.Absence of top predator species or population increase of small mammals. Transects of sightings, quantification of animal tracks, and interviews with local communities. To assess the condition of army ant populations and conduct censuses to determine whether an area has a full complex of ant-following birds. Sample methodologies described in Dirzo and Miranda (1990). Species diversity and composition of soil biota, e.g., mycorrhizae, fungi, microbes, soil mesofauna such as leaf-cutter ants, termites, nematodes, collembola, dung beetles, etc. Fundamental for nutrient cycling and soil structure. Threshold: Statistically significant change in diversity and abundance of soil biota. The threshold marks significant changes in ecosystem processes related to energy flow and nutrient cycling Monitoring litters decomposition and dung decomposition, censuses of termite and ant nests and their viability. Mycorrhizae would be an important indicator but requires technical expertise and equipment. Practitioners can do this after some basic training. Species diversity and composition of understory vegetation. Determine the mechanism of forest regeneration and indicate the health of successional change.Threshold will be defined by baseline and measured variation. Indicators of normal recruitment: no understory vegetation, absence of expected number of understory species, significant reduction of number of individuals in understory (pay special attention to exploited species.) Allows to detect critical situations of forest regeneration patterns. Presence of Invasive alien species such as the citrus in Chuquisaca, Bolivia. (Invasive alien faunal species such as muscoid and fruit flies in Amazonia, European honeybees (particularly africanized), and teju lizards. [Experts, please indicate scientific names of mentioned invasive alien faunal species. Thanks.] Overabundance of particular dominant species such as Brosimum alicastrum trees in Mexico and Central American forests. In the case of direct-use conservation units, a significant decline of exploited plants. Establish permanent plots to track changes in species composition that differ from baseline situations. Low Ecological integrity factors for size Table xxx. Ecological integrity factors for size of Lowland Moist Non-Flooded Broadleaf Forest 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 PriorityExample: Mean and Maximum Fire Disturbance Area Fire is the principal disturbance regime and occurs with regularity.20,000 ha = MDA Good = 2x MDA Very Good = 3x MDA20,000 ha is the maximum recorded fire disturbance for this system; most fires affect areas smaller by a factor of 10. Considering our uncertainty about future interactions of fire and invasive species in grasslands, it is believed a buffer of 3X the minimum dynamic area is ideal (Desired Future Condition)Aerial photography at 3-5 year intervalsHigh Literature Cited Coley, P.D. 1982. Rates of herbivory on different tropical trees. In Leigh, E.G. Jr., A. S. Rand, and D.M. Windsor (eds.). The ecology of a tropical forest: seasonal rhythms and long-term changes. 2nd edition. Smithsonian Institution. 123-132. De la Cruz, M. and R. Dirzo. 1987. A survey of the standing levels of herbivory in seedlings from a Mexican rain forest. Biotropica 19: 98-106. Dirso, R. and A. Miranda. 1990. Contemporary Neotropical defaunation and forest structure, function, and diversity- a sequel to John Terborgh. Conservation Biology 4 (4): 444 447. Dirso, R., 2001. Tropical forests: biodiversity, ecological processes, and global environmental change. In Chapin, F.S. and O. E. Sala (eds.) Future biodiversity scenarios. In press. Denslow, J.S. 1996. Functional group diversity and responses to disturbance. In Orians, G.H., R. Dirzo, and J.H. Cushman (eds.) Biodiversity and ecosystem processes in tropical forests. Springer-Verlag, Berlin, Heidelberg, New York. Pp. 127-151. Emmons, L.H. 1991. Jaguars. In J. Seidensticker & S. Lumpkin (ed.) Great cats. Fog City Press, San Francisco. Ewel, J. 1983. Succession. In Golley, F.B. (ed.), Ecosystems of the World 14A, Tropical rain forest ecosystems: structure and function. Elsevier Scientific Publication Company, New York. Pp. 217-223. Ewel, J.J. and S.W. Bigelow. 1996. Plant life-forms and tropical ecosystem functioning. In Orians, G.H., R. Dirzo, and J.H. Cushman (eds.) Biodiversity and ecosystem processes in tropical forests. Springer-Verlag, Berlin, Heidelberg, New York. Pp. 101-126. Hartshorn, G.S. 1990. An overview of Neotropical forest dynamics. In Gentry, A.H. (ed.), Four Neotropical rainforests. Yale University Press, New Haven. Pp. 585-599. Henderson, A., G. Galeano, and R. Bernal. 1995. Field guide to the palms of the Americas. Princeton University Press, Princeton, New Jersey. Janzen, D.H. 1983. Food webs: who eats what, why, how, and with what effects in a tropical forest? In Golley, F.B. (ed.), Ecosystems of the World 14A, Tropical rain forest ecosystems: structure and function. Elsevier Scientific Publication Company, New York. Pp. 167-182. Laurance, W. F. & R. O. Bierregaard. 1997. Tropical forest remnants: ecology, management and conservation of fragmented communities. University of Chicago Press. Nepstad, D.C., P.R. Moutinho, C. Uhl, I.C. Vieira, and J.M.C. da Silva. 1996. The ecological importance of forest remnants in an eastern Amazonian frontier landscape. In Schelhas, J. and R. Greenberg (eds.) Forest patches in tropical landscape. Island Press, Washington, D.C. Pp.133-150. Por, F.D. 1992. Sooretame, the Atlantic rainforest of Brazil. The Hague: SPB Academic Publishing. Prance, G.T. 1989. American tropical forests. In Lieth, H. and M.J.A. Werger (eds). Ecosystems of the World 14B. Tropical rain forest ecosystems: biogeographical and ecological studies. Elsevier Scientific Publication Co., New York. Pp. 99-132 Rejmnek, M. 1996. Species richness and resistance to invasions. In Orians, G. H., R. Dirzo, and J.H. Cushman (eds.) Biodiversity and ecosystem processes in tropical forests. Springer-Verlag, Berlin, Heidelberg, New York. Pp. 153-172. Whitmore, T.C. 1990. An introduction to tropical rain forests. Oxford York University Press, New York. Additional resources Challenger, A. 1998. Utilizacin y conservacin: de los ecosistemas terrestres de Mxico, pasado, presente y futuro. CONABIO. Condit, R. 1997. Forest turnover, diversity, and CO2. Trends in Ecology and Evolution 12: 249-250. Gentry, A.H. (ed.), 1990. Four Neotropical rainforests. Yale University Press, New Haven. Hunter, M.L.(ed.), 1999. Maintaining Biodiversity in Forest Ecosystems. Cambridge University Press. Laurance, W. F. and R. O. Bierregaard. 1997. Tropical forest remnants: ecology, management and conservation of fragmented communities. University of Chicago Press. Malhi, Y. & Grace, J. 2000. Tropical forests and atmospheric carbon dioxide. Trends in Ecology and Evolution 15: 332-337. McDade, L.A. et al. (eds.), 1994. La Selva. University of Chicago Press, Chicago. pp 486. Orians, G.H., R. Dirzo, and J.H. Cushman (eds.), 1996. Biodiversity and ecosystem processes in tropical forests. Springer-Verlag, Berlin, Heidelberg, New York. Phillips, O. 1997. The Changing Ecology of a Tropical Forest. Biodiversity and Conservation 6: 291-311. Voss, R. S. & L. H. Emmons. 1996. Mammal diversity in Neotropical lowland rainforest: a preliminary assessment. Bull. Amer. Mus. Nat. Hist. 230: 1-115. Witmore, T.C. and J. A. Sayer (eds.), 1992. Tropical deforestation and species extinction. Chapman & Hall, London. 4.2. Lowland moist flooded broadleaf forest General description and geographic variation Flooded forest is "any wetland with a significant component of woody vegetation that is inundated or saturated by surface- or groundwater, freshwater, or tidal water at such a frequency and duration that under natural conditions it supports organisms adapted to poorly aerated or saturated soil." Structural characteristics that recur in flooded forests are presence of monospecific stands such as palm swamp, even-canopied forests, and sharp vegetation zonations. It is common to find that trees in flooded forests develop sclerophylly (firm, thickened leaf) due to poor nutrition or water limitations, or gas exchange structures such as pneumatophores, lenticels, knees, aerial roots, swelling of base of trees, surface or aerial roots in order to overcome poor soil aeration, or support structures, e.g., plank buttresses and stilt roots to provide stability in muddy or steep conditions. Tree heights can vary greatly, between 1-50 m in mangrove stands, depending on the severity of environmental stress (Lugo, 1990). Flooded forest systems are a regular feature of the low-lying coastal areas in the West Indies, southern Mexico, Central America and northern South America. In Mexico, most forested wetlands are concentrated in the coastal plain of southern Veracruz, Tabasco, and Campeche. In Central America, most of the lowlands and the major river systems are found on the Atlantic slopes (De la Rosa, 1995). Along Atlantic and Pacific coasts, swamps of fresh or brackish water with Rhizophora, Conocarpus, Pelliciera, Prioria and the palms Manicaria and Raphia are found in Guatemala, Belize, Honduras, El Salvador, Nicaragua, Costa Rica, and Panama. Inland swamp forests are found in the NW of Progresso and SW of Lago de Izabal in Guatemala, and along the eastern shore of Lake Nicaragua and south of El Castillo in Nicaragua (Davis et al., 1999). In South America, the greatest extent of flooded forest occurs in Amazonia, covering at least 150,000 square kilometers, about 3% of the total Amazon rainforest (Goulding, 1993). Other flooded forests are found in the Orinoco basin, the alluvial overflow plains in central depressions of the Llanos of Venezuela and Colombia, the Choc biogeographic region of Colombia including the delta of the Rio Atrato to the north and the delta of the Ro Pata to the south, the wetlands in SW of Lago Maracaibo in Venezuela, the Llanos de Moxos of Bolivia, and the Pantanal. These areas are all characterized by complex mosaics of wetlands, savannas, and flooded forests. In the Caribbean woody swamp vegetation is reported from the Greater and Lesser Antilles. Small stands of swamp forests occur in the Lesser Antilles (Bacon, 1990). The total rainfall and its distribution patterns are the principal climatic factor that determines the geographic variation of the size of flooded forest ecosystem. In Central America, streams are short with small catchment basins and low total discharge. The western part of Central America receives much less precipitation than the east, the riparian forests are in general fairly small. In northern South America, eastern Central America and the upper Amazon, the amount of rainfall allows medium size flooded forest ecosystems. In the lower Amazon, with rivers originating in the Andes, the aseasonal high rainfall is the cause of the largest and most extensive flooded forest system. From Cerrado to the Gran Chaco, with diminishing precipitation in the interior of South America, the flood forests are relatively small ecosystems. Community types/zonation and major gradients within the system (patterns) Flooded forests can be grouped into riverine, fringe and basin types. Riverine flooded forest depends on seasonal river floods such as vrzea or gapo. Fringe flooded forest grows on oceanic and lake shorelines where water flows are bidirectional, e.g. tidally flooded forest, or freshwater tidal swamp forest. Basin flooded forest is found in depressions where water accumulates and may fluctuate in depth depending on the balance of rainfall runoff and evapotranspiration, e.g. aguajal (Lugo, 1990). Based on the length of the hydroperiod, flooded forests can be grouped into permanently inundated swamp forest and periodically inundated marsh forest. Swamp forest is usually found on soils that have high water table, e.g., Mauritia flexuosa (palm) swamp in Trinidad grows on land perpetually inundated with 30 to 100 cm of water; while marsh forest occurs in areas subjected to inundation during rainy season. Species richness generally decreases with increasing hydroperiod. Based on the type of dominant species, swamp forests can be conveniently divided into two types: forests dominated by hardwood species and those dominated by palms. Dominance by palms becomes stronger with increasing hydroperiod or soil moisture conditions (Bacon, 1990; Lugo et al., 1990). Ecological integrity factors for landscape context Table xxx. Ecological integrity factors for landscape context of Lowland Moist Flooded Broadleaf Forest Key FactorJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold) Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityFlood regime: variation of water level due to hydroperiodHydroperiod- the duration, magnitude, frequency or season of flooding- regulates the succession of ecological processes and physical, chemical and biotic changes in both river, floodplain, and estuary environments. Species diversity is strongly correlated with gradients of flooding depth and duration (Goulding, 1993; Lewis et al., 2000; Lugo et al., 1990).Study patterns of tree rings (e.g., Macrolobium acaciifolium, Genipa spruceana) to help reconstruct historic range of flooding intervals in the past 100-150 years. Duration of flooding: A few days to permanent inundation, depending on the area, e.g., in Amazon floodplain: 6-7 months (Goulding, 1993). Flood (normal or catastrophic) return interval: x year(s), e.g., the Cienaga Grande in Colombia has extreme cyclic flooding of 6-7 years (Bacon, 1990). Depth of water level: 7-13 m /year in Amazon basin (Goulding, 1993).Population decline of keystone (faunal sp.) or dominant tree species. Monitor species dependent on rare events (habitat complexity measure).[Experts, would you please give a few examples of species and methods of measuring habitat complexity or habitat use efficiency or references on this subject. Thanks.) Water level gauge for measuring annual pulse.[Pulse triggers the reproduction of catfish and characid, caiman and turtles.) Remote sensing to measure nutrient flux in large river systems. Consultation with local communities on pattern and frequency of hydroperiod.HighTopographyTopography- position on the landscape e.g. in depressions or on ridges, and elevation- determines extent of tidal penetration, and local flood regimes ( water level, water retention time, and extension of flooding), variation in river flow, erosion and sediment deposits, habitats and vegetation zones (Daly and Mitchell, 2000; Godoy et al., 1999; Lewis et al.,2000; Lugo et al., 1990).Altitudinal intervals (in meters) that define different vegetation zones.See topographic mapHighSlope Affects fluvial dynamics.Channel dynamics (= fluvial dynamics?)The rate of channel change or lateral migration of meanders is important for creating habitat mosaics such as oxbow lakes, levees, seasonal lakes, canals, forested terraces, diverse biotic communities and successional patterns (Henderson and Robertson, 1999).Frequency of natural disturbance, e.g., tectonic activities of the Andes (Linna, 1993). Extent of natural disturbance, e.g., accreation of 100 m/yr of sea level on the Surinam coast (Bacon, 1990).Indicator species of early succession on recent sediments (allogenic successional species) in the upper Amazonia: Gynerium sagittatum (Daly and Mitchell, 2000) and Salix humboltiana (Salix needs new sediment to establish roots). Indicator species of autogenic succession: all species that cannot grow on sediments. [Experts, please give a few species names as examples. Thanks.] HighGeomorphology/ soil type Determine river types, e.g., black water river, white water river, and clear water river, nutrient contents, pH, sediment loads, and species composition and diversity of biotic communities (Daly and Mitchell, 2000; Goulding, 1993; Terborgh and Andresen, 1998). (The hard iron oxide and sandstone in Amazon floodplain may be the reason that rivers in floodplain do not meander freely.) Range of pH in white water rivers: >=7 (basic) See geology, hydrology and soil mapsMedium Upland, upstream & downstream vegetation continuityImportant for providing refugees for terrestrial animals and maintaining population size of organisms that rely on water as dispersal agent for their propagules during flooding season.Distance/area of the flooded water extending into the adjacent upland forest on river bank: ca. 20 km2 in the Amazon basin.See vegetation mapMediumLateral connectivity within wetland associated system. Affects interaction between aquatic and terrestrial systems. High lateral connectivity: permanent exchange of water between river and floodplain, but little change of species composition. Medium connectivity: connected once a year during peak floods, with limited exchange of nutrients and biota. Low connectivity: connected during extreme floods, only once in 20-30 years. Extreme events can change biota completely.Fire regimeFire-flood stress during high water and fire stress during low water can occur. (e.g., in Pantanal)Need studies of fire scars and tree ring structure to reconstruct fire history and natural fire intervals to establish site-specific baseline for fire management.Hydrological connectivityAffects fish movement during inundation along river channels to colonize additional floodplains.Watershed shape Ecological integrity factors for condition Table xxx. Ecological integrity factors for condition of Lowland Moist Flooded Broadleaf Forest Key FactorJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold)Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityAge of floodplain forestsIndicates the successional stage of floodplain forest and its ecological stability.Floodplain forest reaches maturity in >= 400 years.Mature forests contain tree species of 400 years old (evidence from tree rings).Species composition of floodplain forestA good indicator of the health of the forest. Affects forest canopy litter production. Maintain mutualism between fruit-eating fishes and fruiting trees. Tree species diversity varys with flood gradient. Higher species diversity has been observed in high levees. In Manaus, about 200 tree species; while in Tefe area, about 400 tree species. (Experts, please indicate the unit of the area (x ha. or xxxm2) of measured tree species, the size class of tree species (DBH), and literature citation. Thanks.)Indicators of forests that have been selectively cut: presence of abundant fig trees. [Experts, any particular species of fig trees? ] or hollow-centered hardwood tree species.Species composition of fish community: fruit-eating fishes.Important for maintaining diversity of tree species that depends on fishes as dispersal agents. About 20% of fish species of the Amazon floodplain are fruit-eaters. (Experts, please provide literature citation. Thanks.] Spectrum of age/size classes: presence of all age or size classes with high proportion of small size individuals. Consultation with fishermen on fish populations and changes in harvested amounts. Monitor the population of tambaki (Colossoma macropomum), an obligatory fruiteating species that will leave the area where is deficient in fruit trees. Change in size classes of fish is often an indicator of change in available food and trophic structure. Species composition of fish community: migratory fishes.Sensitive to large-scale connectivityThreshold: maintain connectivity up to 2000 km for some catfishes [Experts, please indicate scientific names of some catfishes. Thanks.)Consultation with fishermen on fish populations and changes in harvested amounts. Monitor the population of catfishes (900,000 tons /yr. fishes harvested in the entire Amazon basin). [Experts, please provide literature citation for 900,000 tons/yr. Thanks.] Commercial fishing of catfishes? Paiche? has reduced from 50,000 tons to 5,000 tons/yr. [Experts, please provide literature citation for the figures. Thanks.)Water chemistryThe water quality determined by pH, salinity, dissolved and suspended sediment, and nutrient quantities of nitrogen, carbon and phosphate, affects species composition and biodiversity of biotic communities of flooded forests (Daly and Mitchell, 2000; Goulding, 1993).  Rich, medium, low Dominant swamp species, e.g., mangle or palm species or hardwood species.HighWater transparencyControls the optical environments important for phytoplankton growth, nutrient cycling, species abundance and composition of fish communities by nonvisual predators or visual predators (Goulding, 1993; Lewis et al.,2000)Presence of phytoplaktonHighWater current flow rateAffects species composition and diversity of fish communitiesHighWater sourceDetermines the energy level of aquatic systemRainfall: low energy Upland river: high energy.Biotic omponents: host species, resident biota, migrants, and canopy fauna. (Experts, please give a few examples of host species, resident biota, migrants and canopy fauna of the sites you know. Thanks).Important to the function of wetlands. Host species are wetland-dependent. Without wetlands, there will not be any host species. They are the most important part of wetland biodiversity. Migrants are occasional visitors and canopy fauna specializes on specific trees but independent of wetlands. They all contribute to the array of species diversity of the flooded forests. Experts, please provide some examples of thresholds from sites you know or from literature. Thanks.Presence of viable populations of herbivores, e.g., capybaras, manatees, and turtles. Feed on aquatic macrophytes that support invertebrate community, production and succession. Herbivores are Important for trophic structure and nutrient cycling.Census of populations of manatees, primates, and invertebrates.MediumPresence of viable population o, top predators (e.g., otters, dolphins, caimans, jaguars, or osprey).Important for trophic structure and nutrient cycling. Sensitive to biocumulative toxins.Estimated (an educated guess) minimum number of breeding pairs in a local population for maintaining genetic pool is 500 pairs. [Experts, what species are you referring?] Jaguars often take preys in wet and waterside habitats. Studies indicate that they spent about 70% of their time in forest located at a mean distance of 0.5 km from water (Emmons, 1991). Estimated jaguar density in relatively undisturbed forests: 1/15 km2 in Belize, 1/64 km2 in the Brazilian Pantanal. Populations of black caiman have declined significantly due to hunting. The density of black caiman in 6 localities of Amazonian Ecuador range from 5 to 280 per km2. (Experts, please provide literature citation. Thanks.).Census of populations of birds, boto dolphin (Inia geoffrensis), caiman nests. Consultation with local fishermen and hunters.MediumPresence of exotic species, e.g., water buffaloes.Affects reparian vegetation. Water buffaloes feed on shoots, and therefore have detrimental effects on vegetation.  Ecological integrity factors for size Table xxx. Ecological integrity factors for size of Lowland Moist Flooded Broadleaf Forest 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 PriorityExample: Mean and Maximum Fire Disturbance Area Fire is the principal disturbance regime and occurs with regularity.20,000 ha = MDA Good = 2x MDA Very Good = 3x MDA20,000 ha is the maximum recorded fire disturbance for this system; most fires affect areas smaller by a factor of 10. Considering our uncertainty about future interactions of fire and invasive species in grasslands, it is believed a buffer of 3X the minimum dynamic area is ideal (Desired Future Condition)Areal photography at 3-5 year intervalsHighMinimum size of floodplain forest Forests serve as current breaks or fire breaks. Floodplain forest is an important habitat to canopy fauna or invertebrates adapted to canopy. Shading by forests affects phytoplankton production. Refuge for animals during low water. Sufficient forest cover to help reduce energy in the river system during flood, and maintain ecological processes of biotic exchanges.Experts, please provide descriptions or some examples of thresholds from the site you know well or from literature. Thanks.High Additional information Table xxx. Known occurrences or range distribution of large community types of the Flooded Forest System in LAC Type of flooded forests in LACCommon nameLocationAcoelorraphe wrightii Tique palm swampTikal (Honduras)Patchily distributed in the Caribbean (southern Florida and Cuba), and on the Atlantic coast of Mexico (the Yucatn Peninsula, Tabasco, and southeast Veracruz), Beliz, Guatemala, Honduras (Rio Platano), Nicaragua (South of El Castillo), and Costa Rica.Annona glabra associationWidely distributed, from central Veracruz to western Tabasco, and in isolated patches in Campeche and southern Quintana Roo.Attalea butyracea/ A. cohune palm swampWidespread in Mexico (Oxaca, Chiapas, Veracruz, Tabasco, Campeche, the Yucatn Peninsula. EO: Bajo de Papaloapan , along the Ro Usumacinta and its tributaries in N and NE Chiapas and adjacent areas of Tabasco), Central America (Atlantic and Pacific coasts of Guatemala, Pacific coast of Nicaragua and Costa Rica (dry forests of Guanacaste and Puntarenas), and northern South America in Trinidad, Tobago, Colombia (valleys of Ro Cauca and Magdalena, Caribbean lowlands, eastern plains and Amazon region), the Llanos of Venezuela.Bactris swampPantanal, Honduras, Colombia (Magdalena river in Caribbean region.)Bravaisia integerrima forestCanacoital (Mexico)Southern Tabasco and northern Chiapas, MexicoBucida buceras associationPuktal (Mexico)Isolated patches between the Mamantel and Candelaria rivers in Campeche, in southern Champotn of Campeche, and around the Ascensin and Espritu Santo Bays in Quintana RooCampnosperma panamensis forestSajal (Colombia)Colombia (Nario, Valle del Cauca, Choc)Copernicia alba palm swampThe Chaco region of Paraguay and adjacent countries: Brazil (Mato Grosso, Mato Grosso do Sul), Bolivia (Beni, Santa Cruz), and Argentina (Chaco, Formosa, Santa F)Carapa guianensis dominated forestTangarial (Colombia)Colombia (Choc Biogeographic region) Elaeia oleifera palm swampSmall aggregations throughout Central America and northern South America. Erythrina glauca swampHonduras, Panama, Trinidad (western margin of Nariva Swamp), Surinam, Guyana, Venezuela (on levees in the lower Orinoco delta, and in the upper delta)Euterpe oleraceae Naidi swampNaidizal (Colombia)Colombia (Choc Biogeographic region, in the delta of the Ro Pata)Freshwater tidal swamp forestHondurasHaematoxylum campechianum forest/ swampTintal/ Bajos or Acalches (Mexico)Cover large areas, ca. 1/3 of central and southern Campeche and central Quintana Roo, MexicoIgapoAmazon basin (Ro Negro, the lower Ro Branco), Orinoco basin Manicaria saccifera palm swampTemichal (Venezuela),Truli bush (Guyana)Patchily distributed throughout the Caribbean coasts of Central America and northern South America: in Guatemala, Belize (Temesh River estuary), Honduras, Nicaragua, Costa Rica (between Tortuguero and Barra del Colorado, a density of 663-910 stems/ha in Tortuguero), Panama, Guyana, Venezuela (behind the mangrove covered Lower Orinoco delta and Cerro Marahuaca),Trinidad and Tobago.Marsh forestWest Indies, TrinidadMauritia flexuosa palm swampMorichal (Colombia, Venezuela), Cananguchal (Amazona Colombiana), Aguajal, (Peru), Buritizal (Brazil)Widely but sporadically distributed throughout northern South America lowlands, east of the Andes, especially in the Amazon region. Large patches are found in Venezuela (Orinoco River basin, especially the lower and central delta), Colombia Llanos, eastern Peru (e.g. Rio Ucayali, with total area greater than 1 million hectares; in the Peruvian Amazon, 6-8 million hectares), and Brazil (Rio Branco and delta of Amazon).Metopium brownei association Chechenal (Mexico)Northern YucatanMontrichardia arborescens Arracacho matorralColombia (Nario, Choc Biogeographic region, in the delta of the Ro Pata)Mora excelsa forest (marsh forest), &/ or M. megistosperma Nato forestNatal (Colombia), Mora forestWest Indies, Trinidad, the Guianas, Costa Rica (Osa Penisula), Panama (Darien), Colombia (Nario).Otoba gordoniaefolia swampGuandal (Colombia)Colombia (Choc Biogeographic region, in the delta of the Ro Pata)Pachira aquatica associationZapotonal (Mexico)Mexico (along Gulf of Mexico from Veracruz to Yucatan and Quintana Roo, and Pacific coast from Nayarit to Chiapas, large stand in La Encrucijada, Chiapas)Palm Marsh forestWest Indies, Trinidad, Guyana, Jamaica (in the Black River morass area)Perched water table forestColombia (Magdalena valley)Prioria copaifera Cativo swamp forestCativo swamp or Catival (Panama, Colombia)Belize, Panama (Darien), Costa Rica (on the Caribbean shore, Cerro Tortuguero, Rio Colorado), Colombia (Choc Biogeographic region, in the delta of the Ro Atrato)Pterocarpus officinalis Suela swamp forestCosta Rica (Limon), Panama, Puerto Rico ( between Luquillo and Bergen), Guadeloupe, Trinidad, the Guianas, Venezuela (lower Orinoco delta), Colombia (tributaries of the lower Magdalena River, Cienaga Grande, and the delta of Ro Atrato)Raphia taedigera Pangana palm swampPanganal (Colombia), Yolillal (Costa Rica)Large patches found in the Atlantic and Pacific coasts of Central America in Nicaragua (Zelaya), Costa Rica (e.g., the Golfo Dulce region, the total area of yolillal in Costa Rica is about 600 km2), and Panama. In South America, large stands are found in the estuary of Ro Atrato, the Golfo de Uraba region, in northwestern Colombia (Antioquia, Choc) and the Amazon River delta (Par).Roystonea oleracea palm swampLesser Antilles (Guadeloupe, Dominica, Martinique, Barbados), Trinidad, Tobago, northern Venezuela, and northeastern ColombiaSeasonally flooded riparian/ riverine/ gallery forestLowlands of Veracruz, Tabasco, and parts of Campeche, MexicoSwamp gallery forestNortheastern Mato Grosso, BrazilSymphonia globulifera forest (marsh forest)Coastal plain of Guyana and Surinam, Venezuela (lower Orinoco delta)Tidal varzeaEstuaries of the Ro Amazonas, and the Ro OrinocoTriplaris surinamensis- Bonafousia tetrastacha forest (marsh forest)Coastal plain of Guyana and SurinamVarzeaAmazon basin (Ro Solimes-Maraon, Ro Solimes-Amazonas, the Madeira, the Purus, between the Japur and I rivers), Orinoco basinVirola spp.Cuangarial (Colombia)Colombia (Choc Biogeographic region, in the delta of the Ro Pata)Sources of Information: Alverez-Lopez, 1990; Bacon, 1990; Brinson, 1990; Daly & Mitchell, 2000; Greller, 2000; Henderson et al., 1995; Lot and Novelo, 1990; Lugo, 1990; Myers, 1990; Pennington and Sarukhn, 1998; Polak, 1992. Table xxx. TNC Platform Sites with lowland moist flooded broadleaf forest TNC platform site CountryBocas del Toro Conservation AreaPanamaRio Bravo Conservation and Management AreaBelizeSerra Do Divisor National ParkBrazilRio Platano Biosphere ReserveHondurasSian Ka'an Biosphere ReserveMexicoCanaima National ParkVenezuelaPacaya_Samiria National ParkPeruLaguna MadreMexico List of globally threatened or endemic species Plant species with restricted distribution are: Tabebuia pallida, which is endemic to the Lesser Antilles, and found in swamp forests; and Pelliciera rhizophorae of mangrove swamps, found in coastal Belize, Atlantic and Pacific coastal Honduras, Nicaragua, Costa Rica, Panama, and coastal Ecuador. The following mammals and reptiles are threatened (Emmons,1990; Lewis et al., 1995). TaxaDistribution Conservation statusAmazonian manatee Trichechus inunguisThe Amazon and lower reaches of its tributaries from Ecuador and N Peru to the estuaries at its mouth and the Rupununi and Essiguibo rivers of Guyana.CITES Apprndix I, US-ESA endangeredWest Indian manatee Trichechus manatusThe coasts of Georgia and Florida (USA), Mexico and Central America; the north coast of South America from Colombia to the mouth of the Amazon in Brazil; and the drainages of the Rios Cauca and Magdalena in Colombia and the Orinoco in Venezuela.CITES Apprndix I, US-ESA endangeredCrocodylus rhombiferCubaCrocodylus morelettiSouthern Mexico to Guatemala and BelizeCaiman crocodylusFind out from website or booksMelanosuchus sp.Find out from website or booksPaleosuchus sp.Find out from website or booksBoutu or pink river dolphin or Amazon river dolphin: Inia geoffrensisIn Amazon and Orinoco river systems from headwaters to oceansCITES Apprndix ITucuxi or gray dolphin: Sotalia fluviatilisSouth America: rivers draining into the Atlantic and Caribbean, and coastal waters to PanamaCITES Apprndix ISouthern river otter Lutra longicaudisCentral and South America: N Mexico south to Uruguay. To 3,000 m elevationCITES Apprndix I, US-ESA endangeredGiant otter, Pteronura brasiliensisSouth America: east of the Andes from southern Venezuela and Colombia south to northern Argentina.CITES Apprndix I, US-ESA endangeredJaguar, Panthera oncaNorth, Central and South America: Mexico to Argentina, to 2,000 m elevation. Overhunted for the fur trade and loss of habitat by deforestation.CITES Apprndix I, US-ESA endangeredRed howler monkey, Alouatta seniculusEast of the Andes in Colombia, Venezuela, Trinidad, the Guianas, and Brazil north of the Amazon, and Ecuador, Peru, Bolivia, and Brazil west of the Purus. To 1,200 m elevation. Intensively hunted for meat.CITES Apprndix IIPygmy marmoset, Cebuella pygmaeaEast of the Andes in Colombia, Ecuador, Peru, and Brazil (Acre), from the base of the Andes east to the Rio Purus. Patchy distribution, threatened due to deforestation.CITES Apprndix IIRed or white uakari monkey, Cacajao calvusCentral Amazon Basin of Brazil, Colombia, and Peru. The white form is found on the large delta island in the Rio Solimes below the mouth of the Rio Japur; the red form found south of the Solimes and west of the Juru, and between the Ro Ucayali and Putumayo in Peru. Hunted for food in Peru and for bait in Brazil.CITES Apprndix I, US-ESA endangeredBlack uakari monkey, Cacajao melanocephalusThe upper Amazon Basin north of the river in SE Colombia, S Venezuela, and adjacent Brazil. Hunted for food in ColombiaCITES Apprndix I, US-ESA endangeredGiant Amazonian turtle or arrau, Podocnemis expansaOn few islands in midstream of both the Orinoco and the Amazon Rivers. Egg collecting for food. Other threatened riverine fauna due to hydrologic alteration reported recently by Pringle et al. (2000) are: TaxaDistributionFreshwater shrimps: Macrobrachium, Atya, and Xiphocaris spp.In streams of the Caribbean islands and mainland tropical streamsCoporo: Prochilodus and Semaprochilodus spp.In major river basins of South America Literature Cited Alverez-Lopez, M. 1990. Ecology of Pterocarpus officinalis forested wetlands in Puerto Rico. Pp.251-265. In A.E. Lugo, M. Brinson and S. Brown (eds.) Ecosystems of the world 15. Forested wetlands. Elsevier. Bacon, P.R.1990. Ecology and management of swamp forests in the Guianas and Caribbean region. Pp. 213-250. In A.E. Lugo, M. Brinson and S. Brown (eds.) Ecosystems of the world 15. Forested wetlands. Elsevier. Brinson, M.M. 1990. Riverine forests. Pp. 87-141. In A.E. Lugo, M. Brinson and S. Brown (eds.) Ecosystems of the world 15. Forested wetlands. Elsevier. Daly, D.C. and J.D. Mitchell. 2000. Lowland vegetation of Tropical South America. Pp. 391-453. In D. L. Lentz (ed.) Imperfect balance: landscape transformations in the Precolumbian Americas. Columbia University Press, New York. Davis, S.D., V.H. Heywood, O. Herrera-MacBryde, J. Villalobos, and A.C. Hamilton (eds.) 1997. Centers of plant diversity: a guide and strategy for their conservation. Volume3, The Americas. World Wide Fund for Nature and World Conservation Union, Cambridge, United Kingdom. De la Rosa, C. 1995. Middle American streams and rivers. Pp. 189-218. In C.E. Cushing, K.W. Cummins and G.W. Minshall (eds.) Ecosystems of the World 22: river and stream ecosystems. Elsevier. Emmons, L.H. 1991. Jaguars. In J. Seidensticker & S. Lumpkin (ed.) Great cats. Fog City Press, San Francisco. ___________. 1990. Neotropical rainforest mammals: a field guide. The University of Chicago Press, Chicago. Godoy, J.R., G. Petts and J. Salo. 1999. Riparian flooded forests of the Orinoco and Amazon basins: A comparative review. Biodiversity and Conservation 8(4): 551-586. Goulding, M. 1993. Flooded forests of the Amazon. Scientific American 266 (3): 114-120. Greller, A.M. 2000. Vegetation in the floristic regions of North and Central America. Pp. 39-87. In D. L. Lentz (ed.) Imperfect balance: landscape transformations in the Precolumbian Americas. Columbia University Press, New York. Henderson, A., G. Galeano and R. Bernal. 1995. Field guide to the palms of the Americas. 352 Pp. Princeton University press, Princeton, New Jersey. Henderson, P.A. and B.A. Robertson. 1999. On structural complexity and fish diversity in an Amazonian floodplain. Pp.45-58. In C.Padoch, J.M. Ayres, M. Pinedo-Vasquez and A. Henderson (eds.) Vrzea: diversity, development, and conservation of Amazonias whitewater floodplains. The New York Botanical Garden. Lewis, W.M.Jr., S.K. Hamilton and J.F. Saunders III. 1995. Rivers of northern South America. Pp. 129-256. In C.E. Cushing, K.W. Cummins and G.W. Minshall (eds.) Ecosystems of the World 22: river and stream ecosystems. Elsevier. Lewis, W.M.Jr., S.K. Hamilton, MA. Lasi, M. Rodrguez and J.F. Saunders III. 2000. Ecological determinism on the Orinoco floodplain. BioScience: 50(8): 681-692. Linna, A. 1993. Factores que contribuyen a las caracteristicas del sedimento superficial en la selva baja de la Amazonia peruana. Pp.87-97. In Kalliola, R., M. Puhakka and W. Danjoy (eds.). Amazonia Peruana: vegetacin humeda tropical en el llano subandino. Gummerus Printing, Jyvskyl, Finland. Lot, A. and A. Novelo. 1990. Forested wetlands of Mexico. Pp. 287-297. In A.E. Lugo, M. Brinson and S. Brown (eds.) Ecosystems of the world 15. Forested wetlands. Elsevier. Lugo, A.E. 1990. Introduction. Pp.1-14. In A.E. Lugo, M. Brinson and S. Brown (eds.) Ecosystems of the world 15. Forested wetlands. Elsevier Lugo, A.E., S. Brown and M.M. Brinson 1990. Synthesis and search for paradigms in wetland ecology. Pp. 447-460. In A.E. Lugo, M. Brinson and S. Brown (eds.) Ecosystems of the world 15. Forested wetlands. Elsevier. Myers, R.L. 1990. Palm swamps. Pp.267-286. In A.E. Lugo, M. Brinson and S. Brown (eds.) Ecosystems of the world 15. Forested wetlands. Elsevier. Pennington, T.D and J. Sarukhn. 1998 rboles tropicales de Mxico. Ediciones Cientificas Universitarias. Polak, A.M. 1992. Major timber trees of Guyana: a field guide.The Tropenbos Foundation, Wageningen, The Netherlands. Pringle, C.M., M.C. Freeman and B. J. Freeman. 2000. Regional effects of hydrologic alterations on riverine macrobiota in the New World: tropical-temperate comparisons. BioScience: 50(9): 807-823. Terborgh, J. and E. Andresen. 1998. The composition of Amazonian forests: patterns at local and regional scales. Journal of Tropical Ecology 14: 645-664. Recommended resources Balslev, H., J. Luteyn, B. Ollgaard and L.B. Holm-Nielsen. 1987. Composition and structure of an adjacent flooded and floodplain forest in Amazonian Ecuador. Opera Botnica 92: 37-57. Duivenvoorden, J. M. L. 1993. Ecologa del paisaje del medio Caquet. Memoria explicativa y mapas. En : J.G. Saldarriaga and T. van der Hammen (eds.) Estudios en la amazona Colombiana. TROPENBOS-COLOMBIA IIA: 301 pp. y 11 mapas. Bogot. Galeano, G. 2001. Estructura, riqueza y composicin de plantas leosas en el golfo de Tribug, Choc, Colombia. Caldasia 23(1): 213-236. Galeano, G., S. Suarez and H. Balslev. 1998. Vascular plant species count in a wet forest in the Choc area on the Pacific coast of Colombia. Biodiversity and Conservation 7:1563-1575. Goulding, M. 1980. The fishes and the forest: explorations in the Amazonian natural history. University of California Press, Berkeley. Goulding, M. 1993. Flooded forests of the Amazon. Scientific American 266(3): 114-120. Goulding, M., N. J. H. Smith and D. Mahar. 2000. Floods of fortune: ecology and economy along the Amazon. Columbia University Press. New York. 184 pp. Junk, W.J. 1989. Flood tolerance and tree distribution in central Amazonian floodplains. In L.B. Holm-Nelsen, I. C. Nielsen, and H. Balslev (eds.), Tropical forests, botanical dynamics, speciation, and diversity, pp. 47-64. Academic Press. San Diego, California. Junk, W.J. 1993. Wetlands of tropical South America. In D.F. Whigham et al. (eds.), Wetlands of the World I pp. 679-739. Junk, W.J. 1997. The Central Amazon Floodplain: Ecology of a Pulsing Sysytem. Springer-Verlag, New York. Junk, W.J., P.B. Bayley and R.E. Sparks. 1989. The Flood pulse concept in river-floodplain systems. In D.P. Dodge (ed.), Proceedings of the International Large River Symposium (LARS). Canadian Special Publications in Fisheries and Aquatic Science 106:110-127. Junk, W.J., J.J. Ohly, M.T.F.Piedade and M.G.M. Soares (eds.). 2000. The Central Amazon floodplain: actual use and options for sustainable management. Leiden, Backhuys Publishers. 590 pp. Padoch, C., J. M.Ayres, M. Pinedo-Vasquez, and A. Henderson. 1999. Vrzea: diversity, development and conservation of Amazonias white-water floodplains. New York Botanical Garden Publications, New York. 407 pp. Rangel-Ch., J.O. 1995. Colombia Diversidad Bitica. I. INDERENA Universidad Nacional de Colombia. Urrego, L. E. 1990. los Bosques inundables del Medio Caquet (Amazona Colombiana) Ph.D. Tesis. 240 pp. University of msterdam. The Netherlands. Tambin publicados en TROPENBOS (1994). 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 4.4. Montane cloud forest General description and geographic variation Cloud forests are easily distinguishable from other forest types by the abundance of epiphytes and reduction in woody climbers. Epiphytes such as bryophytes (liverworts and mosses), ferns and allies, and flowering plants (mainly Bromeliaceae, Orchidaceae and Araceae) are significant in both species diversity and biomass (Luteyn and Churchill, 2000). With increasing elevation, the canopy height of cloud forests is lower than that of lowland forests; trees exhibit compact crowns and gnarled trunks; buttresses, lianas, palms, and leaves tend to be smaller, thicker, and harder, apparently an adaptation to suppressed transpiration due to high atmospheric moisture (Bruijnzeel and Proctor, 1995). The geographic distribution of cloud forests is here used to represent the entire geographic distribution of montane broadleaf forest. Neotropical cloud forests extend from 230N to 250S, roughly from mid-Mexico to northwestern Argentina. The typical cloud forest, humid and dense, is generally found on mountain ranges, from 1000 to 3000 m, with relatively continuous cloud covers at the vegetation level, blanketing the forest. The northernmost stand of cloud forest appears to be the Rancho del Cielo, at 230N in the Sierra Madre Oriental of Mexico, between 1000 and 1500m. The southernmost cloud forest located at approximately 250S in tropical northwestern Argentina (Webster, 1995). According to Webster (1995), major cloud forest biogeographical regions are (1) cis-Tehuantepec Mexico (Tamaulipas to Oaxaca and Vera Cruz), (2) trans-Tehuentepec Mexico (Chiapas and Tabasco) and northern Central America (Guatemala to Nicaragua), (3) Costa Rica and western Panama, (4) northern Andes (Venezuela, Colombia and Ecuador), (5) central Andes (Peru and northern Bolivia), (6) southern Andes (southern Bolivia, Argentina and Chile), (7) coastal ranges of southeastern Brazil, (8) Guayana Highlands, (9) coastal Venezuelan ranges, (10) West Indies (Cuba, Hispaniola, Jamaica, Puerto Rico and the highest peaks of the Lesser Antilles). In Mexico, cloud forests generally appear as isolated patches and are surrounded by xeric vegetation. Cloud forests are most extensive and species diverse in the northern Andes (Venezuela, Colombia and Ecuador), where they form vegetation belts between lowland rain forests and elfin forests or pramos. In the southern Andes (southern Bolivia, Argentina, and Chile) they grade into temperate rain forests or puna (Luteyn and Churchill, 2000; Webster, 1995). In the northern Andes there are cloud forests on both western and eastern slopes. Increasing aridity south from the equator limits cloud forests to an ever narrower band on the western slope. South of 70S, forests are restricted to isolated zones that are climatically buffered, and the predominant slope vegetation becomes chaparrals, thorn scrubs and deserts (Davis et al., 1997). The available data set from fifty-three 0.l-ha sample plots shows that Andean forests are floristically similar to lowland Amazonian forests up to 1500 m. Species of Lauraceae, followed by Melastomataceae and Rubiaceae, are predominant woody floristic elements of Andean forests between 1500 and 2900 m. Floristically, Central American montane forests differ from Andean forests in their greater numbers of Laurasian (Northern Hemisphere) taxa (Gentry, 1995). Many botanists consider the montane forest of the northern Andes a region of exceptional biological diversity, greater than that of the Amazon basin (Webster, 1995). In eastern Peru, major altitudinal changes in species composition of birds, bats, mice and flora are found at around 1500 m (Young and Len, 1999); this altitudinal threshold may be repeated in other localities. Recent studies reveal that the eastern slope of the tropical Andes has the highest bird diversity in the world, with most species found near the Equator. Information on endemic and threatened birds in the worlds tropical montane forests indicates that South America has the highest proportion of restricted-range species confined to cloud forest habitats. Species that are regional endemics and restricted to cloud forest habitats are listed in Table 3 (Long, 1995). Mammal diversity, too, is high in cloud forests. Some threatened mammals of cloud forests are listed in Table 4 (Eisenberg, 1989). Community types/zonation and major gradients within the system (patterns) On tropical mountains, montane forest vegetation forms distinguishable zonation along elevation gradients, from about 600 to1500m to a range between 3000 and 3800m. Ecologists recognize three vegetation belts within montane forests: lower montane rain forest, upper montane rain forest (cloud forest), and subalpine rain forest. Table 1 compares the salient features of the three vegetation belts of montane broadleaf forest, lower montane rain forest, upper montane rain forest, and subalpine rain forest, mentioned in Luteyn & Churchill (2000) and Webster (1995). Synonyms for the three vegetation belts are listed in Table 2. Table 1. FeaturesLower montane rain forestUpper montante rain forestSubalpine rain forestAltitudinal range500 2500 m2500 3500 m3500 4000 mMean annual temperature20-120C12 60C8 50C Mean annual precipitation1000 5000 mm1500 2000 mm (with relative humidity > 90%)1000 2000 mmCanopy height15 - 45 m20-30 m12 20 m StrataTwoOne or twoOneButtresses & stilt roots on treesInfrequentRare noneNoneCauliflory Infrequent (few bat-pollinated species)Rare noneNoneDrip tips on leavesFrequentRare noneNoneDominant leaf-size class of trees and shrubsMesophyll (2000 18000 mm2)Microphyll (200 2000 mm2)Nanophyll (20 -200 mm2)Vascular epiphytesAbundantAbundantOccasionalNonvascular epiphytesOccasionalAbundantAbundantCharacteristic montane forests taxaLaurasian (northern hemispere) taxa: Aquifoliaceae, Buddleiaceae, Clethraceae, Cyrillaceae, Illiciaceae, Magnoliaceae, Rhamnaceae, Sabiaceae, Staphyleac eae, Styracaceae, Symplocaceae. Gondwana (southern hemisphere) taxa: Bromeliaceae, Brunelliaceae, Lobeliaceae Clusiaceae, Cyclanthaceae, Hymenophyllopsidaceae, Melastomataceae, Metaxyaceae, Myrtaceae.Holarctic taxa: Berberidaceae, Betulaceae, Caprifoliaceae, Cornaceae, Ericaceae, Hamamelidaceae, Juglandaceae, Myricaceae, Rosaceae, Ulmaceae. Antarctic taxa: Cunoniaceae, Dicksoniaceae, Elaeocarpaceae, Escalloniaceae, Loxsomataceae, Lophosoriaceae, Podocarpaceae, WinteraceaeBamboos, Coriaria, Drimys, Gaultheria, Weinmannia, PolylepisPlant endemism MediumHighLow Table 2. Synonyms for the three vegetation belts of montane forests. Montane Forest SystemSynonymsLower montane rain forestPremontane forest, paratropical forest, selva submacrotermica, selva mesotermica, tierra templada, subandean rain forest, mist forest, bosque montano inferior (Colombia), yungas (Bolivia), laurel forest of northern Argentina.Upper montane rain forestSelva andina, selva pluvial, submesotermica, tierra fria, cloud forest, bosque montano superior (Colombia), ceja (Colombia), medio yungas (Bolivia), nogal-pino forest (Argentina)Subalpine rain forestBosque andino, mata andina, ceja andina, ceja de monte Table 3. Reported threatened bird species that are regional endemics and restricted to cloud forest habitats. Scientific NameCommon NameRegion (country)NotesDendroica angelaeElfin-woods WarblerCaribbean (Puerto Rico)Amazona imperialisImperial ParrotCaribbean (Dominica)Population < 100 in 1992Leucopeza semperiSempers WarblerCaribbean (St. Lucia)Not found recentlyOreophasis derbianusHorned GuanCentral AmericaOtus clarkiiBare-shanked Screech OwlCentral AmericaMargarornis bellulusBeautiful TreehunterCentral AmericaChlorospingus inornatusPirre Bush TanagerCentral AmericaChlorospingus tacarcunaeTacarcuna Bush TanagerCentral AmericaCampylopterus ensipennisWhite-tailed SabrewingSouth America (Venezuela, Tobago)Vulnerable/ Rare (IUCN)Hylonympha macrocercaScissor-tailed HummingbirdSouth America (Paria Penisula, Venezuela)Vulnerable/ Rare (IUCN)Premnoplex tateiWhite-throated BarbtailSouth AmericaBasileuterus griseicepsGrey-headed WarblerSouth America (Cordillera de Caripe, Venezuela)Endangered (IUCN) Myioborus pariaeYellow-faced RedstartSouth America (Paria Penisula, Venezuela)Endangered (IUCN)Pauxi pauxiHelmeted CurassowSouth America (Venezuela, Colombia)Endangered (IUCN)Grallaria chthoniaTachra AntpittaSouth America (Venezuela)Endangered (IUCN)Grallaricula cucullata Hooded AntpittaSouth AmericaGrallaria kaestneriCundinamarca AntpittaSouth AmericaAglaiocercus coelestisViolet-tailed SylphSouth AmericaBangsia aureocinctaGold-ringed TanagerSouth AmericaPyrrhura orcesiEl Oro ParakeetSouth AmericaXenoglaux loweryiLong-whiskered OwletSouth AmericaAulacorhynchus huallagaeYellow-browed ToucanetSouth America (Departments of San Martn and La Libertad, Peru)Grallaria blakeiChestnut AntpittaSouth AmericaGrallaria carrikeriPale-billed AntipittaSouth AmericaGrallaria przewalskiiRusty-tinged AntpittaSouth AmericaScytalopus macropusLarge-footed TapaculoSouth AmericaHemispingus rufosuperciliarisRufous-browed HemispingusSouth AmericaOtus marshalliCloud-forest Screech OwlSouth AmericaAndigena cucullataHooded Mountain-toucanSouth AmericaHapalopsittaca fuertesiAzure-winged ParrotSouth America (Colombia, Ecuador)Endangered (IUCN)Hapalopsittaca pyrrhopsRed-faced ParrotSouth America (Peru)Endangered (IUCN)Eriocnemis nigrivestisBlack-breasted PufflegSouth America (Colombia, Ecuador)Endangered (IUCN)Grallaria milleriBrown-banded AntpittaSouth America (Colombia, Ecuador)Endangered / Extinct (IUCN) Table 4. Some reported threatened mammals of cloud forest. Scientific NameCommon NameDistributionTremarctos ornatusAndean bearFrom Panama through Peru to BoliviaTapirus pinchaqueWoolly tapirIn pramos and cloud forests of Colombia, Ecuador and Peru Lagothrix flavicaudaYellow-tailed woolly monkeyIn Intact montane forests above 2000 m in Ecuador Departments of Amazonas and San Martn, Peru Aotus miconaxNight monkeyAmazonas, Peru Ecological integrity factors for landscape context Table xxx. Ecological integrity factors for landscape context of montane cloud 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 cloud forest vegetation type.More than 1,000 mm annual rainfall (without including horizontal precipitation) Horizontal precipitation (mist/cloud): 5-20% or more of annual rainfall ( Bruijnzeel and Proctor, 1995) Maximum # of days without clouds. [Experts, please provide an estimate of max. # of days without clouds or literature citation. Thanks.] Evapotranspiration rate: 570-770 mm/year (Bruijnzeel and Proctor, 1995). Mean annual temperature: 8-220CAbundance of epiphytes (bryophytes or lichens). Observations of clouds. Measurement of desiccation of mosses and bryophytes.Topography Relief (slope position and stability) and landslide regime Slope position can influence the characteristics of cloud forests Slope stability determines the frequency of landslides triggered by earthquakes and high rainfalls (Stern, 1995). Landslide regime in turn determines landslide disturbance patterns and creates landscape heterogeneitySlope position: e.g., on a convex or concave slope, or on a cliff, a ridge or in a riverine. The tallest cloud forests are often found on mid-slope with moderate steepness moderate and relatively deep soils (Young and Len, 1999). Slope position can influence the characteristics of cloud forests Monitoring landslide factors: date of occurrence, causes, size of affected area, direction of landslide.Dynamics of tree falls or branch fallsMaintain the diversity and composition of species and forest vertical structure Branch falls and tree falls provide appropriate seed bed conditionsForest canopy turnover rates: 50 -300 years (Kappelle et al, 1996b). If forest canopy turnover rate is too high, then we prevent adequate development of big trees and the structural diversity that supports various life forms such as epiphytes.Presence of abundant lianas, dead trees, fallen woody materials, tip-ups, seedlings and saplings indicate that forest canopy turnover rate is very high.Succession after disturbance (wind, hurricanes, landslides).Increases heterogeneity of vegetation structure and between-habitat (beta) diversity, important for patch dynamics. Pioneer species vary accordingly to different sites (e.g., shade-intolerant colonizing species, Blechnum, Equisetum, Piper, Baccharis, Senecio, Miconia, Chusquea, are found on Pasochoa Volcano, Ecuador (Stern, 1995). 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.Connectivity with montane forest systems or other natural systems. Important for gene flow, seeds dispersion, faunas daily and seasonal migrations. Estimate the distance threshold between two forest fragments. [Experts, please provide concrete examples of distance thresholds from sites you know or from literature. Thanks. Maarten Kappelle: This really depends on the species; there is no info yet available.] Presence of species (e.g., monkeys and large predators) that use other ecosystems.See Thomas van der Hammen (unpublished)- Proyecto demonstrativo de restauracin en Sabana de Bogot, Colombia. Eileen Helmer, Costa Rica/Puerto Rico mapas de fragmentos de bosque de montaa (Helmer, 2000). Produce forest maps and perform analysis of fragmentation and fragment configuration. Studies of indicator species e.g., Oak trees (Quercus), Jaguars (Felis). Presence of populations of indicator or sensitive species (e.g., Coyote, Canis letrans, can be an indicator of fragmentation in Central America.) Evidence of human activities in the area.Hydrological regime and fluvial dynamics Along with the vegetation cover and structure, hydrological regime and fluvial dynamics determine the water yield and runoff rates. Water yield: annual streamflow totals = XX% of annual precipitation Run-off rate: XX See references: Hunzinger, Argentina 1997, Moutain Research and Development [Experts, please complete the citation in the References section. Thanks.];, Veneklaas PhD thesis, International Journal of Hydrology, Journal of Tropical Ecology, Zadroga (1981), and Bruijnzeel and Proctor (1995). Indicators 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 montane cloud forest 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 PriorityViable populations of carnivores (important carnivores include jaguar, Panthera onca; puma, Puma concolor ; yaguarondi , Herpailunus yaguarondi; Andean bear, Tremarctos ornatus; tayra, Eira barbara; weasels, Mustela spp. , and hog-nosed skunks, Conepatus spp.)Regulate the populations of small mammals, and impact food chain Control the density, distribution, and condition of rodents that might harm plant populations.Minimum population sizes of Felis spp. that control rodent communities. [Experts, if you know of any published studies, please cite. Thanks.]Fecal transects of Felis spp. Community-based hunting recordsViable populations of evergreen treesMaintain the continuous accumulation of leaf litter due to partial defoliation.Minimum population sizes of evergreen trees. [Experts, if you know of any published studies, please cite. Thanks.]Record the depth of the leaf litter layer at regular time intervals. Viable populations of pollinatorsRegulate pollination of pioneer, secondary and primary forest tree, shrub and herb species.Minimum population sizes of insects, hummingbirds and bats. [Experts, if you know of any published studies, please cite. Thanks.]A few pollination studies in cloud forests suggest a decreasing abundance and diversity of bees and lepidopterans and increasing plant taxa adapted to dipteran and hummingbird pollination at higher elevation. For example, hummingbirds Trochilinae, abundant in cloud forests at 1000 2000 m, are important pollinators for Centropogon, Fuchsia, Vaccinieae, and bromeliads; the dipterans important for Pleurothallid orchids (Webster, 1995).Observational censuses at regular time intervals. Viable populations of epiphytes Important for processes of mutualism and symbiosis e.g., bromelia tanks support tadpoles and insects. Generate tree and branch fall gaps due to mechanical abrasion as a result of heavy weight on tree branches and trunks, and maintain habitats for e.g. frogs. Maintain the natural hydrologic regime: retain water and avoid run-off and soil erosion, and thus reduce upslope soil loss and downslope flooding in lowlands.Minimum population sizes of epiphytes in general. Minimum population sizes of tank bromeliads, mosses and hepatics. [Experts, if you know of any published studies, please cite. Thanks.]Estimate epiphytes (e.g., bromeliads) abundance and biomass at regular time intervals. Presence of sensitive species or species susceptible to changes (e.g., amphibians). Estimate forest edge run-off at regular time intervals.Viable populations of frugivorous and granivorous species Regulate seed dispersal of pioneer, secondary and primary forest tree and shrub species.Minimum population sizes of frugivorous and granivorous species. See Nadkarni and Wheelwright (2000) for quetzal and Lauraceae. also Guatemala, various papers in Biotropica, Carolina Murcia. [Experts, please complete the citation in the References section. Thanks.] Observational censuses at regular time intervals. Viable populations of 'key' species (e.g. species of Lauraceae).Maintain food availability in times of food scarcity, for a variety of species groups (birds, rodents, etc.) and maintain seasonal species migration routesMinimum population sizes of key species (e.g. lauraceous spp.). See Nadkarni and Wheelwright (2000) for quetzal and Lauraceae. Tree species inventories and phenological records at regular time intervals Viable populations of mycorrhiza Maintain decomposition and symbiotic relations with key tree species such as oaksMinimum population sizes of mycorrhiza. Monteverde, Muller and Halling. [Experts, please complete the citation in the References section. Thanks.Record the presence of mycorrhiza at regular time intervals. Presence of mycorrhiza on top soil. [Experts, please verify. Thanks.]Viable populations of herbivores and decomposers. Maintain the nutrient availability to species at ground level in closed forest.Minimum population sizes of herbivores and decomposers. [Experts, if you know of any published studies, please cite. Thanks.]Censuses of herbivores and decomposers at regular time intervals. Vegetation structure (size/age classes, strata)Provide the diversity of microhabitats and niches within the vegetation. Jaime Cavelier Bosques de alizo comparado con plantaciones. [Experts, please complete the citation in the References section. Thanks. ]Measure the complexity- # of stratus, # of life forms, structure and size, # of functional groups, # of guilds. Ecological integrity factors for size Table xxx. Ecological integrity factors for size of montane cloud forest 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 PriorityEXAMPLE: Minimum Dynamic AreaMuch geographical variation.Consider the average size of primary disturbance (e.g., landslides, storms, etc.) and the average return intervals. Buffer this by a multiplier depending on the confidence in your data and the variability of the key factor attributes (size, return interval, etc) around the geometric mean. Information gaps and caveats There is a major gap in information on the natural range of variation and integrity thresholds of key factors. Recommended priorities for conservation-driven research agenda and next steps We should orient our efforts on restoring secondary forests and agroecosystems, creating new reserves and establishing biological corredors (Brown, A.D. & M. Kappelle. 2001. Introduccin a los Bosques Nublados del Neotrpico: Una Sntesis Regional. In: M. Kappelle & A.D. Brown, eds., Bosques Nublados del Neotrpico.). Literature Cited Bruijnzeel, L. A. and J. Proctor. 1995. Hydrology and biogeochemistry of tropical montane forests: what do we really know? Pp. 38-78. In L. S. Hamilton, J. O. Juvik, and F.N. Scatena (eds.) Tropical montane cloud forests. Springer-Verlag, New York. Cavelier, Jaime. Year? - Bosques de alizo comparado con plantaciones. Davis, S.D., V.H. Heywood, O. Herrera-MacBryde, J. Villalobos, and A.C. Hamilton (eds.) Centers of plant diversity: a guide and strategy for their conservation. Volume3, The Americas. World Wide Fund for Nature and World Conservation Union, Cambridge, United Kingdom. Eisenberg, J.F. 1989. Mammals of the Neotropics. The Northern Neotropics. Volume 1: Panama, Colombia, Venezuela, Guyana, Suriname, French Guiana. The University of Chicago Press, Chicago and London. Gentry, A. H. 1995. Patterns of diversity and floristic composition in Neotropical montane forests. p.103-126. In S.P. Churchill, H. Balslev, 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. Guariguata, Manuel. Year? Helmer, E.H. 2000. The landscape ecology of tropical secondary forest in montane Costa Rica. Ecosystems 3: 98-114. Hunzinger, Argentina? 1997. Moutain Research and Development Kappelle, M. 1996a. 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. 336 pp. Kappelle, M., T. Geuze, M. Leal y A.M. Cleef. 1996b. Successional age and forest structure in a Costa Rican upper montane Quercus forest. Journal of Tropical Ecology 12: 681-698. Long, A. J. 1995. The importance of tropical montane cloud forests for endemic and threatened birds. Pp. 79-106. In L. S. Hamilton, J. O. Juvik, and F.N. Scatena (eds.) Tropical montane cloud forests. Springer-Verlag, New York. 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. Muller and Halling. ? Monteverde. Murcia, Carolina. Year? various papers in Biotropica ? Myster, Randy reference ? Nadkarni, N.M. & N.T. Wheelwright 2000. Monteverde: Ecology and Conservation of a Tropical Cloud Forest. Oxford University Press. New York. Stern, M.J. 1995. An inter-Andean forest relict: vegetation change on Pasochoa Volcano, Ecuador. Mountain Research and Development. Vol. 15 (4): 339-348. Van der Hammen, Thomas (unpublished) Year? Proyecto demonstrativo de restauracin en Sabana de Bogot, Colombia. Veneklaas, E.J. 1990. Rainfall Interception and Aboveground Nutrient Fluxes in Colombian Montane Tropical Rain Forest. Ph.D.Thesis. University of Utrecht. 109 pp. Webster, G.L. 1995. The panorama of Neotropical cloud forests. Pages 53-77. In S.P. Churchill, H. Balslev, E. Forero, & J.L. Luteyn (eds.). Biodiversity and Conservation of Neotropical Montane Forests. The New York Botanical Garden, Bronx. Young, K.R. and B. Len 1999. Perus humid eastern montane forests: an overview of their physical settings, biological diversity, human use and settlement, and conservation needs. 97 pp. DIVA, Technical Report no 5. Zadroga, F. 1981. The hydrological importance of a montane cloud forest area of Costa Rica. Pp. 59-73. In R. Lal y E.W. Russell, eds., Tropical Agricultural Hydrology. J. Wiley y Sons. Ecoandes book Volume 1? Recommended resources Churchill, S.P., H. Balslev, E. Forero & J.L. Luteyn. 1995. Biodiversity and Conservation of Neotropical Montane Forests. The New York Botanical Garden. New York. 702 pp. Grau, A. & A.D. Brown 2000. Development threats to biodiversity and opportunities for conservation in the mountain ranges of the upper Bermejo river basin, NW Argentina and SW Bolivia. Ambio 29: 445-450. Grau, H. R. & T.T. Veblen 2000. Rainfall variability, fire and vegetation dynamics in neotropical montane ecosystems in north-western Argentina. Journal of Biogeography 27: 1107-1121. KAPPELLE, M. & A.D. BROWN. 2001(in press). Bosques Nublados del Neotrpico. INBio IUCN FUA. Editorial INBio. Santo Domingo de Heredia, Costa Rica. c. 702 pp. Hamilton, L.S. 1995. Montane cloud forest conservation and research: a synopsis. Mountain Research and Development 15: 259-266. Hamilton, L.S., J.O. Jurik and F.N. Scatena (eds.). 1995. Tropical Montane Cloud Forests. Ecological Studies 110, Springer Verlag, New York. Nadkarni, N.M. & N.T. Wheelwright 2000. Monteverde: Ecology and Conservation of a Tropical Cloud Forest. Oxford University Press. New York. Stadtmuller, T. 1987. Los Bosques Nublados en el Trpico Hmedo. University of the United Nations (Tokyo) and Centro Agronmico Tropical de Investigacin y Enseanza CATIE. Turrialba. 85 pp. Soma. D & Perovic, P. 1999. Planificacin regional de la conservacin en los bosques montanos del noroeste Argentina. En Matteucci, Morelo, Solbrig, Eds. [Experts, please provide the title and publisher of the edited book. Thanks.] 4.5. Dry forest General description and geographic variation Tropical dry forests generally have high temperatures throughout the year, an annual precipitation of less than 1,600 mm with one or two long and pronounced dry seasons. The duration and intensity of drought govern the distribution of dry forests. Physiognomically, dry forests during the rainy season are often similar to tropical humid forests but are generally shorter in stature, lower in biomass, diversity, density of epipyhtes and lianas. However, they are rich in habitat diversity (Frankie, 1997). Typically, tropical dry forests have ca. 50-70 species greater than 7.5 cm DBH in 0.1 ha sample plots. In some of the most arid tropical dry forests, monospecific stands can occur (e.g. Loxopterygium huasango in Tumbes, Peru). Tropical dry forests usually have a high level of endemism. They grow on soils of significantly higher fertility than savannas, usually have a closed canopy with woody floras dominated by Leguminosae and Bignoniaceae and a sparse ground flora with few grasses. Floristically, there is a greater richness and abundance of species of Capparidaceae, Cactaceae, Erythroxylaceae, Zygophyllaceae, Anacardiaceae, Asteraceae, Malvaceae, Lamiaceae, and Leguminosae. Common genera include Acacia, Caesalpinia, Cassia, Mimosa, Tabebuia, Capparis, Byrsonima, Lysiloma, Ceiba, Aspidosperma, and Erythroxylon. Most of the woody plants in dry forests are deciduous (Daly & Mitchell, 2000; Pennington et al., 2000). Most woody plants in riparian habitats of dry forests are functionally evergreen (Frankie et al., 1974). Dry forests occur along the Pacific coast of Mesoamerica, from Mxico to Costa Rica (Guanacaste Province), and also on the Yucatn Peninsula. In South America dry forests dominate the Caribbean coast of Colombia and Venezuela (Provincia Guajira of Cabrera & Willink, 1980), the Pacific coast of northern Peru (Departamento Tumbes) and southern Ecuador, the Caatinga of northeastern Brazil, the Chiquitana region of Bolivia, and the Gran Chaco of SE Bolivia, W Paraguay and north-central Argentina. Parts of the dry forest ecosystem are the piedmont forests of NW Argentina and SW Bolivia, Paran subtropical semideciduous forests in southern Brazil (W of States of Sao Paulo, Paran and Santa Catarina), eastern Paraguay and NE Argentina. Dry forests are also found in Cuba and other Caribbean islands. (Note: For this workshop, we excluded mezquitales, matorrales" and valles secos interandinos.) Community types/zonation and major gradients within the system (patterns) Carmen Josse: Gradientes como humedad y disturbio tambien son importantes en los bosques secos. En el caso de alteraciones antropogenicas frecuentes y de larga historia, hay una tendencia hacia los tipos de matorral, es decir que el bosque poco a poco cambia su estructura y las formas arbustivas empiezan a dominar. Este no es el caso de las quemas, sino de alteraciones como tala selectiva, claro para parcelas agrcolas, y cierto grado de ganadera. Cuando la ganadera se vuelve mas intensa y hay quemas involucradas, estos bosques tienden a convertirse en sabanas. La humedad puede ser un factor importante para la diversidad de comunidades y el gradiente puede darse tanto con relacin a ros y bosques de galera (fuentes de agua), como con relacin a la presencia de cerros que por su posicin captan humedad atmosfrica. Cuando es as, pequeas diferencias de altitud pueden traer importantes cambios en composicin y estructura. Ecological integrity factors for landscape context Table xxx. Ecological integrity factors for landscape context of dry forest Factores ClaveJustificacin para el Factor de SeleccinUmbral Ecolgico: (Umbral de Integridad Mnimo) Justificacin para Umbral de Determinacin (Ej. Rango Natural de Variacin)Indicadores para Monitoreo de CampoFactor De PrioridadHabitats MosaicIt is important to maintain the diversity of animals and plants that use different habitats, but also for special organisms that use a specific type of habitat. It is required to have large areas of dry forest that maintain sufficient patches of the different habitats of sufficient size to maintain the diversity of vertebrates that require different habitats through their cycle of life. This mosaic is maintained by diverse factors as edafics, hydrologic, topographical, precipitation, natural perturbations and anthropogenics. It is necessary that the managers establish their base line in relation to the mosaics, identifying and characterizing the different habitats, for the purpose of monitoring their changes in the time.Connectivity and fragmentationIt is important for species that require large areas to maintain ecologically functional populations as carnivorous and large herbivorous . Migratory animals (Lat.): butterflies, bats, births of north Mexico or America Central (warblers, flycatchers). High (elevated) migration: butterflies, bats, and hummingbird. Marine turtles that use the bays of the coastal dry forest. Continuous and gradient of vegetation? It is required to maintain the connectivity toward inside the dry forest (for example among patches of the same habitat, toward springs, bodies of water separated). Also it is necessary the connectivity for species that present high (elevated) migrations. This conectivity should be maintained toward other systems. In Caatinga exist species that require free water during all year, of such form that they perform migrations toward higher parts ('brejos') in the epoch of dryness.Monitoring changes of use of the soil and habitats fragmentation of infrastructure of roads, mining, hydraulic, etc. Some possible species of monitoring to evaluate the functional connectivity of the system are: jaguar, lipped peccary, and some greedy birds . Nutrients cycleOur group does not possess sufficient experience to develop this factor, although we consider important to mention it. History of human use The Spanish conquerors founded its first cities (many of them are today the capital of Latin America, for example in America Central) upon lands occupied originally by the dry forests. This enlarged its use for firewood or of the ground for agriculture (its soils are the most fertile of the tropics!), and accelerated the destruction and almost disappearance, to times, of this ecosystem.Decrease of the area occupied by these forests by deforestation; disappearance of endemism; lack of new collections of indicator species in the national and regionals herbals.Precipitation Regime In this system the rain is seasonal and predictible up to a point. In these systems also long periods of droughts can be seen. In this way a determinant factor in the structure of the forest is the duration of those periods of drought.Related to climatic change and should be evaluated to a global or regional level.Local rainfall patterns Rainfall patterns from a regional level in a long term. Change evaluation in relation to global phenomenas. Example: El NioFire RegimeThe Dry Forests are not modeling by fires. Their presence can have essential effects. The presence of exotic species such as gramineas (ej. Melinis minutiflora, Hyparrhenia rufa) increases the frequency and intensity of fires in the Dry Forest. [Darin Prado: Attention: Hyparrhenia rufa is not exotic in Brazil, where the botanical type was collected, and asked me if it was in the rest of Latin America.]A dense cover of exotic species (as the buffalo grass in the Sonorense Forest [Experts, please you indicate you the scientific name of "zacate buffel." Thanks.] and the jaragua grass (Hyparrhenia rufa) in Costa Rica (to see Frankie et al., 1997) exceed the threshold [Experts, if you know of any published studies on this specific threshold, please cite. Thanks.] and provokes a change in the composition and structure of the forest.Monitoring of exotic cover. Fire frequency. Monitoring of composition and structure of vegetation burned areas and associated or key fauna. Underground water regimeAvailability of water for deep plants of roots. Maintain springs and dependent bodies of water of the water of the under ground water. These bodies of water are important for aquatic species of flora and fauna - endemic, migratory and native, especially during epochs of droughts.The decrease of the aquiferous below the reach of the most susceptible roots, but are fed from underground water. The drop of the sufficient phreatic level to feed the springs and other dependent bodies of water of springs.Note: The knowledge related to the subsoil water plants dependence is limited. With springs a rank doesnt exists, just the presence or absence of the spring. Monitoring of the phreatic level in wells. Monitoring the changes effect of the phreatic level upon plants and dependent bodies of water of the subsoil water. Hydrologic regime and galery forest connectivityThe maintenance of the integrity of the riparian hbitats is highly dependent of the superficial water flow states. These riparian habitats are runners inside the forest for birds and other animals. The "flash floods" (quick floods provoked by heavy rains) accelerate the processes of erosion when the original vegetable cover has diminished . In the areas of Costa Rica dry forest, the riparian habitats provide accommodation to twice thediversity than the remainder of the forest, this is due to the availability of water, food and refuge and areas of nesting. It is necessary that the managers consider the protection of large trees as of the thicket forest in these habitats.It is necessary to conserve the original structure of the riparian habitats. The riparian habitats are very susceptible to stockbreeding since although the large trees are maintained, small species are lost; also in these places is common to observe the substitution of native trees by exotic fruit trees. Another possible impact is provoked by the development of roads infrastructure that from time to time obstruct the free flow of the water; in these cases is necessary the construction of drains. When the wide of the riparian corridor is diminished for establishment of grazing land, clearing of trees or by cutting them, increase the hydraulic processes of erosion and aeloic, and increase the vulnerability of the trees before the effect of the wind.There is a need to know the minimum width required by these habitats to consider them functional, [Experts, if you know of any published studies, please cite. Thanks.] since in many countries exist laws that set the minimum width acceptable under arbitrary criteria. Connection to systems of dry forest and adjacent mountain: water, gradient or elevation. Ecological integrity factors for condition Table xxx. Ecological integrity factors for condition of dry forest Key Factors Justification for SelectionEcological Thresholds: (Minimum Integrity Threshold) Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityDepredation upon large herbivorous Is important because of its effect upon the composition and structure of the vegetation and as source of food for local people (for example through systems of populations, source, and drain) and native predators. In general terms populations of herbivorous are under their ideal carrying capacity. The carrying capacity capacity of large herbivorous would be able to become 10 times higher in dry forests that in humid forests. This index has been measured in the National Park Kaa-iya in Bolivia and in the Chiquitana (Estancia San Miguelito). Monitoring densities of large herbivorous (metodology of Andrew Noss, et.al.) [Experts, please complete the citation in the References section. Thanks.]Presence of Large Carnivorous Important to maintain populations of species of low level in the trophic chain.In general terms the populations of large carnivorous are very diminished.Information gap: is difficult to establish the minimum levels of threshold.Monitoring of carnivorous through the metodology of the handbook WCS (Wildlife Conservation Society) at present in development. Changes in soil strutureSeed bank of short duration Breeding season/ Phenology -Seeds -Fishes -Bees -ReptilsNon-sexual reproductionPolinization: specially bees, moths, and bats. The majority of the woody and succulent plants are dioecious and/or hermaphrodites and auto-incompatibles, that bloom in certain periods of the dry season or rain season, and therefore they depend on the pollinators for their sexual reproduction. Particularly in dry forests the bees are an important group of pollinators (in America Central around the 70% of the plants are pollinized by bees). In general, large bees build nests in dry season and small bees during two seasons of the year. Other important nocturnal pollinators are bats and small and large moths (Sphingidae).Many of the bees travel large distances and need the diversity of habitats of the dry forest, by which is very important the connectivity of the system. Due to the diversity of plants, pollinators, habitats and the relations among these factors, is therefore difficult to define the most minimum thresholds of integrity. More specifically, pollinators need diverse habitats because of the diverse floral resources, and other life history resources that communities offer, .e.g. plant/soil materials for bee nests; appropriate larval host plants for developing moths, etc." Some bees can survive among years under the soil in case of drought, of such form that the absence of bees in a specific moment, necessarily does not signify a reduction of its populations. Due to the high diversity of species and habitats the effective reproductive population (from 500 to 1000 individuals) will be dispersed upon a very large area. Due to the intrinsic variability of the system, is difficult to determine if a reduction exists in the pollinators and in the levels of fructification that are out of the natural rank of variability.Monitoring the phenology of flowering. It is possible to monitor bees, bats and moths with established methodologies in the literature. (See Frankie et al., 2002 and Haber & Frankie, 1989) for bees and moths; and La Val, 2002 and Heithaus et al., 1975 for bats.) Monitoring the fructification of indicator plants Monitoring large bees key groups. Monitoring bats. Dispertion of seeds The main dispersers of seeds are birds, mammals (bats, deer, tapirs, coatis, raccoons, coyotes and foxs) and ants. The survival of many species of plants depends on their mobility out of the patch where they are found. The reduction of animals by selective hunting, fire (of high intensity in areas with exotic grass) and competence with the cattle. In the Chaco, the over-overgrassing favored the increase of the density of Bromeliads, this provoked that the seeds scattered by the wind never arrived to the soil, being reduced thus the growth of seedlings. It is supposed that similar processes can occur in other dry forests (as exotic gramineas in Mexico and America Central). At present the populations of species for hunting are very reduced (perhaps very near the threshold), presumably this is altering the seeds dispersion processes, is possible this to be reflected in the next 10 or 30 years. The seeds production fluctuation in long-lived plants is not a significant factor; however, if a tendency of reduction in the long time limit exists, this could affect the forest structure. Monitoring the structure of ages of the arboreal species of the forest (Dan Janzen and the CDC of the University La Molina in Peru). [Experts, please complete the citation in the References section. Thanks]. There should be more knowledge about the succession of arboreal species and in which sseral stages are different in the patches. Monitoring the relative abundance of the representatives of the different guilds of dispersors. Bodmer and Damian Rumiz (Bolivia), and Natural Museum Noel Kempff Market. Low Animal Biomas At present populations of species for hunting are very reduced (perhaps very near the threshold). [Experts, if you know of any published studies, please cite. Thanks.] Presumably this is altering the seeds dispersion processes. It is possible this could be reflected in the next 10 or 30 years.Seeds Depredation Groups of granivorous and frugivorous birds and mammals, coleopteral (bruchidos), are the most important groups in the depredation and dispersion of seeds. The unsteadiness in the number of seeds pillagers will produce at the same time an unsteadiness in the processes of dispersion and growth of new seedlings. It is known that ecologically they have disappeared some guilds of seeds pillagers, [Experts, if you know of any published studies, please cite. Thanks.] However the results will be seen to the next 20 years.The effect upon the forest structure is not known for lack of good prior studies to the ecological changes. Herbivory over seedlings or mature plants.Herbivorous is a very important factor upon seedlings since is one of the most vulnerable phases in its cycle of life. The survival of seedlings determines the structure of the Dry Forest. The stockbreeding of goats and cows is affecting the survival of seedlings. This effect is notorious in all the Chaco (Morello and Saravia Toledo, 1959; Lewis, 1991). In Costa Rica the cows are eating leguminous in a selective way, affecting the composition of the forest. The lack of older natives herbivorous is changing the composition of the forest.In Caatinga is notorious the effect of the over-shepherding, being observed that when the cattle is removed (goats and cows) a quick recuperation of the forest is given.Morello and Saravia Toledo (1959). The Forest Chaqueo. II. The Stockbreeding and the Forest in the East of Salta. Rev. Agron. of the Northwest Argentino, Vol. 3, 209-258. Studies being developed in Chamela, Mexico. (Bullock, 1995). Also in Las Gamas, Argentina (Lewis, 1991). Vegetation Structure This factor is one of the most important factors that maintains the functionality of the Forest. Exotic Animals: cows, horses, donkeys, goats, and tilapias (Oreochromis spp.) In the case of cows, goats (very harmful due to that they eat also the roots), horses and donkeys, these affect changing the structure of vegetation and of soil to trampling and favoring the processes of erosion. In the case of tilapias, these affect pillaging or by competence by resources with native species of fish, affecting the structure and composition of the communities of this group. In Ecuador the over-overgrassingchanged the dry forest to savanna or thorny brushwood in some places. [Experts, please specify the places in Equator. Thanks.] Studies in Africa show that the over-overgrassingcontributes to the desertification; in general many systems are not recoverable after the over-shepherding. The experience has shown that the tilapia has provoked strong impacts in the aquatic systems. Its introduction has been favorable for human consumption or simply has been scattered for hydraulic works. There are examples like the ones from Caatinga, in which the start of the recuperation of the forest has been observed (over-shepherding) after a year of stockbreeding had been removed. Also it was observed that when deforestation for artificial meadows and overcharge those plots, recuperation is not observed, and the desertification processes start. See Gordon Frankie for reference of a thesis of Cornell. [Experts, please complete the citation in the References section. Thanks.] In Costa Rica Dan Janzen is handling the stockbreeding to recuperate the Dry Forest. [Experts, if you know they Give' s publication on this subject, please cite. Thanks.] Monitoring of vegetable cover and compact of the soil in cattle areas.  Information gaps and caveats Carmen Josse: Una aproximacin posible a esta falta de informacin, puede ser el hacer evaluaciones comparativas de reservas o areas protegidas con este tipo de ecosistema. Donde se pueda evaluar condicin, tamao y contexto bajo diferentes situaciones. Recommended priorities for conservation-driven research agenda and next steps Gordon Frankie (Frankie et al., 2002, in press): 1. Need more basic plant inventory work. Plant inventory surveys in the Neotropics are needed in order to know the full range of species present in given regions, ecosystems, or habitats. In addition to species lists, the surveys help to: a) determine frequencies of species occurrence, b) define geographic and habitat limits of species, c) construct phenological patterns, and 4) identify which habitats are in urgent need of attention. 2. Need information on generalized versus specialized pollination. The very common occurrence of the small bee/generalist and general insect systems in the dry forest raises many questions about the effectiveness of each visitor type to effect pollination (Johnson and Steiner 2000). More specifically, we need to know the comparative capacity of each visitor type for carrying pollen and which visitors are moving between plants to promote outcrossing. We also need to know more about the periods of stigmatic receptivity in relation to visitor activity, especially for the insect groups that cross over to other pollination systems. This information may become important in dry forests that become heavily impacted by humans in the future. 3. Need to monitor and restore pollinators. There is little doubt that at least some pollinator types, such as large bees, are declining in the dry forest (Frankie et al. 1997) as habitat loss through human development continues. In large areas that still contain substantially protected natural habitat, there is a need to develop long-term monitoring programs for the important pollinator groups, especially large and small bees, bats, and moths. At the same time basic biological and ecological information must be developed for use in restoring populations of pollinators where downward trends are obvious. Sites for conducting this type of research and application must be carefully selected to insure a high probability of success. Historical records of vegetative cover and land use will play an important role in this research. It is important to stress that this kind of work must start NOW while there are still extant and appropriate habitat and pollinator populations (see also discussion on this topic by Janzen, 1974). 4. Need to apply biologists' knowledge to conserve and restore pollinators. There are at least three courses of action that pollinator biologists could pursue to assist declining pollinator populations. First, they can collaborate with other biologists who are also concerned about decline of their specific organisms (e.g. birds, mammals) and habitat. Building a coalition of concerned biologists with integrated management plans for protection of habitat for several threatened species could be effective if land stewards, associated with the habitat, were receptive and willing to participate in some way as stakeholders. Second, biologists could also work in a variety of ways towards conserving areas known to naturally harbor healthy populations of pollinators, and preferably several types. As in the first case, this kind of project will require that pollinator biologists collaborate with land stewards to insure that high quality habitats will be given special attention. Once again, the land steward (owner or manager) should be brought into this kind of project as a participating stakeholder. In the case of a private landowner, the option of creating a conservation easement to protect special areas should be given serious consideration. In areas where much is known about the requirements of pollinators and where decline is strongly suspected, restoration is another possible course of action. Restoration in this case should have the goal of adding known floral and other resources preferred by pollinators. Hands-on work of planting plants or actively enhancing other pollinator resources (e.g. nesting material for bees) has great appeal to private landowners and some public land stewards in contrast to just "setting habitat aside for wildlife." Restoration is proactive and highly visible and can be understood and appreciated by most landowner/stewards. Literature Cited Andrew Noss, et.al. On page 4. Bodmer and Damian Rumiz (Bolivia), y Museo Natural Noel Kempff Mercado?? On p. 6. Bullock, S.H., Mooney, H.A. and Medina, E. (eds) (1995). Seasonally dry tropical forests. Cambridge University Press, Cambridge, U.K. 521 pp. Cabrera, A.L. and A. Willink. 1980. Biogeografa de Amrica Latina. Serie de Biologa, 2nd ed., Secretara General de la Organizacin de los Estados Americanos, Washington, DC. Daly, D.C. and J.D. Mitchell. 2000. Lowland vegetation of Tropical South America. In D. L. Lentz (ed.). Imperfect balance: landscape transformations in the Precolumbian Americas. Columbia University Press, New York. Pp. 391-453. Dan Janzen esta manejando la ganadera para recuperar el Bosque Seco en Costa Rica.?? On p. 8. Dan Janzen y el CDC de la Universidad la Molina en Peru ?? On p. 6. Frankie, G. W. 1997. Endangered havens for diversity. BioScience 47: 322-324. Frankie, G. W., H. G. Baker, and P.A. Opler. 1974. Comparative phenological studies of trees in tropical wet and dry forests in the lowlands of Costa Rica. J. Ecol. 62: 881-919. Frankie, G.W. , W.A. Haber, P.A. Opler, and K.S. Bawa. 1983. Characteristics and organization of the large bee pollination system in the Costa Rican dry forest. In: C.E. Jones and R.J. Little (eds.) Handbook of experimental pollination biology. pp. 411-447. Van Nostrand, Reinhold Inc., New York. Frankie, G.W., S.B. Vinson, M.A. Rizzardi, T.L. Griswold, S. OKeefe, and R. R. Snelling. 1997. Diversity and abundance of bees visiting a mass flowering tree species in disturbed seasonal dry forest, Costa Rica. Journal of the Kansas Entomological Society. 70(4): 281-296. Frankie, G.W. , W.A. Haber, S.B. Vinson, K.S. Bawa, P.S. Ronchi, and N. Zamora. 2002. Flowering phenology, breeding systems, and pollination systems, diversity in the seasonal dry forest. Chapter in G. W. Frankie, A. Mata, and S. B. Vinson (eds.) Biodiversity conservation in Costa Rica: learning the lessons in a seasonal dry forest. University of California Press, Berkeley, CA. (In press). Haber, W.A. and G.W. Frankie. 1989. A tropical hawkmoth community: Costa Rica dry forest Sphingidae. Biotropica 21: 151-172. Heithaus, E. R., T. H. Fleming, and P.A. Opler. 1975. Foraging patterns and resource utilization in seven species of bats in a seasonal tropical forest. Ecology 56: 841-854. La Val, R. K. 2002. An ultrasonically silent night- the tropical dry forest without bats. Chapter in G. W. Frankie, A. Mata, and S. B. Vinson (eds.) Biodiversity conservation in Costa Rica: learning the lessons in a seasonal dry forest. University of California Press, Berkeley, CA. (In press). Lewis, J.P. 1991. Three levels levels of floristical variation in the forest of Chaco, Argentina. J. of Vegetation Science 2: 125-130) Morello, J. y C. Saravia Toledo 1959. El Bosque Chaqueo. II. La Ganadera y el Bosque en el Oriente de Salta. Rev. Agron. del Noroeste Argentino, Vol. 3, 209-258. Pennington, R.T., D.E. Prado and C.A. Pendry. Neotropical seasonally dry forests and Quaternary vegetation changes. Journal of Biogeography: 27: 261-273. Recommended resources Bullock, S.H., Mooney, H.A. and Medina, E. (eds). 1995. Seasonally dry tropical forests. Cambridge University Press, Cambridge, U.K. 521 pp. Frankie, G.W., H.G. Baker, and P.A. Opler. 1974. Comparative phenological studies of trees in tropical wet and dry forests in the lowlands of Costa Rica. Journal of Ecology. 62: 881-919. Frankie, G.W. , W.A. Haber, P.A. Opler, and K.S. Bawa. 1983. Characteristics and organization of the large bee pollination system in the Costa Rican dry forest. In: C.E. Jones and R.J. Little (eds.) Handbook of experimental pollination biology. pp. 411-447.Van Nostrand, Reinhold Inc., New York. Frankie, G.W., S.B. Vinson, M.A. Rizzardi, T.L. Griswold, S. OKeefe, and R. R. Snelling. 1997. Diversity and abundance of bee visiting a mass flowering tree species in disturbed seasonal dry forest, Costa Rica. Journal of the Kansas Entomological Society 70(4): 281-296. Frankie, G.W. , W.A. Haber, S.B. Vinson, K.S. Bawa, P.S. Ronchi, and N. Zamora. 2002. Flowering phenology, breeding systems, and pollination systems, diversity in the seasonal dry forest. Chapter in G. W. Frankie, A. Mata, and S. B. Vinson (eds.) Biodiversity conservation in Costa Rica: learning the lessons in a seasonal dry forest. University of California Press, Berkeley, CA. (In press). Janzen, D.H. 1974. The deflowering of Central America. Nat. Hist. 83: 48-53. Janzen, D.H. (ed.) 1983. Natural history of Costa Rica. University of Chicago Press, Chicago, Illinois. 816 pp. Kessler, M., K. Bach, N. Helme, S.G. Beck, J. Gonzales. 2000. Floristic diversity of Andean dry forests in Bolivia an overview. In: Siegmar W. Breckle, B. Schweizer and U. Arndt. (eds.) Results of worldwide ecological studies. Proceedings of the 1st Symposium of the A.F.W. Schimper-Foundation established by H. and E. Walter, Hohenheim, October 1998. Verlag Gnter Heimbach, Stuttgart. Prado, D.E. and P.E. Gibbs. 1993. Patterns of species distributions in the dry seasonal forests of South America. Ann. Missouri Bot. Gard. 80: 902-927. Prado, D.E. 1993. What is the Gran Chaco vegetation in South America? I. A review. Contribution to the study of flora and vegetation of the Chaco. V. Candollea 48(1): 145-172. Prado, D.E. 1993. What is the Gran Chaco vegetation in South America? I. A redefinition. Contribution to the study of the flora and vegetation of the Chaco. VII. Candollea 48(2): 615-629. Prado, D.E., P.E. Gibbs, A. Pott, and V.J. Pott. 1992. The Chaco-Pantanal transition in southern Mato Grosso, Brazil. In: Furley, P.A., J. Proctor and J.A. Ratter. (eds.) Nature and dynamics of forest-savanna boundaries. Chapman and Hall, London. Prado, D.E. (2000): Seasonally dry forests of tropical South America: from forgotten ecosystems to a new phytogeographic unit. Edinburgh Journal of Botany 57 (3): 437-461. Ruiz Zapata, T. and M. T. K. Arroyo. 1978. Plant reproductive ecology of a secondary deciduous tropical forest in Venezuela. Biotropica 10(3): 221-230. 4.6. Savanna General description and geographic variation The savanna is a structurally simple but spatially patchy ecosystem in tropical and subtropical regions, characterized by a layer of herbaceous plants-mainly C 4 grasses and sedges, and C3 forbs- with varying degrees of shrubs and/or trees. Based on a broad definition of the term, savanna includes associated grasslands and woodlands. The high root/shoot ratio due to a predominant herbaceous layer is a feature that provides savanna ecosystems resistance to stress and disturbance from drought, fire, and herbivory. It is believed that savannas have evolved under disturbance factors like fire, herbivory and drought. Persistence of savannas may depend on such disturbance to preserve stabilizing components and properties (Baruch et al., 1996; Coupland, 1992; Silva, 1996). Savannas occur in hot climates with rainfall varying between 750 to 2000 mm and a dry period of two to six months, forming a transition zone between moist forest and xerophytic vegetation. Rainfall distribution is a major determinant of the savanna vegetation types. Tropical savanna usually develops on nutrient deficient, acidic soils with aluminum toxicity and pronounced alteration of wet and dry conditions (Sarmiento, 1983). In Mexico savannas are best developed in the southeast, in Campeche, Tabasco, Chiapas, and Veracruz. They also occur, much reduced in size, on the Pacific coast from Chiapas to Sinaloa. In Mesoamerica, patches of pine savanna occur in SE Chiapas, Izabl and central Petn of Guatemala, and along the Atlantic coasts of Belize, eastern Honduras, and eastern Nicaragua. The vegetation is characterized by open stands of Pinus caribaea, Curatella ameircana, and Byrsonima crassifolia in a matrix of grasses, sedges and forbs (mainly legumes and composites) (Greller, 2000). In the Caribbean, sizable savannas are found in Cuba. Seasonal and hyperseasonal savannas occupy between 10 and 20% of Cuba. They occur in western Cuba and the Isla de Pinos, and the highlands (at 400 to 600 m) of the Sierra de Nipe in eastern Cuba. Some of them are floristically similar to continental savannas, but many have a high degree of endemism due to different soil types, either silicious or derived from serpentine (Sarmiento, 1983). In South America, savannas cover an area of approximately 2.5 million km2 [see sum of numbers given in the table below]. They include The Cerrados mostly in the tablelands uplands (1,000800 ~ 1,500 m) of central Brazil, extending towards SE and NE in areas with lower elevations. Outliers are found in Paraguay and Bolivia. The Llanos del Orinoco in the central lowlands of Venezuela and northeastern Colombia. The Gran Sabana of southeastern Venezuelan Guayana, n Guayana the Roraima-Rupununi savannas of Brazil and Guyana), and as well as the coastal savannas of the three Guayanas (also called Guianan savannas). The Pantanal region that includes extensive areas of wetlands and savanna in Brazil, Bolivia, and Paraguay. The Llanos de Moxos of northern Bolivia drained by the Beni, Mamor, and Guapor to form the Ro Madeira (covers an area between 11~160S, and 64~690W). Others: Amazonian campos in northern Brazil, Llanos of the Magdalena valley, Colombia (Solbrig, 1996; Daly and Mitchell, 2000). Salient features of several South America savannas reported in Daly (2000), Solbrig (1996), and Sarmiento (1983) are summarized in the following table. SavannaCerrado sensu latoLlanos of Venezuela and ColombiaLlanos de MoxosGran Sabana / Roraima-Rupununi savannasPantanalArea (km2)1,800,000500,000Savannas ca. 150,000 and forests ca. 120,000 km2 54,000 150,000 170,000Elevation range100 ~ 1,500 m0-300 m130-235 m100~1,3500 m~100 mPhysiognomic units or savanna typesCampo limpo, campo sujo, campo cerrado, cerrado sensu stricto , cerrado (sclerophyllous woodland).Hyperseasonal savanna, wetlands (esteros) and seasonal savanna (dunes)Hyperseasonal savanna, wetlands (esteros) and seasonal savannaHerbaceous swamp savannas on the tepuys; seasonal savannas and swamps on white sand, lower tablelands; alluvial fans,hyperseasonal savannas on river flat. Physionomically similar to Llanos, a complex mosaic of wetlands, savannas, and forests. It includes cerrado vegetation with an entire continuum from campo limpo to cerrado. Average annual precipitation1,500 mm (rainfall ranges from 750 mm to 2,000 mm, and increases toward NW and SE).1,000 mm (eastern Llanos) ~ 2,200 mm (SW at the Guaviare River in Colombia). Rainfall is highly seasonal.1,300 mm (E) ~ >2,000 mmLow rainfall (1000-2500 mm; northern end of the roughly NW-SE transverse central eastern Amazonia dry belt).1,000-1,400 mm, mostly during the rainy season (November- April).Dry season3-4 to 7 months (May ~ September). The wet period is likely to be interrupted by a short period of drought that may last for 1 to 3 weeks.1-6 months along a SW-NE gradient.2-3 months dry season (June ~ August) in summer. A long short to medium dry season (2-6 months, December/January to April/May)3-4 monthsTemperature Frost occurs at southern limit of Cerrado.No frost260C in summer, but can reach 6 0C in winter with cold fronts.No frostSoil typeMost dystrophic oxisols, very poor in N, P, Ca, high in aluminum, low pH and cation exchange capacity. On richer mesotrophic soils, dry forest or cerrado may occurs.Mostly alluvial and aeolian sediments, highly leached oxisols and ultisols, low in exchangeable bases and high in aluminum. On richer clays and shales, dry forest occurs. Alluvial sediments, different origin and age, ranging from dystrophic oxisols in the north to less alterated (verwittert) and more nutritive cambi-luvisols, and gleysol, solonetz and hydromorfic soils in the south.Mostly well-drained, but highly oligotrophic soils; many swampy areas with sandy or organic soils and an impermeable hardpan below.Most alluvial and aeolian sediments.Characteristic taxaDominated by woody plants: Dystrophic cerrado: Hirtella glandulosa, Emmotum nitens, Aspidosperma macrocarpon, Vochysia haenkiana, Xylopia sericea Mesotrophic cerrado: Magonia pubescens, Callistheme fasciculata Cerrado sensu stricto, campo cerrado, campo sujo; many species of Fabaceae, Poaceae, Asteraceae, Orchidaceae, Rubiaceae, Myrtaceae, Melastomataceae, Apocynaceae, and Leguminosae. Widespred trees and shrubs are: Qualea grandiflora, Kielmeyera coriacea, Copaifera langsdorffii, Caryocar brasiliense, Byrsonima coccolobifolia, B. crassifolia, Curatella americana. Palms: Butia leiospatha, Syagrus acaulus, Astrocaryum campestre. Grasses: Andropogon, Aristida, Axonopus, Elionurus, Paspalum, Echinolaena and Trachypogon. Dominated by herbaceous plants, especially grasses: and sedges: Trachypogon, Andropogfon, Aristida, Axonopus, Panicum , Paspalum, Mesosetum, Eliionurus, Sporobolus, Trachypogon, and sedges: Bulbostylis, Rhynchospora, Scleria. Palms: Copernicia tectorum, Mauritia flexuosa. Fauna: waterfowl, ibises, storks, capybaras.Dominated by herbaceous plants and shrubs in southern wetlands: Cyperus giganteus, Thalia geniculata, Paspalum,Panicum, Axonopus and palmares of Copernicia alba. In the northern Llanos cerrado like vegetation with grasslands of Leptocoryphium lanatum, Trachypogon plumosa, Macairea scabra, Mesosetum penicillatum, Bulbostylis juncoides and trees and shrubs as Qualea, Byrsonima, Palicourea rigida, Kielmeyera. In depressiones Palms: Mauritia flexuosa, Mauritiella aculeata and grasslands with Burmanniaceae, Eriocaulaceae, Xyridaceae Termites savanna are commun.In grass savannas: Trachypogon, Axonopus, Echinolaena, Aristida, Panicum, Paspalum, Mesosetum, and sedges: Bulbostylis, Rhynchospora, Scleria, Lagenocarpus, Hypolytrum. Palm swamps: Mauritia flexuosa, Andropogon, Scleria, Xyris, Eriocaulaceae, Rapatea, Drosera, Cephalostemon, Abolboda.In hyperseasonal savanna: Woody plants: Curatella americana, Bactris glaucescens, Licania parvifolia, Copernicia alba. Herbaceous plants: Poaceae (Axonopus purpusii, Paspalum almum, Paratheria prostrata, Reimarochloa brasiliensis, Elionurus muticus), Cyperaceae, Asteraceae, etc. On murundus (forest islands on decayed termite mounds): Tabebuia aurea. Gallery forest: Guarea macrophylla, Abuta grandifolia, Mouriri guianensis, Tapirira guianensis, Hymenaea stilbocarpa, Protium heptaphyllum, Xylopia emarginata, Mauritia flexuosa. Dry deciduos or semideciduos forests on limestone outcrops in Central Brazil: Cavalinesia arborea, Tabebuia impetiginosa, Myracruodruon urundeuva. Savannas that were discussed in the Experts Workshop: Sabanas Centroamericanas Llanos de Colombia y Venezuela Sabana Escudo Guayano Llanos de Moxos Cerrado Community types/zonation and major gradients within the system (patterns) Sarmiento (1983) recognizes three types of Neotropical savannas: Semiseasonal savanna which occurs in a rather humid climate with one or two short dry seasons and less frequent fires (some of the Amazonian and Guianan savannas), Seasonal savanna characterized by a severe dry season and frequent fires (e.g., the Cerrado and the Llanos), Hyperseasonal savanna which is the result of excessive drought and fires during dry season and severe flooding during the wet season. This savanna type is common in the poorly drained bottomlands of the Pantanal, Llanos de Moxos, the Roraima-Rupununi savannas, and part of Llanos. Palm stands are often found in water-logged areas. The savanna system is usually heterogeneous, consisting of a mosaic of pure grasslands, patches of trees or shrubs, dry deciduous or semideciduous forests, gallery forests, and sometimes wetlands (Daly and Mitchell, 2000). The distribution of various plant communities in a savanna landscape often follows edaphic gradients (e.g. soil types or levels of water table) and fire regime. Community types within the eight savanna systems that the experts identified. Sabanas Centroamericanaos: 1) hmeda o seca, 2) variacin altitudinal (0-1,500 m), 3) variacin latitudinal Sabanas Antillanas (en Cuba y Repblica Dominicana; palmares en general se encuentran en lugares hmedos, eastacionalmente inundados). Llanos de Colombia y Venezuela: 1) Inundable (e.g., Lllanos de Apure-Arauca: palmares, bancos, bajos y esteros, bajillos), 2) no-inundable (e.g. Llano Alto: sabanas arbustivas: con Curatella americana, Byrsonima crassifolia, Bowdiachia virgilioides; y sabanas abiertas o lisas: con Trachypogon spicatus, Axonopus anceps, Bulbostylis paradoxa, etc.). Sabanas del Escudo Guayano (con alta frecuencia de termiteros). Sabanas Amaznicas: 1) suelo mal drenado, 2) suelo bien drenado (sabana no-inundable). Llanos de Moxos: 1) Sabana del nordeste de Beni, agua estancada por lluvia, parecido a la flora de cerrado (sensu lato). Tambin incluir bosques abiertos, variacin de pastizal hasta bosque abierto con diferente regimes hdricos. Sobre substrato antiguo, muchos termiteros. 2)Sabana del central y sur de Beni: sabana inundable por ro y lluvia, comunidad vegetal se cambia con nivel topogrfico: i) bajura, ii) semi-altura, iii) altura de algunos metros, con suelo aluviales, algunos semi-alturas con pH hasta 9. Cerrado: 1) herbceo: i) bien drenado: campo limpio, campo sujo, campo cerrado, ii)mal drenada: campo limpo hmedo, vereda, campo de murudum. 2) leosa: i) cerrado sensu stricto: latosol, sandstone, ii) cerradao: mesotrfico, distrifico. Sabanas Andinas: incluye Valle de Magdalena y Valle Sinu. Hay manchas pequeas de Trachypogon plumosuso, entre bosque seco y hmedo, zona de colonizacin incaica. Ecological integrity factors for landscape context Table xxx. Ecological integrity factors for landscape context of savanna Key FactorJustification for SelectionNatural Range of Variation and Ecological ThresholdsJustificationIndicatorsFactor RankFire regimeImportant in shaping savanna properties at a local scale. Changes the mix of habitat types by increasing or decreasing the dominance of one type over the other. Modifies community or floristic structure by affecting mortality of many species, especially young plants. Recurrent frequent fires cause nutrient impoverishment.Fire frequency (Frost and Robertson, 1987; Braithwaite, 1996): In the moist savannas fires generally occur at intervals of 1-5 years, whereas in the more arid savannas, the intervals are about 5-50 years. Fires increase patchiness or habitat diversity within the savanna. By optimizing the fire mosaic, savannas can be managed to increase biodiversity. Long-term fire exclusion experiments in Calabozo, Venezuela show (Bulla, 1996): New tree species invade the savanna and increase the diversity of the tree layer. Tree density increases. In the open savanna, evergreen tree species increase rapidly, whereas deciduous trees showed a long lag of at least 16 years. In the groves, evergreen species change little and deciduous species increase in the first 8 years. Fire intensity (Frost and Robertson, 1987; Braithwaite, 1996): Factors that affect fire intensity are fuel types, fuel loads, the occurrence of periods of above-average rainfall, seasons, landscape units and vegetation types. Long intervals between fires increase the fuel load and therefore the probability of a wild fire event. The fire intensity will cause changes in patchiness of habitats and the vertical structure of upper shrub or tree layer and ground herb or shrub layer. Dominance shifts towards to upper shrub or tree layer if fire intensity is low. Conversely, dominance shifts to ground herb or shrub layer if fire intensity is high. Type of fires (Frost and Robertson, 1987): most savanna fires are surface fires, burning through the herbaceous layer, and burn patchily as a result of varying wind speed, topography and distribution of fuel loads. Time of natural fires (Frost and Robertson, 1987; Braithwaite, 1996; Ramos-Neto and Pivello, 2000): Natural fires occur mostly at the beginning of the wet season when the frequency of lightning is highest. Early dry season fires: create greatest patchiness in the ground herb or shrub layer and impact negligibly the upper shrub or tree layer. High intensity late dry season fires typically scorch high into trees and are likely to cause maximum mortality. Impact on fauna (Braithwaite, 1996): Burned sites show increase in ants species diversity due to structural changes in the habitat caused by fire, especially the level of litter accumulation and insulation on the ground. Many bird species that feed on the ground such as granivores, omnivores and carnivores are attracted to areas that have been recently burned. Species of Lepidopterans, herpetofauna, and mammals, respond differently to different fire regimes.Corticolous lichens have been used as potential bioindicators of fire history in the cerrado of central Brazil (Mistry, 1998).Humidity regime This factor should certainly rank in the first place of importance in this Category: Landscape context, before the Fire.Important in shaping savanna structure at a landscape scale. Modifies plant available moisture (PAM). It is related to the length of dry season, precipitation and woody cover (Solbrig, 1996).Pulses in rainfall or drought spells induce important changes in savanna composition. Observations support the following hypotheses. Early onset of rains favors species blooming very early in the wet season (precocious species). A prolonged rainy season favors species blooming at the end of the wet season (late species). A shortening of the rainy season would be detrimental to late species but favorable to species blooming in early or the middle of the wet season (intermediate species) Drought favors deciduous trees or trees with smaller leaf size. A higher rainfall pulse will increase grass biomass during that season, increase standing dead biomass during the next dry season, and increase the probability of fires. [If no fire, shading will increase during the following wet season and reduce grass growth, but increase tree growth and recruitment, alter the tree/grass ratio in the following years, affect grass growth and reproduction, and influence fire and grazing regimes.]Soil type or fertilityImportant in shaping savanna structure at a landscape scale. Modifies plant available nutrients and moisture. Affects savanna productivity.Savanna soils vary widely in particle size, structure, profile, and depth. The nutrient status of the soil is related to the age of the sediments, parent material, and topography. Recent studies show that estimates of productivity of tropical savanna grasslands approximate the figures for tropical forests (Solbrig, 1996).Interactions between plant available moisture (PAM) and nutrients (PAN).Important in shaping savanna structure at a landscape scale.Woody elements dominate where plant available moisture and nutrients have high values. As moisture and/or nutrient levels increase, the savanna gives way to a moist forest eventually. Xerophytic elements prevail where moisture and/or nutrient have low value. If the values of moisture and nutrient get very low, the savanna is replaced by a semidesert (Solbrig, et al., 1996). Interactions between plants and humidity (rainfall, drought) and herbivory during post fire recovery phase (Frost and Robertson, 1987).Cattle grazing (Baruch, 1996): [Cattle were introduced to the Americas in the 16th century.]Important in shaping savanna properties at a local scale. Modifies community or floristic structure.Woody species / grass species ratio or density of trees affects the sensitivity of grass layer to grazing and overgrazing. Grazing pressure by domestic animals plays a role in the dynamics of savanna communities dominated by native and alien grasses, and may favor alien grasses that have higher defoliation tolerance, e.g. African grasses.Connectivity by gallery forest networksImportant corridors linking patches of similar or different ecosystem types, e.g., gallery forest networks of cerrados linking the Amazonian forests to the Atlantic coastal forests of Brazil (Oliveira-Filho and Ratter, 1995; Daly and Mitchell, 2000). Ecological integrity factors for condition Table xxx. Ecological integrity factors for condition of savanna Key FactorJustification for SelectionNatural Range of Variation and Ecological ThresholdsJustificationIndicatorsFactor RankDiversity of above-ground plant functional groups (species that share morphological, chemical, structural or life history characteristics)(Denslow, 1996) Determines the role of biodiversity in ecosystem response to disturbance such as water stress (drought or flood), fire, shading, competition, and herbivory or grazing. Woody species: density and spatial patterns of trees determine the spread of fire and its erratic performance. Herbaceous species: Rapid and homogeneous fires are produced in communities that have a high biomass of grasses and a low fuel biodiversity. The architecture of dominant grasses defines the vertical distribution of fuel. Taller grasses produce higher flame which increases the probability of damage to tree crowns. C4 and C3 plants in savannas result in patchiness of herbaceous layer in terms of productivity and efficiency of water and nutrient use. In seasonal savannas most dominant grasses have the C4 syndrome, and only in very wet environments do C3 grasses become abundant (Baruch et al., 1996) Disturbance-resistant vs. disturbance-sensitive species: Fire-resistant woody species: usually evergreen, sclerophyllous (hard, thick, leathery leaves), and with thick bark or protected buds, or escaping adaptation by underground woody structure such as xylopodia or lignotubers (e.g., Andira humilis and Anacardium humile). They are usually reproductively active during the dry season; and adapt to water or nutrient stress. Fire-sensitive woody species: usually deciduous and mesophyllous. Drought -escaping ephemeral annuals or decicuous perennials: active only in the wet season. Survive the period of drought stress as seed or by going dormant. Drought-enduring evergreens or perennial graminoids and deep-rooted phreatophytes: continue being physiologically active during the period of drought as long as there is sufficient soil moisture. Phreatophytes, represented by the majority of savanna trees and shrubs, can gain access to the water table by their deep roots. Drought-resistant succulents: special morphological and physiological features to maintain physiological activity, even under conditions of drought stress. Species reproduced by seeds vs. vegetative propagules: populations of plants that recover by seeds after fire will be more susceptible to change than those that recover by resprouting. Abundance of dominant species in herbaceous layer and/or woody layer. In Cerrado sensu stricto: 40-100 woody species and three times more herbaceous. One hectareof cerrado contains from 60 to 100 woody species and two times more herbaceous species. One hactare of gallery forest contains from 80 to 200 woody species and a similar number of herbaceous species. One hactare of dry deciduous forest contains 40 to 80 woody species and a similar number of herbaceous species. One hactare of campo limpo contains from 200 to 300 herbaceous species. Dominance of species from the genera Sclerolobium, Mimosa, Solanum and Ouratea can serve as disturbance indicator.Community composition/ Diversity High beta diversity is usually found in areas with mosaic habitats that are created by difference in topography (e.g., valley or ridge crests), or soil types or disturbance of various intensity e.g. fire or flood. The patchy diversity of the community affects fire impacts on species composition (Bilbao et al., 1996).Plant species composition and structureDetermine the fuel mixture, fuel quality and degree of flammability during the event of fire. Affect the types and intensity of fires. Determine the productivity of the savanna. High biodiversity increases the probability of redundancy within functional groups. This may buffer the negative impacts of perturbations to ecosystem processes and products (Denslow, 1996).The available plot data suggests that the range of species diversity for tropical savanna is 7-100 species of trees and shrubs/ha with lower diversity in the Amazonian savannas, e.g. Amap, and higher diversity in central Brazilian cerrado and Roraima-Rupununi savanna (Daly and Mitchell, 2000). Cerrados are the most species rich savannas of the world. About 6.500 species of higher plants have been identified in the Cerrado Dominium (Vania Pivello, Pers. Com.). Plant species composition in savanna often relates to available moisture and nutrients, herbivory, and fire disturbance (Bilbao et al., 1996). Low productivity, stress-driven savanna communities and high productivity, competition dominated ones will have lower species diversity than intermediate ones (Bulla, 1996). Herbaceous plants that are commonly found in the Neotropical savannas are usually members of grass and sedge families. In the Poaceae, Andropogoneae: Trachypogon, Andropogon, Hyparrhenia and Saccharum are usually dominant in the core savanna area; Panicoid: Digitaria, Panicum, Paspalum, Pennisetum, Setaria, dominate in drier environments, and Oryzeae (Oryza, Luziola, Zizania) in very wet and swamp savannas. Cyperaceae includes genera of Cyperus, Rhynchospora, Bulbostylis, and Scleria (Solbrig, 1996). Woody plants widespread throughout the Neotropical savannas are Curatella americana, Byrsonima crassifolia, B. coccolobifolia, B. verbascifolia, Bowdichia virgilioides, Anacardium occidentale, Hancornia speciosa, Salvertia convallariodora (Daly and Mitchell, 2000).Fauna diversityAffect food webs of the savanna system. Neotropical savanna avifauna (Fry, 1983) includes the two most species abundant insectivorous Tyrannidae (tyrant-flycatchers), granivorous Fringillidae (finches), and aboreal frugivorous Aquilidae (parrots), Trochilidae (hummingbirds), Furnariidae (spinetails), Icteridae (caciques) and Thraupidae (tanagers). Endemic bird families: characteristic large ground birds of the open savanna: herbivorous Rheidae (rheas), Anhimidae (screamers), Cariamidae (seriema), and ground-foraging granivorous Tinamidae (tinamous). Reptiles (Barbault, 1983): snakes of Colubridae, Typhlopidae, Boidae, Elapidae and Viperidae, lizards of Gekkonidae, Scincidae and Iguanidae. Rodents and lagomorphs (Happold, 1983): Cricetidae (Calomys, Zygodontomys, Phyllotis, Akodon, Baiomys, and Eligmodontia), Caviomorph rodent (capybara, Hydrochoerus hydrochaeris), and endemic lagomorph, Sylvilagus brasiliensis.Biotic interactions: presence of pollinators (bees, butterflies, moths, bats, and hummingbirds), and seed dispersal agents such as fruit-eating birds (e.g. parrots).Pollinators affect food webs.Wind is important seed dispersal agent: almost 50% of cerrado tree species are dispersed by wind. Birds seem to be the most important pollinators and dispersal agents besides wind. Seedling recruitment for grass species mostly depends on yearly seed production. There is no apparent permanent seed bank of native grasses in the savanna soil.Biotic interactions: presence of large herbivores (e.g., branch-antlered white tailed deer Odocoileus virginianus, pampas deer Ozotoceros bezoarticus, swamp deer Blastocerus dichotomus, and capybara Hydrochoerus hydrochaeris. (Ojasti, 1983). Insects' contribution to consumer biomass is much greater than that of vertebrates in the savannas (Lewinsohn and Price, 1996).Herbivory affects woody/herb plant ratio, vegetation stratification, total cover and biomass (Lewinsohn and Price, 1996).Few large herbivores in Neotropical savanna. Folivores can accelerate nutrient release from living plants. Folivorous insects are much more important than vertebrate herbivores, and their species diversity may be dependent on plant species richness due to host-specificity. Meristem, stem or root feeders can change plant architecture and their susceptibility to fire, drought, and frost damage. Propogule feeders can increase patchiness: In cerrados, seed crops are often almost entirely destroyed by predispersal predation. Peak of insect herbivory: the sunnier or drier part of the year.Biotic interactions: presence of anteaters (Myrmecophaga tridactyla) or armadilos (Dasypus spp., Priodontes maximus) (Rodrguez and Rojas-Surez, 1999) or top predators e.g. jaguar (Felis onca), Orinoco crocodile (Crocodylus intermedius).Savanna is an important habitat for anteaters and armadilos. Top predators control the populations of small mammals and herbivores..etc.Biotic interactions: Presence of decomposers (mycorrhizae, fungi, microbes) and soil macroinvertebrates (termites, nonsocial arthropods, earthearths, leaf-cutter ants)Can increase nutrient supply rates and improve soil structure, including nutrients, moisture and oxygen availability (Denslow, 1996).Biotic interactions: presence of alien vertebrate species: cattle (see grazing)Introduce diseases to native fauna, disseminate exotic herbaceous species, and change species composition of native flora. Its impact can vary from low biomass consumption of natural savanna plant communities during rainy season, to massive plant biomass harvesting of regrowth after burning during dry season, to total substitution of natural savanna plant communities by introduced, artificial fodder grasses. Biotic interactions: presence of alien plant species: e.g. the African grasses that are the most aggressive invaders of Venezuela (Baruch, 1996) and Brazilian savannas, Melinis minutiflora in secondary savannas above 600 m; Hyparrhenia rufa in lowland savannas with poor soils and marked dry season; Panicum maximum in humid and relatively fertile areas; and Brachiaria mutica in periodically flooded savannas. The indigenous community is generally dominated by native grasses Trachypogon, Axonopus, and Bulbostylis.Grass invaders can change composition and structure of the grassland (e.g., shortgrass to tallgrass), and modify factors that control the functioning of savannas such as water and nutrient availability, and fire regime. Often decrease biotic diversityIn Venezuela savannas, the invasive plants usually have higher growth rates, establish themselves rapidly and compete successfully for resources, and displace native species from moist, fertile sites. In contrast, native grasses have higher root/shoot ratios with carbon reserves in underground organs and slower growth rates. They can endure invasion and persist in poorer sites. Studies of Venezuela savannas show that the persistence of some African grasses [depending on the grass species] is sustained by a fire and invasion cycle. Space created after burning is invaded by alien grasses, which in turn promote more frequent and intense fires due to higher biomass, and reinforce the process of colonization and elimination of native competitors. To arrive at thresholds, determine whether the success of invasive species is due to differences in competitive abilities or due to the primary disturbance that removes native species. Invaders may not persist if the disturbance is eliminated. Ecological integrity factors for size Table xxx. Ecological integrity factors for size of savanna 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 PriorityEXAMPLE: Minimum Dynamic AreaConsider the average size of the primary disturbance (fire) and the average return intervals (see above). Buffer this by a multiplier depending on the confidence in your data and the variability of the key factor attributes (size, return interval, etc) around the geometric mean.Minimum Population Size of anteaters or armadillos or jaguarsTop predators in this system are critical to control populations of ants or termites, small mammals and herbivores.   Additional information The experts assessed the biodiversity health of seven savanna systems. The results are presented in the following tables. System: Pine Savannas, Central America, Landscape Context, Condition & Size Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold) Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityFire RegimeSystems dynamic is associated to the fire cycle. The pine savanna requires fire; they are maintained by fire; the species have adaptations to endure the fire; many require the fire to regenerate. Without the appropriate fire regime habitats and species are lost. Fire frequency: 5-20 years for Pinus oocarpa and P. caribaea of Central America. Each 3-10 years for Pinus caribaea in the Bahamas and Cuba, P. tropicalis in Cuba, and P. elliottii in the Keys of Florida. Intensity and duration of the fire and period of occurrence. Size of the natural regeneration. For P. oocarpa and P. caribaea approximately a meter of total height (age: 5-8 years). <5 years: open pine forests in Central America. Pinus caribaea (in Cuba and Bahamas), P. elliottii and P. tropicalis support fires of greater frequency (with P. tropicalis almost annually). Fires of high intensity and duration during times of drought destroy the natural regeneration , the organic material, and define the growth of the pine trees. Regeneration disappears with fires of high intensity and duration.Monitoring through permanent plots. Fire history (place, extension, intensity, and time of the year). Fire mapping. Sources of ignition; areas of high riskHydro regime seasonLimitation of water during certain part of the year.Pinus caribaea: 3-9 months of dry season.Extreme periods of drought weaken the pine trees and make them susceptible to plague and/or illnesses attacks. Records of the periodicity and intensity of the drought. Drainage, depth and fertility of the soil.Density of pine stands and floristic composition of the undergrowth.Age structure Determinant to guarantee the dynamics of the system. The age structure allows maintaining populations of birds that guarantee biological control against endemic plagues. Existence of a diametric structure for all the pine forests in the form of inverted J (ordered: number of trees by hectare and abscisa: diametric classes).Without total diametric structure the system degenerates or tends to disappear (under natural conditions) Through temporary samples to level of all the mass. Diversity of associated species Determinant to maintain systems stability and health. Unknown in Central America and the Caribbean. In Florida there is> 20 species by square meter, mainly herbaceous. In all the pine forest the vital function of microorganisms associated to its roots is recognized (mycorrhiza). Systems susceptibility to the attack of plagues and/or illnesses. Continuous inventories through permanent plots.  Exotic species Displacement of native species, especially to a level of undergrowth. Favors fire incidence. Deteriorate the habitat.Investigations required.Loss of biodiversity. Increment of fire risks.Continuous inventories through permanent plots. Irrational timber exploitationAlter pine structure . Damage biodiversity. Provoke genetic erosion. Induces appearance of exotic species. Alters hydrologic regimes to a soil level.The causes and possible effects are known but not the quantification. Degraded pine trees in genetic and economic terms. Deterioration of habitats. High rates of surface water run-off and reduction of infiltration into soil. Continuous inventories through permanent plots and run-off plots.Grassing Damage biodiversity. Induces appearance of exotic species. Alters hydrologic regimes to a soil level. Alters fire regimes.The causes and possible effects are known but not the quantification. Deterioration of habitats. High rates of surface water run-off and reduction of infiltration into soil. Continuous inventories through permanent plots and run-off plots.Connectivity Loss of speciesSize (minimum dynamic area)Maintains systems stability and health.Depends of the origin and diversity of habitats. Large enough to maintain fire regime and genetic base. For example, if the frequency of the fire is 5 years, should not burn more than one fifth of the pines each year.Need of buffers to facilitate fire management.  Savanna Colombia and Venezuela, Landscape Context Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold) Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityPrecipitationClimatic evidence Precipitation: 800-1,600 (2,500) mm / year. Dry season: 6 months max. <800 mm: spiny vegetation >1,600mm: semi deciduous forest to evergreen forestsRainfall measure Specific geohydric regime Soil hydric regime and limit soil nutrients reserves 1. Texture: sandy clayey, the gradient can adapt the type of savanna (rank: consult literature). [Experts, please provide literature citation. Thanks.] 2. Hydro edafico regime: Flooded- non-flooded Apure savanna, classified in river bank, low-lying ground and marsh; 6 months of flood min. Other types: with less or non flood (High plains, oriental plains, Mesa de Guanipa, etc.); variables in their woody component: smooth savanna to wooded savanna.In more fertile soils (clayey) semi-deciduous to deciduous forests tend to be installed . In soils less fertile would be a very open and degraded savanna (e.g., Mesa de Guanipa)Soil composition measure Fire regime Maintenance of mosaic of the present vegetation of savanna and forest. Frequency of fire: 5 years to maintain the present equilibrium (little natural fire). If the fire is eliminated, probably the woody plants increase. Monitoring plots of the relationship of woody / non-woody plants.The great beta and range diversity: distributed upon an area of aprox. 300,000 Km2, extensive variety of vegetable communities; great heterogeneity paisajstica. An important and representative biome of the flora and fauna of north South America. Minimum dynamic area: depends on each type of savanna included in this extensive region. For some types less extended, as for example the plain palm groves of Copernicia tectorum, the minimum area can be of 1-2 km2, while for types of savanna more extensive (for example, oriental savannas of Monagas) they can be required not less than 100-200 km2. In the case of plains Morichales, typically riparian, the minimum area is measured in km of river and should cover all the river extension with this type of vegetation (among 200-300 km). Empirical assumptions. Remote sensors. Level of natural state observed in the field.Floristic composition of Plains (see Aristeguieta, 1966; Ramia 1974) dominated by gramineas (Trachypogon spicatus, Axonopus spp., Panicum spp., Andropogon spp. ) in the herbaceous stratum and by Curatella American, Byrsonima crassifolia and Bowdichia virgilioides in the woody stratum. Few endemic species; the same are mostly concentrated in the gallery forests. Floristic typical group of this biome in Venezuela and Colombia. The herbaceous species conform the 60-70% of the savanna Venezuelan flora . The woody species and the epiphytes and parasitic plants conform the rest. 40-60 species are considered as representatives of most of the Venezuelan savanna types. Indicator species and faithful to a basic floristic nucleus of Neotropical savannas (Huber, 1987).Floristic inventories. Savannas and herbaceous of Escudo Guayanes (azonal system), Condition Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold)Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityLevels of nutrients in the extremely low soil (oligotrofismo edfico extreme). It can be observed in the following three typical and exclusive of the Shield Guayans types of herbaceous ecosystems : Savanna graminous totally open upon soil of very low fertility derived from sandstone (oxisol and ultisol), Herbaceous oligotrophic, upon peat soil (histosol) (relictual vegetation of subtepuy). Herbaceous oligotrophic upon inundated sandy soil (quartzipsamment) Responsible for the presence of this type of savanna in the low and medium soils of the north and east of Escudo Guayanes (Huber, 2000), including the Great Savanna in Venezuela, the state Roraima of Brazil, Savanna Rupununi in Guyana, Sipaliwini Savanna in Surinam (aprox. 60,000 km2). Exclusive and only type of relict subtepuy vegetation in the world; extension of just 1000 km2 in the Great Savanna of Venezuela. Vegetation type exclusive from low and medium soils of the southwest and northwest sector of the Shield Guayanes (aprox. 8,000 km2 in the center of the Amazonas state in Venezuela and Dept. Guainia in southeast of Colombia). (Huber, 1982; Huber, 1995.) Rank of the level of nutrients in the soil should be extremely low. Adequate hydrologic conditions should be assured in the soil that maintain the organic matter in permanent state of saturation. The marked levels of oligotrophisim edaphic should prevail; these levels of soil nutrientes are not measurable with methods of traditional analyses. Obviously the factor nutrients reserve is the principal limiting factor in these ecosystems 1 to 3. With inferior fertility rates, extremely degraded savannas are observed until desertification fenomenous (for example, in Gran Sabana sudeste); with superior fertility rates, savanna forests similar to the savannas in Venezuelan plains or Brazilian cerrados are observed. In case of a prolonged hydric deficits a dry out of the peat is observed, this become inflammable and is frequently destroyed by fire. Seems to be ecosystems adapted to extremely edaphics and environmental conditions in effect in its distribution area. In general: periodic measure of the levels of soil nutrients. Specifically: floristic and ecological inventories for the observation of less evident changes. Floristic composition Highest continuos savannas in Venezuela (e.g., Uper elevation limit of Mauritia flexuosa 990 msnm). High rate of regional endemism. Too high rate of regional endemismo.20-30 species/ gramineas community Herbal subtepuyano should have a minimum of 10-20 spp. of tepuyana flora. The herbal of low soil should have a good representation of endemic species, adapted in their herbaceous and woody behavior to the extreme habitat conditions. Any change in the floristic composition means biologic degradation of the ecosystem. (specially in 2 and 3).Periodic floristic inventories.Intense fire regime High fire frequency (8,000 fires per year in Gran Sabana). 2 y 3. Occasional fires are causing notable impacts.  Apparently, none of the three ecosystems types 1, 2, and 3 are particularly adapted to survive the ecological effects of fire, showing in some cases evident signs of degradation caused by this factor. Therefore, it is estimated.The reduction of the fire frequency seems to be the only way to reduce negative impacts that are observed in these peculiar herbaceous ecosystems.Monitoring of the number of fires.  Cerrado, Landscape Context Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold)Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityFire regime Controls plant species distribution and balance between herbaceous and woody layers. Controls community composition. Interferes with nutrient cycling. Stimulates flowering, fruiting, and germination, Important for herbivore food availability and distribution. Fire frequency: Herbaceous campo limpo, campo sujo, and campo cerrado: well drained: 2- 5 yrs. badly drained: no fire. Cerrado (sensu stricto): 4-6 yrs. Cerrado and associated gallery forest: no fire. (Coutinho, 1990; Pivello & Norton 1996.) < 2 yrs. : Decrease species diversity and nutrient availability. >5yrs. : Accumulate dead organic matters, slow down nutrient cycling, change structure, and decrease herb diversity. Fires decrease species diversity and change composition and structure. <5yrs. : Reduce woody density and fire sensitive species; decreases nutrient availability. >10yrs. : reduce herbaceous layer and fire prone species. Favors Melinis minutiflora, and increases risks and intensity of wild fires. Fires decrease species diversity and change composition and structure; eliminate fire sensitive tree species.For monitoring changes at large scale: satellite imagery; field studies include: Burned stems or tussocks. Palms age distribution, dead standing individuals; presence or absence of organic matter layer. Burned stems and tree trunks; litter accumulation; density of dead standing woody plants. A normal stand should not have >5% of dead standing woody individuals >5 cm basal diameter (See Felfili et al., 1994, 1997 & 2000). Presence or absence of dead Xylopia and Copaifera trees, >5% of dead standing trees.3Drainage and watertable depth Affects plant community composition. Important for maintenance of seasonal lagoons for birds, amphibians and reptiles. Important for maintenance of pasture for mammals and herbivores during dry season.Herbaceous campos, well drained: moist only during the peak of the rainy season and >1 m watertable depth for the rest of the year. Herbaceous campo, badly drained: flooded 9-10 months and moist during dry season. Cerrado and Cerrado : > 5m watertable depth. Gallery forest : Seasonally flooded, flooded during the rainy season and moist the rest of the year. Well drained: moist during the rainy season and well drained during the rest of the year.Influence structure, floristic and faunal composition.Floristic structure and composition.2Climate: seasonal rainfall and frost frequency. Determines presence or absence of cerrado.5-7 months of dry season (May-Sept.) with sporadic frost events occurring in > 3 yrs.Water deficit prevents the establishment of moist forest. Frequent frost prevents cerrado establishment.Rainfall and frost records.1Soil structure and fertility Determines plant community types.Hydromorphic soils or shallow soils with hardpan: herbaceous campos badly drained and associated gallery forest. Shallow or very poor soils: herbaceous campos, well drained. Deep soils: woodland. Mesotrophic: cerrado. Dystrophic: cerrado or cerrado. Rocky soils (without hardpan): Shallow: high altitude cerrado. Rock outcrops on slopes: woodland cerrado. (Roots penetrate cracked rocks.)1. Humid 8-10 months of the year. 2. < 1m depth. (see Goodland & Pollard, 1973; Lopes & Cox, 1977; Haridasan and Ranzani ?? year for nutrient levels.) [Experts, please complete the citation of Haridasan & Ranzani in the References section. Thanks.] 3. 5 m depth. (see Haridasan ??year for nutrient levels.) [Experts, please complete the citation in the References section. Thanks.] Presence or absence of bare rocks or gravels.Soil Texture and color.3Connectivity by gallery forest networks (gallery forests serve stepping- stones-like connections for cerrado fragments.)Maintenance of genetic interchange among isolated populations.Index of connectivity : see Metzger and Dcamps, 1997. Loss of top predators and rare species.Presence of top predators (e.g.,Jaguar). Cerrado, Condition Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold) Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityViable and naturally regulated frugivore bats and birds communities and maned wolf.Help seed dispersal.Identify the minimum set of species that disperse all plant species in the ecosystem and the minimum population size for each species. [Experts, if you know of any published studies, please cite. Thanks.]Absence of some animal species may cause diversity erosion in plant communities.Density and diversity of birds, bats, and density of maned wolf.6Pollinators Help plant reproduction.Same as aboveSame as aboveSame as above5Herbivory (ants, termites, and other insects, Rhea, deer..etc.) Improves nutrient cycling. Prevents woody plant encroachment and fuel build up.Same as aboveSame as aboveSame as above4Cattle grazing Introduces exotic grasses and changes community composition; increase fire frequency. May spread diseases to native fauna.Presence or absence of cattle: the approximate carrying capacity in managed areas : 1 head/ 4 ha (This figure may vary depending on the floristic composition and landscape mosaics of the savanna).Compact soil. Grazing increases soil erosion. Reduce population size of plant species sensitive to grazing. Presencia de carcavas, caminos de pisoteo de ganados.Exotic species Competition and displacement of native species. Reduces biodiveristy.Presence or absence of Melinis, Brachiaria, Andropogon, Hyparrhenia, etc. One or more of these species become the dominant herbaceous species in terms of their high Index of Importance Value among herbaceous species.Partial or total replacement of native herbaceous plants.Calculate Index of Importance Value (See: Pivello et al., 1998.)Minimum dynamic area Maintenance of viability of all biotic elements.> 130.000 ha (Parque Nacional Emas)Maintenance of the landscape structure, function at all trophic levels.Areas needed to maintain viable population size of top predators.1 Plains of Moxos - Savannas of north Beni, Landscape Context Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold)Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityPrecipitationAffects flooding dynamics because of the rainfall in the region and occurrences of termite nests in flooding area. Minimum rainy season: 5-7 months. Flooding period: 3-5 months.The decrease of precipitation and the flooding period change the extension and composition of communities to more xerophytic.Measurement of rainfall regime. Relief and soil (structure: e.g., hard pan, and fertility) Affects flooding dynamic and distribution of the vegetables communities. Depth of hard pan close to the surface: <1 m.High plains with emergence of lateritic crusts and pisolites with campo cerrado vegetation. In ravine dominates the broadleaf evergreen forest partially flooded above lateritic hard pan and fine material.Mosaic of different types of vegetation from the ravine forest to open savannas with cperaceas dominating.Fire regime Affects the mineralization of organic matter and the distribution of vegetable communities. Fire period: <20 years to maintain the system. Change the systems to dryer communities. Domination of pyrophytic species : in the hillock: Trachypogon plumoso (=T. spicata). in the depressions: Paspalum, Leptocoryphium lanatum Plains of Moxos-Savanna of north Beni, Condition Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold)Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityTermite and nests in plains areas bad drained or old riverbeds forming bogs.Maintain the flow of nutrients. Presence of earth and termite nests: 10-40%.Are there marshy or aquatic vegetation found? It is a supposition not proved!Rate of the termite or earth nests coverage. Plains of Moxos sensu estricto (Savannas of south Beni), Landscape Context Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold) Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityPrecipitationDetermine the distribution of species. Minimum rain period: 5-7 months. Flooding periods because of river overflowing and local rains: 3-5 months, up to 8 months, relief dependent. Decrease of the flooding period changes the extension and composition of the communities to more xerophytic.Measurement of the rainfall regime. Fire regime Determine the composition and distribution of vegetable communities - aquatics (lagoons, deep streams), marshy (yomomos, curichis), herbaceal (low-lying ground, herbaceous, and woody (mid elevations to tajibales) (Tabebuia spp.) and forest island. Affects the flooding dynamic. Tusicales (Machaerium hirtum), type of mid elevation community, found only in soils with pH> 8. [I modified the sentence. Experts, please verify. Thanks.] Maintenance of other communities requires 3-9 months of flooding. If pH changes and the flooding regime decrease, the vegetable communities tend to be drier. Measurement of the rainfall regime, flooding and pH. Fire regime Affects the dynamic and composition of vegetation. Cyperus giganteus Marsh (Yomomo o Pirizal): fire period >50 years. The bajos de canuela (Luziola, Panicum, Echinochloa, Hymenachne): should not get burned (is source of foraging, important habitat to a marsh deer). Mid elevation with little organic matter, burned sporadically: <10 years. High elevations towards the Andes: burned each year. Ecosystem degradation, decrease of the Tabebuia insignis island and heron nests and caimans and lizards habitats. dem for marsh deer. In places without tusicales, fire destroys the little organic matter, becomes more xerophytic. If there is no fire for 2-5 years in high elevations, the vegetation changes to woody. Fire frequencyDynamic minimum area Maintains the viability of all biotic elements including gallery forests, lagoons and forests island. Larger flooding areas: >200,000 ha.Large flooding events: every 10 years.Presence of top predators.Salinity Responsible for the presence of tusicales (open and espinoso forest of Machaerium hirtum) that serves as refuge for animals that take salt.Salnity level: see Werner Hanagarth (1993). Size of tusicales: de 0.5 a 5 /10 ha.The community depends of salinity and high pH.Dominion of Machaerium hirtum. Information gaps and caveats Vania Pivello: Food sources and feeding behavior of cerrado mammals and birds are not well known. The fire ecology of cerrados needs more studies, though there have been several research projects on this subject. For a list of information gaps on cerrado ecology and management, see Pivello, V.R. and G. Norton, 1996. Firetool: an expert system for the use of prescribed fires in Brazilian savannas. J. Appl. Ecol. 33: 348-356. Jeanine Felfili: 1- Control of weedy species in conservation units. 2 Non-invasive herbaceous species for recovery of degraded areas. 3 Monitoring of vegetation changes in permanent plots. 4 -- biodiversity assessments. 5 Silviculture of native species. Recommended priorities for conservation-driven research agenda and next steps Vania Pivello: 1. Implement fire management in protected areas that harbor representative cerrado ecosystems. 2. Increase connectivity between cerrado ecosystems and forest patches by adding corridors, buffer zones, etc. Literature Cited Aristeguieta, L., 1966. Flrula de la Estacin Biolgica de Los Llanos. Bol. Soc. Venez. Cienc. Nat.,110: 228-307. Barbault, R. 1983. Reptiles in savanna ecosystems. In Bourlire, F. (ed.) Tropical savannas. Ecosystems of the world 13. Elsevier Scientific Publishing Company, New York.Pp. 325-336. Baruch, Z. 1996. Ecophysiological aspects of the invasion by African grasses and their impact on biodiversity and function of neotropical savannas. In Solbrig, O.T., E. Medina and J.F. Silva (eds.) Biodiversity and savanna ecosystem processes: a global perspective. Springer-Verlag, Berlin, Heidelberg. Pp.79-93. Baruch, Z., J.A. Belsky, L. Bulla, A.C. Franco, I. Garay, M. Haridasan, P. Lavelle, E. Medina, and G. Sarmiento. 1996. In Solbrig, O.T., E. Medina and J.F. Silva (eds.) Biodiversity and savanna ecosystem processes: a global perspective. Springer-Verlag, Berlin, Heidelberg. Pp.175-194. Bilbao, B., R. Braithwaite, C. Dall'Aglio, B. Dias, A. Moreira, P. Oliveira, J. F. Ribeiro, and P. Stott. 1996. Biodiversity, fire, and herbivory in tropical savannas. In Solbrig, O.T., E. Medina and J.F. Silva (eds.) Biodiversity and savanna ecosystem processes: a global perspective. Springer-Verlag, Berlin, Heidelberg. Pp.197-203. Braithwaite, R. 1996. Biodiversity and fire in the savanna landscape. In Solbrig, O.T., E. Medina and J.F. Silva (eds.) Biodiversity and savanna ecosystem processes: a global perspective. Springer-Verlag, Berlin, Heidelberg. Pp.121-139. Bulla, L. 1996. Relationships between biotic diversity. In Solbrig, O.T., E. Medina and J.F. Silva (eds.) Biodiversity and savanna ecosystem processes: a global perspective. Springer-Verlag, Berlin, Heidelberg. Pp.97-117. Coupland, R.T. 1992. Overview of South American grasslands. In Coupland, R.T. (ed.) Natural grasslands. Ecosystems of the world 8A. Elsevier Scientific Publishing Company, New York. Pp. 363-366. Coutinho, L. M. 1990. Fire in the ecology of the Brazilian cerrado. In: Goldamer, J.G. (ed.) Fire in the tropical biota. Springer Verlag. Pp. 63-105. Daly, D.C. and J.D. Mitchell. 2000. Lowland vegetation of Tropical South America. In D. L. Lentz (ed.). Imperfect balance: landscape transformations in the Precolumbian Americas. Columbia University Press, New York. Pp. 391-453. Denslow, J.S. 1996. Functional group diversity and responses to disturbance. In Orians, G.H., R. Dirzo, and J.H. Cushman (eds.) Biodiversity and ecosystem processes in tropical forests. Springer-Verlag, Berlin, Heidelberg, New York. Pp. 127-151. Felfili, J.M., A. V. Rezende, M. C.Silva Junior and M.A. Silva. 2000. Changes in the floristic composition of cerrado (sensu stricto) in Brazil over a nine-year period. Journal of Tropical Ecology 16:579-590. Felfili, J.M., M. C. Silva Jr., A. V. Rezende, P. E. Nogueira, B. W. T. Walter, M. A. Silva, J. I. Encinas. 1997. Comparao Florstica e Fitossociolgica do Cerrado nas Chapadas Pratinha e dos Veadeiros. P. 6-11. In: Leite, L. and C. H. Saito. Contribuio ao conhecimento ecolgico do cerrado. Editora Universidade de Braslia. Felfili, J.M., T. S. Filgueiras, M. Haridasan, M. C. Silva Jr., R. Mendona and A. V. Rezende. 1994. Projeto biogeografia do bioma cerrado: Vegetao e solos. Cadernos de geocincias do IBGE.12: 75-166. Frost, P.G.H. and F. Robertson. 1987. The ecological effects of fire in savannas. In Walker, B. H. (ed.). Determinants of tropical savannas. Pp 93-140. IUBS Monograph Series, No.3. IRL Press Limited, Oxford, UK. Fry, C. H. 1983. Birds in savanna ecosystems. In Bourlire, F. (ed.) Tropical savannas. Ecosystems of the world 13. Elsevier Scientific Publishing Company, New York. Pp. 337-357. Goodland, R. and R. Pollard. 1973. The Brazilian cerrado vegetation: a fertility gradient. J. Ecol. 61: 219-224. Greller, A. M. 2000. Vegetation in the floristic regions of North and Central America. In D. L. Lentz (ed.). Imperfect balance: landscape transformations in the Precolumbian Americas. Columbia University Press, New York. Pp. 39-87. Hanagarth,W. 1993. Acerca de la geoecologa de las sabanas del Beni en el Noreste de Bolivia. La Paz. Happold, D.C.D.1983. Rodents and lagomorphs. In Bourlire, F. (ed.) Tropical savannas. Ecosystems of the world 13. Elsevier Scientific Publishing Company, New York. Pp. 363-400. Haridasan ??year in p. 28 Haridasan and Ranzani for nutrient levels in p. 28 Huber, O. 1982. Significance of savanna vegetation in the Amazon Territory of Venezuela. In Prance, G.T. (ed.) Biological diversification in the Tropics. pp. 221-244. Columbia University Press, New York. Huber, O.1987. Neotropical savannas: their flora and vegetation. Trends in Ecology and Evolution (TREE) 2(3): 67-71. Huber, O. 1995. Vegetation. In Berry, P., B.K. Holst and K. Yatskievich (Eds.). 1995. Flora of the Venezuelan Guayana. Vol.I: Introduction. pp. 97-160. Missouri Botanical Garden. St. Louis & Timber Press, Portland, Oregon. Huber, O. and G. Febres (editores). 2000. Gua ecolgica de la gran Sabana. Troncal 10: Piedra de la Virgen Santa Elena de Uairn. The Nature Conservancy EcoGraph, Caracas. 192 pp. Lewinsohn, T.M. and P.W. Price. 1996. Diversity of herbivorous insects. In Solbrig, O.T., E. Medina and J.F. Silva (eds.) Biodiversity and savanna ecosystem processes: a global perspective. Springer-Verlag, Berlin, Heidelberg. Pp.143-157. Lopes, S. and F. R. Cox. 1977. Cerrado vegetation in Brazil: an edaphic gradient. Agronomy Journal 69: 828-831. Metzger, J.P. and H. Dcamps. 1997. The structural connectivity threshold: an hypothesis in conservation biology at the landscape scale. Acta Oecologica 18:1-12. Mistry, J. 1998. Corticolous lichens as potential bioindicators of fire history: A study in the cerrado of the Distrito Federal, central Brazil. Journal of Biogeography 25 (3): 409-441. Ojasti, J. 1983. Ungulates and large rodents of South America. In Bourlire, F. (ed.) Tropical savannas. Ecosystems of the world 13. Elsevier Scientific Publishing Company, New York. Pp. 427-439. Oliveira-Filho, A.T. and J. A. Ratter. 1995. A study of the origin of central Brazilian forests by the analysis of plant species distribution patterns. Edinburgh Journal of Botany. v. 52, p. 141-194. 1995. Pivello, V.R. & Norton, GA. 199. FIRETOOL: an expert system for the use of prescribed fires in Brazilian savannas. J. Appl. Ecol. 33: 348-356. Pivello, V.R.et al. 1998. Abundance and distribution of native and alien grasses in a cerrado (Brazilian savanna) biological reserve. Biotropica, 31: 71-82 Ramia, M. 1974. Plantas de las Sabanas Llaneras. Monte Avla Editores, Caracas, 287 pp. Ramos-Neto, M.B. & Pivello, V.R. 2000 Lightning fires in a Brazilian savanna National Park: re-thinking management strategies. Environ. Manage. 26: 675-684. Rodrguez, J.P. and F.Rojas-Surez. 1999. Libro rojo de la fauna Venezolana (2da. Edicin). PROVITA, Caracas, Venezuela. Sarmiento, G. 1983. The savannas of tropical America. In Bourlire, F. (ed.) Tropical savannas. Ecosystems of the world 13. Elsevier Scientific Publishing Company, New York. Pp. 245-288. Silva, J.F. 1996. Biodiversity and stability in tropical savannas. In Solbrig, O.T., E. Medina and J.F. Silva (eds.) Biodiversity and savanna ecosystem processes: a global perspective. Springer-Verlag, Berlin, Heidelberg. Pp.161-171. Solbrig, O.T. 1996. The diversity of the savanna ecosystem. In Solbrig, O.T., E. Medina and J.F. Silva (eds.) Biodiversity and savanna ecosystem processes: a global perspective. Springer-Verlag, Berlin, Heidelberg. Pp.1-27. Solbrig, O.T., E. Medina, and J.F. Silva 1996. Determinants of tropical savannas. In Solbrig, O.T., E. Medina and J.F. Silva (eds.) Biodiversity and savanna ecosystem processes: a global perspective. Springer-Verlag, Berlin, Heidelberg. Pp. 31-39. Tania Sanaioti, INPA Recommended resources Beard, J.S. 1953. The savanna vegetation of northern tropical America. Ecological Monographs 23(2): 149215. Beck, S.G., J. Samiento, N. Paniagua Z., C. Miranda and M.O. Ribera. 2000. Humedales de Bolivia, una aproximacin a su conocimiento actual. Pp.119-150. In Homenaje al Acadmico Correspondiente Dr.h.c. C.N. Troels Myndel Pedersen. Academia Nacional de Agronomia y Veterinaria. Tomo LIV. Buenos Aires, Argentina. Berry, P., B.K. Holst and K. Yatskievich (Eds.). 1995. Flora of the Venezuelan Guayana. Vol. I: Introduction. Missouri Botanical Garden. St. Louis & Timber Press, Portland, Oregon. Coutinho, L. M. 1990. Fire in the ecology of the Brazilian cerrado. In: Goldammer, J.G. (ed.) Fire in the tropical biota. Springer Verlag. Pp. 63-105. Felfili, J. M. and M. C. SILVA JR. 1993. A comparative study of cerrado (sensu stricto) vegetation in central Brazil. Journal of Tropical Ecology 9(3): 277-289. 1993. Eiten, G. 1982. Brazilian savannas. In: Huntley, B.J. and B. H. Walker (eds.) Ecology of Tropical Savannas. Springer-Verlag. Pp.? Felfili, J.M. 1995. Diversity, structure and dynamics of a gallery forest in central Brazil. Vegetatio 117:1-15. 1995. Felfili, J.M., J. F. Ribeiro, C. W. Fagg, and J. W. B. Machado. 2000. Recuperao de matas de galeria. EMBRAPAS-CERRADOS. Planaltina. Doc. 21: 1-45. Felfili, J.M., A. V. Rezende, M. C.Silva Junior and M.A. Silva. 2000. Changes in the floristic composition of cerrado (sensu stricto) in Brazil over a nine-year period. Journal of Tropical Ecology 16:579-590. Filgueiras, T. S., J. M. Felfili, M. C. Silva Junior and P.E. Nogueira. 1998. Floristic and structural comparison of cerrado (sensu stricto) vegetation in central Brasil. Pp. 633-647. In: Dallmeyer, F. (ed.) Measuring and monitoring forest biological diversity. Ed. Smithsonian Foundation/MAB. The Parthenon publishing. New York. Furley, P.A., J. Proctor, and J. A. Ratter (eds.) 1992. Nature and dynamics of forest-savanna boundaries. Chapmen & Hall, London. 614 pages. Hanagarth, W. and S.G. Beck.1996. Biogeographie der Beni-Savannen (Bolivien). Geographische Rundschau 48 (11): 662-668. Braunschweig. Huber, O. 1974. The neotropical savannas. Select bibliography on their plant ecology and phytogeography. Istituto Italo-Latino Americano, Roma (Italia). xlii + 855 pp. Huber, O. 1999. Manejo integral de sabanas tropicales [Propuesta tcnica]. En Jornadas sobre desarrollo sostenible del medio rural: en busca del mejor camino (C. Pardo, coord. gen.), pp. 119120. Fundacin Polar, Caracas. Huber, O. 1990. Savannas and related vegetation types of the Guayana Shield region in Venezuela. In Sarmiento, G. (ed.) Las Sabanas Americanas: Aspectos de su biogeografa, ecologa y utilizacin. pp. 5797. Universidad de los Andes, Mrida, Venezuela. Huber, O.1987. Neotropical savannas: their flora and vegetation. Trends in Ecology and Evolution (TREE) 2(3): 67-71. Mendona, R., J.M. Felfili, B. M. T. Walter, M. C. Silva Jnior, A.V. Rezende, T. S. Filgueiras, and P.E.N. Nogueira. 1998. Flora vascular do bioma Cerrado. pp. 287-556. In: Sano, S. and S. Almeida. Cerrado, Ambiente e Flora. EMBRAPA CERRADOS. Ed. Planaltina, EMBRAPA-CPAC. Medina, E. and O. Huber 1992 [2ed ed. 1994]. The role of biodiversity in the functioning of savanna ecosystems. In Solbrig, O.T., H.M. van Emden and P.G.W.J. van Oordt (eds.). Biodiversity and global change. IUBS Monograph 8: 139158. International Union of Biological Sciences, Paris. Pinto, M.N. 1990 Cerrado: Caraterizao, ocupao e perspectivas. Braslia: Editora Universidade de Braslia, +657pp. Pivello, V.R. and L.. M. Coutinho. 1996. A qualitative successional model to assist in the management of Brazilian cerrados. Forest Ecology and Management 87: 127-138. Pivello, V.R. and G. A. Norton. 1996. Firetool: an expert system for the use of prescribed fires in Brazilian savannas. Journal of Applied Ecology 33: 348-356. Ratter, J. A ., S. Bridgewater, R. Atinkson and J. F. Ribeiro. 1996. Analysis of the floristic composition of the Brazilian vegetation of 98 areas. Edinburgh Journal of Botany 58(2):153-180. Ratter, J. A., J. F. Ribeiro, and S. Bridgewater. 1997. The Brazilian cerrado vegetation and threats to its biodiversity. Annals of Botany London 80(3): 223-230. Redford, K.H. and Fonseca, G. A. B. 1986. The role of gallery forests in the zoogeography of the cerrados non-volant mammalian fauna. Biotropica 18 (2): 126-135. Sano, S. and S. Almeida. 1998. Cerrado, Ambiente e Flora. EMBRAPA CERRADOS. Ed. Planaltina, EMBRAPA-CPAC. Sarmiento, G. 1984. The ecology of neotropical savannas (transl. By O. Solbrig). Harvard University Press, Cambridge, Mass. xii + 235 pp. 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). 4.8. Hot xeric systems General description and geographic variation Deserts are found in areas influenced by subtropical anticyclones, in rain shadows created by mountains or in continental interiors far from the ocean (Shmida, 1985). They develop in areas south of tropical latitudes and extend into temperate latitudes to the north. In America, they occur predominantly on the Pacific side of the continent. The hot xeric system includes the following deserts: Chihuahuan, Sonoran, Chilean-Peruvian, Baja Californian, Argentinean Monte, and coast of Venezuela and Colombia. The Mediterranean Shrublands and Patagonia are excluded. The team has no information on the areas of Tehuacan (Mexico) and Argentinean Monte. All deserts have extreme temperatures. Hot deserts differ from cold ones in their average annual temperatures. Deserts generally have low and highly seasonal precipitation, with great interannual variation. The extremes can vary from one to 300 mm (0.0412 inches) a year. Vegetation cover by annual plants varies due to large quantitative and seasonal rain fluctuations. During the dry season the landscape is barren. A large portion of desert vegetation consists of annual plants. One can also find microphyllous shrubs, small succulent trees, plants in rosettes (such as agaves and terrestrial bromeliads) or perennial and semi-deciduous shrubs. Deserts have high levels of endemism at the species and genus level. Insects are very important in desert environments, especially pollinators such as bees. Underground diversity consists of seed banks, termites (Sonoran and Chihuahuan) and micorrhizae. Flowering and faunal reproduction processes are adapted to rainy seasons. Ants and rodents are fundamentally important species (Jim Brown). [Experts, please cite Browns publication in the Literature Cited section. Thanks.] Community types/zonation and major gradients within the system (patterns) Based on seasonal rains, deserts are classified into three types: winter rain deserts, summer rain deserts and winter and summer rain deserts. Winter rains: Baja California, Chile (310 250S latitude, along the coast at 190S). Summer rains: Chihuahuan and Argentinean Monte, north of Atacama above 250S latitude. Winter and summer rains: Sonoran Desert. Based on the source of water, another category, fog desert, can be found along coastal zones of Peru and Chile. Annual precipitation here is close to zero and fog is the main source of water. Habitat diversity in the hot xeric system is spatially very heterogeneous and patchy. It can have dunes, xeroriparian systems, salt lagoons, temporary water bodies, coastal shrublands, bajadas formed by soil gradients, and springs. Ecological integrity factors for landscape context Table xxx. Ecological integrity factors for landscape context of hot xeric systems Key FactorsJustification for factor selection Ecological Threshold: (Minimum Integrity Threshold)Justification for determining the threshold (e.g., Natural Range of Variation)Field-based monitoring indicatorsFactor PriorityPrecipitation regimes. Their variability and patchy patterns determine beta diversity as well as habitats and resources within patches. They determine productivity and vegetation cover.Related to a climatic change and it must be evaluated at a global or regional scale.Variability is good in the desert. High variability in the desert on an annual basis over time (years of drought) is acceptable and should not be altered.Local precipitation patterns. Long-term regional precipitation patterns. Evaluation of changes with relation to global phenomena (e.g. El Nio).Size gradient of soil particles (Bajadas). Bajadas have very important and predictable vegetation gradients that maintain particular species on each of the bands of the gradient. Diversity decreases from the highest parts towards the valleys.It is necessary to maintain connectivity between high parts and the valleys.Fragmentation of bajadas (agricultural fields, roads, mines, etc.). Monitoring indicator species, such as saguaros, which depend on the structure of bajadas.Nitrogen and phosphorous availability.Fire regime.In general, fire is not a natural factor in the desert and its presence can have unpredictable outcomes. Nevertheless, in the Chihuahuan desert, which functions as a grassland with 3-5 year fire regimes, fire suppression favors woody plants and changes the vegetation from savanna to shrubland. Exotic species presence increases the frequency and intensity of desert fires.A dense cover of exotic species (such as buffelgrass Cenchrus ciliarisin the Sonoran desert) surpasses the deserts natural fire regime threshold.Sources generating recurrent fires do not exist in deserts with winter rains (such as the Chilean and Baja Californian). In deserts like the Sonoran and the summer rain deserts of Chile, the source of fires exists (lightening) but there is not enough vegetation cover for them to spread. In the Chihuahuan desert, where both vegetation cover and sources (summer rain lightening) necessary to initiate and propagate fires exist, desert grasslands are maintained by fire regimes.Monitoring exotic plant cover. Fire frequency. Monitoring vegetation composition and structure in burnt areas and associated or key fauna. See Alberto Brquez, UNAM (Hermosillo Campus, Sonora, Mexico). [Experts, please complete the citation in the References section. Thanks.] Underground water regimes.Water availability for plants with deep roots. They maintain springs and water bodies that depend on springs and underground water (cinagas). Water bodies are important to native and endemic aquatic plant and animal species.The lowering of the water table below the reach of the most susceptible roots that feed from underground water. The lowering of the water table below the level needed to feed the springs and other bodies of water dependent on springs.Note: knowledge about plant dependence on underground water is very limited. In terms of springs there is no range, only the presence or absence of the spring.Monitoring the water table in wells. Monitoring the effect of water table changes on plants and bodies of water that depend on underground water.Surface water regimes.Maintaining the integrity of xeroriparian habitats is highly dependent on surface water flow regimes. These xeroriparian habitats are corridors for birds and other animals within the desert. Flash floods create vegetation patches in flooded areas.The minimum acceptable threshold for water regimes is highly dependent on the systems vegetation cover. In other words, the intensity of the impact from any change in surface water regimes depends on the systems vegetation cover. An indication that a minimum acceptable threshold has been exceeded is an increase in trenches (carcavas) that are not restorable, in comparison with similar areas considered in good condition.Monitoring volume and periodicity of surface water flows, if there are conservation targets that depend on these factors (fishes, amphibians). Monitoring vegetation cover, intensity of overgrazing and erosion.Invasion by exotics (local hydrology).See Alberto Brquez. UNAM (Hermosillo Campus, Sonora, Mexico).Connectivity to adjacent dry forest and mountain systems: water and altitudinal gradient. Ecological integrity factors for condition Table xxx. Ecological integrity factors for condition of hot xeric systems Key FactorsJustification for factor selection Ecological Threshold: (Minimum Integrity Threshold)Justification for determining the threshold (e.g., Natural Range of Variation)Field-based monitoring indicatorsFactor PriorityPollination: especially by bees, bats, and humming-birds. The majority of woody and succulent plants are dioic and/or self-incompatible, flowering seasonally and in a patchy spatial pattern, and therefore depend on pollinators for sexual reproduction. Bees are a particularly important pollinator group in the deserts. Bee emergence is highly dependent on precipitation cycles. Bats and hummingbirds depend on flowering to complete their migratory routes. The loss of plant continuity necessary for some migrant pollinators could cause pollinator populations to collapse. There is an information gap with respect to the relation between plants and specific groups of pollinators.Some bees can survive for 8 or 9 years underground during droughts, so that their absence in a given time does not necessarily mean a reduction of their populations. The effective reproductive size of plants tends to be larger than the real population size, given that it is common that not all individuals flower each year. In some years there is no synchronization between flowering and the pollination process. Due to the intrinsic variability of the system, it is difficult to determine if a reduction of pollinator populations and fruiting levels is outside of the natural range of variation. Monitoring fructification of indicator plants. [Experts, please provide a few examples of indicators or cite references. Thanks.] Monitoring key groups of large bees. [Experts, please provide a few examples of the key groups of large bees? Thanks.] Monitoring bats. Monitoring flowering phenology.Seedling establish-ment. In order to establish seedlings, some plant species depend on the confluence of several factors that are, by nature, variable, e.g., precipitation amount and pattern, temperature, pests, nurse plants, and seed bank conditions. Seedling establishment is the most critical factor to maintain plant composition in desertsSmall changes in climate can have enormous effects on community structure and composition.Not all the factors required for seedling establishment in deserts are known. Establishment events are naturally very sporadic.Monitoring species representing different life forms (e.g., plants with seed banks vs. plants without them). Information gap: we need to know what the factors are for establishment of key desert species.Hervibory on seedlings and seed predation. Hervibory is a very important factor for seedlings and seeds, since these are the most vulnerable stages of plant life cycles. Seed predation is part of the process of species dispersal. Rodent and ant guilds are the most important groups for seed predation.The disequilibrium in the number of seed predators (ants and rodents) will produce a similar disequilibrium in the dispersal and growth processes of new seedlings.Variability in deserts is very high, which makes it difficult to differentiate factors that are within their natural range of variation from those that are not.See the work of Jim Brown. [Experts, please cite Jims publication in the References section. Thanks.] Search for information and permanent plots of IBCD [Experts, please provide the full name of IBCD. Thanks.] established in the 70s. [Experts, please provide literature citation. Thanks.]Maintenance of seed banks (guilds that use the soil: burrowing rodents, ants, termites, beetles, etc.) Many desert plants depend on seed banks, particularly during drought years. These seeds can survive many years until appropriate conditions occur. Rodents and ants are keystone species very important for maintaining the seed-bank dynamics and, consequently, the vegetation structure.It is very important that the vertical structure of soils be maintained, because if it is lost, some of the seeds could remain on the surface and be lost or germinate all at once, causing the loss of the seed bank; or they could end up too deep in the ground, where they will not germinate. Variability in deserts is very high, which makes it difficult to differentiate factors that are within their natural range of variation from those that are not.Monitoring soil use changes. Monitoring rodents and ants. See Jim Brown. See Julio Gutirrez (seed bank in Chile). [Experts, please cite Jims and Julios publications in the References section. Thanks.] Vegetation structure: nurse plantsMany of the succulent plants depend on nurse plants for survival, because these create a favorable microclimate for them (retaining humidity, protecting against freezing and hervibory).Any factor affecting shrub cover (logging, overgrazing) has a greater effect, because it reduces the capacity of shrubs to serve as nurse plants to other plants.Species acting as nurse plants are resistant to natural variations in the desert with the result that ranges of variation are more influenced by human-related factors.Monitoring vegetation cover changes: See Susan Anderson and Peter Warren. [Experts, please cite the publication by Anderson and Warren in the References section. Thanks.]Seed dispersal.The group lacks the experience to address this factor.External migration patterns: birds, bats, and butterflies.Many of the migration patterns of birds are known. Birds use deserts as part of their migration routes and also have specific patterns within the desert. Little is known in the case of bats and butterflies, but new information is being generated.Each migrant group uses particular plant species. Thus, it is important to maintain connectivity over large desert areas (e.g. hummingbirds use Fouquieria, bats use Agave and columnar cacti, and butterflies use flowering plants including herbs). In the case of butterflies, migration patterns involve different seasons of the year. Rodrigo Medelln and Arnulfo Moreno (bats in Mexico). Ted Fleming (bats and saguaro) Bill Calder (Hummingbirds and Fouquieria). [Experts, please complete the citation in the References section. Thanks.]Internal migration patterns related to patchy resources habitat availabilityCryptogamic crusts.They are important for maintaining soil fertility and for avoiding soil loss through erosion. Are they nitrogen fixers? This discussion group lacks information on this subject.Restoration of these crusts is very difficult after disturbance has occurred.Guano deposits on coastal deserts (sea birds) Studies could show their function within deserts.Exotic animals: cows, goats, horses, donkeys, fishes, snails and lagomorphs (South American deserts)Cows, goats (very damaging because they also eat roots), horses and donkeys change the structure of vegetation and soils through trampling, favoring processes of erosion. Fishes and snails predate on native species or compete with them for resources, affecting the structure and composition of native communities. Some studies have shown that without grazing, deserts such as the Sonoran could recover. But in deserts with low precipitation (relative to other deserts) restoration could be more difficult. Studies in Africa have shown that overgrazing contributes to desertification; in general, many systems are not restorable after overgrazing. Dune systems are particularly susceptible. The same happens with native fish communities.There are information gaps and one cannot generalize from one desert to another due to ranges of variation among different deserts. See Susan Anderson and Peter Warren (effect of overgrazing on vegetation) Se Campo Experimental La Campana (Chihuahua, Mexico). ). [Experts, please complete the citation in the References section. Thanks.] NOTES: There is an emerging problem related to the invasive potential of exotics, particularly Cactoplastis (a coleopterus insect used for biological control of Opuntia in Australia). It is apparently moving rapidly from the U.S. towards Mexico and a possible catastrophe is predicted for Opuntia. Literature Cited Anderson, S. y P. Warren. ?? year?? (monitoring changes on vegetation cover). On page 4. Anderson, S y P. Warren (effect of overgrazing on vegetation). On p.5. Brown, Jim. ? Year? On p. 4. Brown, Jim. ?Year. On p. 4. Brquez, Alberto. UNAM (Unidad Hermosillo, Sonora, Mexico). On p. 7and p. 8. Calder, Bill (hummingbird and Fouquieria). On p. 4. Campo experimental La Campana (Chihuahua Mexico). On p. 6 Fleming, Ted (bats and saguaro). On p. 4. Gutirrez, Julio (seed bank in Chile). On p. 4. IBCD: information and permanent plots from IBCD established during the 70s. On p.4. Medelln, Rodrigo and Arnulfo Moreno (bats in Mexico). On p.5. Shmida, A. 1985. Biogeography of the desert flora. In: Evenari, M., I. Noy-Meir, and D.W. Goodall. (eds.) 1985 Ecosystems of the world 12A: Hot deserts and arid shrublands. Pp. 23-77. Elsevier Science Publishers, Amsterdam. Recommended resources 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. Evenari, M., I. Noy-Meir, and D.W. Goodall. (eds.) 1985. Hot deserts and arid shrublands. Ecosystems of the world 12A. Elsevier Science Publishers, Amsterdam. Gutirrez, J.R., P.L. Meserve, J.M. Jaksic, L.C. Contreras, S. Herrera and H. Vquez. 1993. Structure and dynamics of vegetation in a Chilean semi-arid thornscrub community. Acta Oecologia 14: 271-285. Gutirrez, J. R., P. L. Meserve, L.C. Contreras. H. Vsquez and F. M. Jaksic. 1993. Spatial distribution of soil nutrients and ephemeral plants underneath and outside the canopy of Porlieria chilensis shrubs (Zygophyllaceae) in arid coastal Chile. Oecologia 95: 347-352. Medelln- Leal F. 1982. The Chihuahuan desert. In: G. V. Benden (eds.) Reference handbook on the deserts of North America. Greenwood Press. Wesport, Connecticut. London, England. Van der Maarel (ed.). 1993. Dry coastal ecosystems: Africa, America, Asia and Oceania. Ecosystems of the World 2B. Elsevier Science Publishers, Amsterdam. Van der Maarel (ed.). 1997. Dry coastal ecosystems: General aspects. Ecosystems of the World 2C. Elsevier Science Publishers, Amsterdam. 4.9. Mangroves General description and geographic variation The text that follows was adapted from the pre-workshop document of Yara Schaeffer-Novelli, with additional information contributed by Aaron Ellison and Elizabeth Farnsworth. Mangrove ecosystems, also referred to as Mangal, are characterized by woody, tropical halophytes that are obligate to the system. The quintessential mangrove system is composed of trees with prop roots that spend much of their time inundated by tides. However, there are frequent physiognomic variations that depend on local biotic and edaphic factors and gradients in which the mangroves are found. Mangrove ecosystems are critical nursery grounds, last refuges for a number of large mammals of lowland forest, and refugia from predation for a number of commercially important fish and invertebrates (spiny lobsters). The mangrove ecosystem is found panglobally. There is substantial continental and regional variation in species richness within mangal; in the Neotropics richness is highest on the Pacific coasts of Colombia, Panama and Costa Rica and declines with latitude (Duke et al. 1998; Ellison and Farnsworth, 2000). Caribbean mangal hosts the worlds richest mangrove associated invertebrate fauna, and provides habitat for a number of globally endangered species. The coast of the Yucatan Peninsula and Belize, with abundant rainfall, occasional hurricanes, and a <1m tidal amplitude, has a particular high invertebrate and tree diversity. Ellison and Farnsworth (1996) provide critical data on the extent of mangrove forests in the Caribbean. In Central America, Mangroves provide habitat for threatened epiphytes, including the orchids Brassavola nodosa and Schomburgkia tibicinis (see Murren and Ellison, 1996; Rico-Gray et al., 1989) and several bromeliads (see Gomez and Winkler, 1991.) I dont know if these species are globally endangered, but theyre all threatened, and on CITES restricted lists. Manatees depend on mangroves and adjacent seagrass beds for habitat as well. Other species to check on include the mangrove warbler, American crocodile, and caiman. For a list of rare species supported by mangal around the world, please see Appendix, pp. 30 & 31. New World mangroves attain their greatest structural development where rainfall and tidal subsidies are large such as near equatorial areas where intense convective activities of the Intertropical Convergence Zone occur, and mesotidal (tidal range 2 to 4 m) or macrotidal (tidal range > 4 m) regimes dominates. Conditions associated with high annual rainfall (> 2,000 mm/yr) and large tidal amplitudes (> 2 m) occur on the northwest seaboard of the South American continent along the Pacific coast of Colombia, Ecuador and northern Peru, and on the eastern seaboard of the continent, from south of the Gulf of Paria (Venezuela) to So Lus (Brazil). It is within this moist and dynamic region where the greatest mangrove development occurs (Schaeffer-Novelli & Cintrn-Molero et al., 1990). This belt is roughly restricted to within 10 degrees of the equator, except on the Pacific coast of South America. There the Peruvian collision-type coastal margin is dominated by high mountains and small watersheds; and is characterized by reduced delivery of sediments to a narrow coastal plain (Schaeffer-Novelli & Cintrn-Molero, 1993). Atmospheric stability arising from the subsidence of the easterly winds flowing down the Andes and the westerly winds cooled by the cold Humboldt Current suppresses convective activity and creates extremely xeric conditions close to the coast (Linacre & Geerts, 1997). Thus, the southernmost mangrove stands at the western coast of South America barely reach 05o30' S, at the mouth of the Piura River (Pea & Vasquez, 1985). Along the Brazilian coast mangroves occur from the border with French Guiana (04o30' N) to points well beyond the Tropic of Capricorn, reaching 28o30'S, near Laguna (Santa Catarina, Brazil), where they eventually become limited by low temperatures and sporadic frost events (Cintrn-Molero & Schaeffer-Novelli, 1981&1983). Community types/zonation and major gradients within the system (patterns) Mangrove ecosystems exhibit distinct zonation, with certain tree species within mangrove systems occurring along tidal gradients, likely attributable to species-specific differences in tolerance to edaphic factors/physical gradients that vary with tidal height; sorting of propagules; interspecific competition; and seed predation (Ellison and Farnsworth, 2000). The exact ecological processes shaping this zonation are not conclusive. There are about nine species of mangroves in the Neotropics, Rhizophora mangle, Avicennia germinans, Laguncularia racemosa, and Conocarpus erectus, which are common and widespread, and A. schaueriana, A. bicolor, R. racemosa, R. harrisonii, and Pelliciera rhizophorae, which have more restricted distribution. Avicennia schaueriana occurs in the Caribbean Islands, A. bicolor on the Pacific coast of Central America. Rhizophora racemosa and R. harrisonii (= R. mangle x R. racemosa) are found on the Pacific coast of Central and South America. Pelliciera rhizophorae occurs sporadically in Nicaragua, Costa Rica, Panama, Colombia and Ecuador. It has been included in The 1997 IUCN Red List of Threatened Plants (Walter and Gillett, 1998). Pelliciera rhizophorae was formerly widespread throughout the Neotropics, but climatic changes during the Holocene have dramatically reduced its range (Graham, 1995; Ellison, et al., 1999; Tomlinson, 1995.). The physiognomy and functional characteristics of mangrove stands reflect the response of the component species to local ecophysiological factors. Lugo & Snedaker (1974) recognize six physiographic types of mangrove stands: fringe mangroves, basin mangroves, riverine mangroves, overwash mangroves, scrub mangroves, and hammocks. Each of these types is influenced by sets of similar energy signatures so that within each forest type, similar levels of structural development are attained. Cintrn-Molero et al. (1985) suggest that this classification could be reduced to three broad types - fringes, riverine and basins. Overwash mangroves are basically self-enveloping fringes, whereas scrub and hammocks are considered special types of basins. The classification of mangroves by Lugo & Snedaker has been largely superceded by one developed by Twilley (1995). Fellers work illustrates the importance of nutrient limitation on mangrove structure, especially contrasting scrub, fringe, and basin, and suggests that all is not geomorphology and flooding (Feller, 1995). Hogarth (1999) recognizes the following types of mangrove systems. Fringe Mangroves (tide-dominated): characterized by a high tidal range over a shallow intertidal zone that is often colonized by mangrove trees. Tidal water is typically full strength sea water, but wave action is diffused quickly by passage over a stepped intertidal zone. Sediment and mangrove soils are likely to be more dynamic as tides deposit and remove sediments from the sea and from inland river estuaries. Receive less runoff of terrestrial nutrients compared to riverine forests. Basin Mangroves: On the landward side of fringing mangroves in estuaries. Sheltered from wave action, and inundated infrequently. Highly variable salinity depending on rainfall, groundwater flow, and local tidal surges. Often exhibit high evaporation rates, which can result in hypersaline soils. Due to low currents and little turbulence, basin mangroves can be sinks for nutrients and sediment. Riverine Mangroves: Many large expanses of mangroves are located at river deltas where soils and salinity are amenable to mangrove community development (e.g. Amazon delta). Have low tidal ranges, and strong freshwater flow carrying substantial sediment loads, much of which is deposited within the mangrove communities. Characterized by shifting river channels, and typically mangal expanding inland as well as outward in the shifting, sediment-driven river deltas. Scrub Mangroves: Found in extreme environments where nutrients and freshwater may be limiting. Hammock Mangroves: Relative isolation from rivers or the sea leads to a domed accumulation of organic peat over depressions, where mangroves take root. Carbonate Setting Mangroves: On low-energy coasts where carbonate has accumulated from coral reef breakdown, resulting in lime sediment and silt accumulation. Inland Mangroves: Areas where the mangroves are completely cut-off from the sea, often in sink holes or other depressions. Ecological integrity factors for landscape context Mangroves are naturally disjunct habitats occurring along coastlines and rivers. They tend to have linear or small patch distrbutions. Though mangrove development encompasses both riverine and coastal types in the coastal plain of Belize and eastern Central America, mangroves do not generally occur as large matrix habitats. Mangrove species disperse well with waterborne propagules (often viviparous). They tend therefore not to be as sensitive to habitat fragmentation as many other coastal forest habitats, as long as major ecological processes are intact. However, fringing mangroves in particular may be affected by rising sea level, as their landscape context is extremely limited and linear. These types of mangrove systems will be limited by geology, and by human fragmentation, with in situ loss of the remaining fringing systems from physiological effects of rising sea levels (Ellison & Farnsworth, 1997). Table xxx. Ecological integrity factors for landscape context of mangroves Key FactorJustification for SelectionEcological ThresholdsJustification (Natural Range of Variation)IndicatorsFactor Relative Priority Wave/Tidal ExposureLeeward sites more speciose in mangrove root epibiont communities (Farnsworth & Ellison, 1996).Massive sponges, colonial tunicates indicate stable salinity, low tidal action and amplitudes.Tidal CyclesImportant for larval dispersal of mangrove invertebrate and fish communities, Contribute to the dispersal of mangrove seedlings and the zonation patterns of mangrove communities within the system. Affect soil and water salinity ranges.Sea LevelSea level rise from climate change will result in some inland migration that will be highly constrained by geology, and human impediments, and will likely cause significant die-off (Ellison & Farnsworth, 1996b). Simulated experimental rises in sea-level resulted in decreased photosynthesis, growth, and productivity of mangrove tree species (Ellison & Farnsworth, 1997). Throughout the Holocene, mangroves have responded to small changes in sea level < 8-9 mm/yr through landward and seaward migration, mediated by local topography (Ellison & Farnsworth, 1997). Now the migration is no longer possible with dense human settlement along coasts.Freshwater DischargeThough halophytes, mangroves need freshwater input.15 ppt seawater seems to be about the threshold in the neotropics. Benthic epifauna drop out at <35 ppt; freshwater lenses resulting from strong rainfall will kill epibionts near the surface.SedimentationGrowth of mangrove saplings on coral cays declines significantly with sedimentation rate (Ellison & Farnsworth, 1996a). The structure of mangrove roots leads to increased sedimentation and settling of suspended silt. Some 80% of suspended sediment brought in from coastal waters is trapped in mangroves naturally (Hogarth, 1999). The muds that result from the process are critical not only as substrate to mangrove trees and invertebrate communities, but are the site of much nutrient cycling in the system, and unicellular algae and cyanobacteria within these muds exhibit high photosynthetic rates.Accretes in several mangrove systems between 3-6 mm/yr (Ellison & Farnsworth, 1996a), with maximum reported of 8 mm/yr (Hogarth, 1999). Sedimentation rate correlates with pneumatophore density in many mangrove forests. Some 80% of suspended sediment brought in from coastal waters is trapped in mangroves naturally (Hogarth, 1999).High rates of sedimentation associated with urban development cause loss of ascidians and alter epibenthic community structure (Ellison & Farnsworth, 1996b).Hydrographic ProcessesDeemed critical for other successional processes to occur, and affects other landscape context key factors such as sedimentation, freshwater discharge, seedling disperal, etc. Distributes nutrients among adjacent matrix ecosystems such as coral reefs and sea grasses (Ellison & Farnsworth, 2000).NEED INFO. See B. Kjerfve et al., 1997, for extensive and detailed studies. Thresholds are affected by road-building and other processes that divert fresh water from mangroves.See Wolanski et al., 1992; Lugo & Snedaker 1974; Twilley 1995.Integrity of Catchment BasinsCan affect hydrographic processes, freshwater inflow from rivers, runoff, sedimentation rates, and indirectly affect human pressures in coastal mangrove forests.Nutrient and Biotic Transport among neighboring systems Especially interactions with seagrass beds and coral reefs. Significant carbon transfer and flux occurs between mangal and seagrass beds (Hogarth, 1999). In addition, crustaceans and fish often move between the 2 ecosystems, either regularly or between different phases of their life cycles. Corals are vulnerable to sedimentation and to eutrophication, which are both mediated by mangroves. Mangroves serve as nurseries for many early life phases of coral reef fish species. Fish and invertebrates may also move between coral reefs and mangroves within phases of their life cycle as well (Hogarth, 1999).There is a broad range of variation in this. Some mangroves are tightly linked to adjacent systems, while others are much less so. Degree of litter export is extremely variable among systems. Ecological integrity factors for condition Mangroves exhibit vivipary, or the precocious growth of seedlings while still attached to the parent tree. When abscised, the propagules are tough, buoyant, and readily water-dispersed. Competition between mangrove and non-mangrove tree species is rarely a key factor in mangal because the unique hydrologic and edaphic conditions of mangrove ecosystems make it difficult for non-mangrove spp to invade mangal. The peaty muds typical of mangroves have a very high silt content and tend to be fairly inhospitable to most suspension and filter-feeding invertebrates. However, mud-dwelling sesarmid, portunuid, and ocupodid crabs are extremely common (Ellison & Farnsworth, 2000). Table xxx. Ecological integrity factors for condition of mangroves Key FactorJustification for SelectionEcological ThresholdsJustification (e.g., Natural Range of Variation)IndicatorsFactor Relative Priority TemperatureMangroves grow where the average monthly minimum temp. is 20 deg. Celsius. Water temperature Upper threshold = > 31 deg. Air Temperature upper limit: 38 deg. C Temperature fluctuations greater than 10C over short periods and below freezing for more than a few hours have serious negative effects, including structural damage, reduced leaf area (Myers & Ewel, 1990). Reduces leaf area, results in achlorophylly, reduces net photosynthesis, and increases photorespiration (Ellison & Farnsworth, 1996b). Water temps >31 deg C result in a 50-90 percent decline in sessile and mobile invertebrate communities associated with mangroves. Waterborne Dispersal of Mangrove PropagulesAaron Ellison: For state-of-the science, see paper by Rabinowitz (1978). Most recently see paper by P.J. Clarke in Journal of Ecology (August 2001), albeit for the palaeotropics. He reviews the current literature well, though.Dispersal of mangrove epibiont communitiesInfluences the species richness and abundance of mangrove invertebrate and algal communities around mangrove roots (Farnsworth & Ellison, 1996).Insect HerbivoryHerbivores and herbivory are very important regulators of community and physical structure of mangrove ecosystems (Farnsworth & Ellison, 1991). Shorten leaf life span and can hasten premature abscission of Rhizophora propagules (see Farnsworth & Ellison, 1997a.) by increasing abscission rates and decreasing plant productivity; can increase nutrients in soil and near root water communities (Farnsworth & Ellison, 1996). Herbivory negatively impacts seedling growth, influences leaf form and secondary compound chemistry, alter branch architecture and structure, and influences competitive interactions between Avicennia and Rhizophora (Ellison & Farnsworth, 1993).Overall primary production across mangrove ecosystems that is consumed by insects ranges from 10-25%. However, the average is across the globe generally under 5%. Specific study results: Insects consumed 11.1 +/- 32.99% of leaf tissue in Avicennia seedlings (Farnsworth & Ellison, 1996). Herbivores damaged 4.3-25.3% of Rhizophora leaf area and 7.7-36.1% of Avicennia leaf area in Belize (Farnsworth et al., 1996). Herbivory tends to be greater at Medium Water levels vs. at Lower levels. Insect communities of Neotropics are an order of magnitude lower than in the Paleotropics.Elizabeth Farnsworth: It is important to note that neotropical insects in mangrove have not been studied at all to date; several rare species may be found there, and surveys are warranted.Outbreaks of root-boring isopodsReduce root growthCan reduce root growth rates by up to 50% (Ellison & Farnsworth, 1990).Isopod attacks are reduced in presence of certain epibionts (sponges) that repel them chemically. So where these epibionts disappear, isopod attacks may increase.PHSoil chemistry affects not only mangrove tree species themselves, but the ability of algal and invertebrate symbionts to colonize and survive in mangrove ecosystems.pH of pore water within mangrove peat is often acidic, ranging from 4.9 to 6.8, though presence of carbonate materials can elevate pH to7.0 When drained, mangrove peat undergoes rapid acidification (down to 3.5-5.0) which complicates use of mangroves as agricultural land.Dissolved OxygenLimits occurrence of certain species, thereby affecting zonation and community abundance patterns. Rhizophora spp. Can tolerate low oxygen levels better than Avicennia spp. (Ellison & Farnsworth, 1993).Suspended SolidsElizabeth Farnsworth: Heavy metals may be damaging to mangroves; see Ed Klekowski's work [Citation?] on mutational load in Rhizophora mangle. Heavy metals may be damaging to mangroves; see Ed Klekowski's work on mutational load in Rhizophora mangle (Klekowski, et al. 1994a, 1994b, 1994c & 1999; Lowenfeld & Klekowski, 1992).Phosphorus levelsPrimary nutrient limiting growth of some spp (e.g. Rhizophora) at or above mid-water levels (Ellison & Farnsworth, 1996a).SalinityLimits occurrence of certain species, thereby affecting zonation and community abundance patterns. Despite living in saline environments, mangroves require fresh water for growth (acquired through freshwater inflow, as well as through physiological adaptations salt glands, succulent leaves, ultrafiltration processes).Adaptation to saline environments may result from the reduced competition with freshwater species experienced in saline environments (Myers & Ewel, 1990).Avicennia spp. can tolerate high salinity levels better than Rhizophora spp. (Ellison & Farnsworth, 1993). Rhizophora can tolerate soil salinities up to 60-65 ppt, whereas Avicennia may grow at salinities >80-90 ppt (Myers & Ewel, 1990).Post-dispersal PredationImportant factor in structuring mangrove communities Usually occurs by grapsid crabs, coleopteran beetles, and lepidoptera (larvae). Tends to be higher in mangroves near human population centers and at high intertidal sites (Farnsworth & Ellison, 1997a).Data on local predation rates available from Farnsworth and Ellison (1997a).Mutualistic algal and invertebrate epibiont communities in mangrove roots.Many spp. are restricted to mangrove prop roots; contributes to the diversity of mangrove communities. Mutualistic interactions with mangroves: epibionts cause prop roots to produce adventitious rootlets that increase nutrient uptake (esp. N) by mangrove trees, whereas sponges on mangroves grow 1.4-10 x faster than same spp on other substrates (Ellison et al., 1996). Sponges also reduce frequency of isopod attack to roots that reduces root growth rates (Ellison & Farnsworth, 1990). Percent cover of Belizean mangal by root fouling epibionts averages >90% (Ellison et al., 1996). Sponges may account for up to 5-10% of the nitrogen uptake of mangrove trees (Ellison & Farnsworth, 2000). Densities of grapsid crabs may range up to 50-70/m2 Distributions controlled by local larval recruitment dynamics and by larger scale physical factors, such as current regimes. Mutualistic algal and invertebrate epibiont communities in mangrove mud.Crabs oxygenate muddy silt-laden soils, and alter the distribution of toxins within these soils (Ellison & Farnsworth, 2000). They also consume substantial leaf litter quantities and are responsible for significant nutrient recycling. Densities of grapsid crabs may range up to 50-70/m2Note that these crabs are often harvested in large quantities throughout the world.Avian Communities of MangalContribute inorganic nitrogen to mangrove ecosystems; some spp globally restricted to coastal mangrove forests during all or part of their life cycle. Species can contribute Diversity and abundance of resident and migrant birds. Size of nesting colonies of waterbirds. Mangrove pollinationCritical for reproduction and gene flow, dependent on animal communities.Rhizophora mangle is cleistogamous (self-pollinating). Avicennia germinans is out-crossed and can be a local honey-producer. Pollination biology of mangroves is generally poorly studied. It deserves much more research attention.Tree Stand StructureMangrove Tree Abundance/ Canopy CoverCan prevent the regeneration and establishment of some invasive species, especially the pantropical mangrove fern Achrostichum aureum and Brazilian pepper Schinus terebinthifolius in Everglades which can dramatically alter mangrove seedling recruitment. Also limits the growth of any understory plants in order to maintain the simplified architecture that characterizes mangal (Ellison & Farnsworth, 2000).90% canopy cover within mangrove stands (Ellison & Farnsworth, 2000), though this can be highly plastic depending on local edaphic conditions. Invertebrate mud-dwelling species among mangrove rootsMutualistic interaction: inverts receive habitat and protection; through their burrowing, they oxygenate and nutrify soils for mangrove trees.Consumption of litter fall by detritivores (crabs)Critical for nutrient cycling.30-80% of fallen leaves, branches, fruits, flowers, etc. are consumed by detritovores (Ellison & Farnsworth, 2000). Presence, Abundance, and Diversity of EpiphytesInclude orchids, mistletoes, ferns, bromeliads (Ellison & Farnsworth, 2000). These epiphytic species can then harbor important insect communities. Abundance and richness is substantially lower than in upland tropical forests. Epiphytes can support their own mutualisms, e.g., with ants (Rio-Gray et al., 1989).PollinationFlowers of some mangrove spp. support a diverse fauna of native pollinators (Ellison & Farnsworth, 2000), including bees and bats. Maintaining pollination is critical to maintain these pollinating species, as well as maintain heterozygosity, fitness, and gene flow among mangrove trees.Avicennia is probably important to honey bees, which are significant pollinators of adjacent crops.Mangrove Fungal communitiesDominant agents of decomposition of mangrove wood and leaves. Play a significant role in nutrient cycling (Ellison & Farnsworth, 2000).Reptile communitiesEspecially crocodiles and alligators, as top predators.Mangrove PathogensFairly poorly studied, but known to cause massive die-back of some mangrove communities when in outbreak. May become more frequent when mangal is exposed to pollution (Ellison & Farnsworth, 2000).  Ecological integrity factors for size Table xxx. Ecological integrity factors for size of mangroves Key Factor or Disturbance TypesJustification for SelectionAverage size of disturbance and range. Return Intervals of Disturbance Recovery Time Or, if dictated by a species: Minimum population Size Area required for Min Pop SizeRecommendations for Calculations of Minimum Dynamic AreaVG, G, F, P Recom-mendations (including multipliers for ratings of VG, G, F, P)Tree fallsLightning StrikesRegular source of canopy disturbance. Likely to cause changes to soil characteristics and invertebrate communities.Return interval: 8-100 year range Size of Disturbance: << 1km2 Disturbance duration/Recovery time: 1-50 yearsHurricanesMangroves afford protection to upland systems from cyclones/hurricanes, but can be killed by wind damage and tidal surges. Very few species resprout after damage to the main trunk regeneration is almost entirely from seedlings. Hurricanes can cause changes to the species zonation in a mangrove community given resorting of seedlings and differential mortality of species (e.g. Rhizophora survival>Avicennia> Laguncularia in the Neotropics. They also cause direct damage, and can alter sedimentation patterns in both positive and negative ways. While sediment may be removed from extant mangroves along the coast, increased sedimentation from catchment rivers can increase alluvial fans and areas for mangrove expansion (Hogarth, 1999). Recovery Times: 10-50 years Return Intervals: 50-100 years. Size of Disturbance: 1-100 km2Tidal erosion and deposition / Geomorphologic changesSize of Disturbance: 50-500 km2  Additional Information The experts assessed the biodiversity health of mangrove systems. The results are presented in the following tables. Mangrove, Landscape Context Key FactorJustification for SelectionEcological ThresholdsJustification (Natural Range of Variation)IndicatorsFactor Relative Priority Habitat PhysiographyMangroves occupy tropical coastal areas protected from the surf (low energy environments), little deep, and with tendency to accumulate sediment. Those areas act like reservoirs of nutrientes and therefore, they serve of refuge and/or areas of reproduction for many marine, estuaries, migratory and freshwater organisms. It is recommended to avoid modifications in the coastal Physiography (coastal morphology) without the necessary knowledge of the site dynamics . For example: to avoid dredged, backfill, infrastructures and excessive accumulation of sediment.The rates of sedimentation / erosion in the mangrove systems are naturally high, but at the same time extremely variables at site level. Mangroves used to adapt to elevations of the relative sea level up to> 10mm/year. The extraction of water of the phreatic stratum and the exploitation of petroleum contribute to the sinking of the soil, resulting in a "relative" elevation of the sea level, and an increase in rates of erosion. The physiological stress of the mangrove forest is more evident in the external border (in contact with the water), as well as in interfaces of the mangrove with the adjacent terrestrial system.Defoliation can be monitored through aerial photography; the external and internal edges of the mangrove are going to present answers before, the center delays in responding. Indicator: leaves floating after being fallen of the canopy. Identify geomorphologic processes (if in passed years they have seen changes by surveys or interviews) Monitoring of the development and planning of local and river basin infrastructure. 1 Hydrologic Regime: Circulation patternsRegulate the sedimentary dynamic and the regimen of flood Controls salinity, levels of oxygen, dispersion of nutrientes and propagules. Control the balance of freshwater and salted water allowing the optimum physiological and ecological levelsThe threshold is zero. It is necessary a minimum laminar flow without interruptions (ex. Barriers), guaranteeing the maximum area of flood, considering extreme and episodic cyclic events (not necessarily periodic). The nil threshold indicates that any interference in the flow compromises the viability of the system. Assess the velocity of the flow guaranteeing to be laminar (<1m/s), open flow without interruptions. Changes in infrastructure. Monitor the hydraulic behavior of basins (Reference: Elizabeth Farnsworth: Consult Robert Twilley about hydrology and importation/exportation in mangroves). Hydraulic model that describes laminar flow in mangrove and "mudflats". It is important to assure that the flow in the canals in emptied moments to be faster than the laminar flow) see Windevoxel and Alfonso Aguirre for references [Experts, please complete the citation in the References section. Thanks.].2Hydrologic Regimen: Flooded cycleRegulates the balance of freshwater and salted waters permitting the optimum physiological and ecological levels. Determines the competence among the different species of plants and animals, resulting in zoning in the successional processes .To maintain the natural environment of flood (conserving the amplitude for the flood of the river as for the tide, besides of guaranteeing the accumulation of nutrientes and propagules) In each area a natural variation of the flooded area exists, thus is not possible to establish a threshold. The mangrove presents seasonal variations, periodic as not periodic, derived from the climatic, topographical, oceanic a hydrodynamic conditions. Indicators can be defined when you analyze the topography, aerial photography, tide table, registrations of river volume, levels and rivers flood periods, and speaking with local people. Where possible, monitor with brief estimation with posts in zones of flood to measure tides system with bottles to different heights that are filled with the tide. See Robertson & Alongi, 1992 and "1996 Australian publication" [Experts, please provide literature citation. Thanks.] about monitoring methods of tropical ecosystems; contact Ivan Jose Nieff-Univ of Corrientes, Argentina. [Experts, please provide Ivan Jose Nieff' s and-mail address or cite her publications in the References section. Thanks.]2Hydrologic Regimen: Quality of waterThis factor is more related with the transportation of sediment (particles) in suspension. The accumulation of sediments is not a physical requisite to the presence of mangroves. In erosive coasts (high energy), the accumulation of sediments maintains relatively constant the water and substrate levels. Mangroves are found in areas practically without contribute of sediments(coral platforms - keys) even in mouths of rivers, still with discharges of sediment in suspension in the order of 150 x 106 tons/year and a discharge of 1,100 km3/ao (Orinoco River)2 Mangrove, Condition Key FactorJustification for SelectionEcological ThresholdsJustification (Natural Range of Variation)IndicatorsFactor Relative Priority Tide (amplitude) Mangroves occupy the coastal areas showered by the flow and reflux of the tides. Tides are a requirement for the establishment of the mangroves, once they guarantee the success of the competence among the facultative halophytes plants (typical of the mangrove) and the glicophytes (of freshwater).Mangroves develop in micromareales environments (< 2m de amplitude); mesomareales (2 - 4 m), and macromareales (>4m). Extreme tides (> 10 meters of amplitude) are associates to the erosion of the coast and to the lost of the mangrove forests and the invertebrates associated. Tides of small amplitude reduce the laminar flow and are associated with water stagnation, contributing to the accumulation of salts, reaching values> 60 ppm. 1Surf (energy) Mangroves occupy coastal tropical protected areas of the surf.In sectors of the coast characterized by high energy of the water (coastal and/or rivers), mangroves develop behind barriers or verge of sand. The strong surf (high energy) produces erosion destabilizing the substrate, collapsing the trees and impeding the establishment of the propagules. 2Freshwater: RiversThe volumes of freshwater represent an important fountain of subsidiary energy to the development of the system.  The excess of freshwater impedes the entrance of the wedge saline creating favorable conditions of competition to the glicophytes with relation to the facultative halophytes (vegetable species of the mangroves)1Freshwater: Rain Mangroves develop mostly where the average of annual rainfall is @ 1,300 mm, throughout the year. Mangroves systems are found mostly in rainfall conditions between > 1,000 mm/year y < 8,000 mm/yearTemperature The distribution of mangroves presents good correlation with the temperatures of the sea surface (> isotherm of the 20o C. Mangroves occur rarely where temperatures fall under the isotherm of the 20o C There has not been defined a threshold for high temperatures, but in St. Marta rose at more than 32 degrees and the mangrove does not recuperate; changes in temp going up affect mainly the fauna. See Walsh for Introduction to the Mangrove Ecology & Project GTZ-SEES for the cienaga St. Marta-Colombia. [Experts, please complete the citation in the References section.All the typical vegetable species of mangroves are sensitive to frosts, and not survive to long periods with temperatures under the 0o C. It is important to consider the temperature factor when there are drastic changes in the climate. Mangroves do not survive where very low temperatures (under the 0o C) have high frequency of recurrence, neither where the photosynthesis is reduce for long periods throughout the year. 2SalinityThe typical vegetal species of the mangroves are facultative halophytes. These occupy the areas where the plants are not tolerant to salt.Salt water per se is not a physical requisite to the presence of typical plants of mangrove.1ETP (potential evapotranspiration )Near the 90% of the mangroves are found in humid regions.Mangroves are practically unknown in dry weather.Mangroves are found occasionally in sub-humid climates and its occurrence is considered exceptional in conditions of semi-dryness2Reproductive successKey for the occupation, recuperation and recolonization of the system. Maintains the diversity of the genetic information flow.Thresholds are specifics to each site. It is fundamental the presence of propagules and seedlings, besides the adequate habitat phisiographyc conditions. The number of propagules necessary to maintain the stand is ( to the rate of loss of live of the forest or stand, more than the rate of loss of live of the seedlings. The number of propagules by species varies seasonally and it depends of the site. Consult: Snedaker & Snedaker (1984)1 Endangered species associated with mangroves: Mangrove rivulus (fish) Rivulus marmoratus. Species of special concern in Florida. See http://floridaconservation.org/pubs/endanger.html (official list, 1997) Florida manatee (Trichechus manatus) Loggerhead turtle (Caretta caretta) American crocodile (Crocodylus acutus) see http://www.nps.gov/ever/ed/eddanger.htm) and http://www.aoml.noaa.gov/flbay/mari95.html for 1995 review of research and management in Florida Bay Atlantic salt marsh snake (Nerodia fasciata taeniata) See http://www.nature.nps.gov/wv/article2.htm from the National Park Service Intertidal Trapdoor Spider (Idioctis yerlata) in Queensland, Australia (see http://www.qmuseum.qld.gov.au/features/endangered/animals/trapdoor_spider.asp for description and management efforts) Wood stork ((Mycteria americana) nests in mangroves in the Florida Everglades Asian small-clawed otter Atlantic hawksbill turtle uses mangroves in southeast U. S. -- from U. S. National Marine Fisheries Service (see report at http://environment.about.com/library/weekly/blturt2.htm?once=true&) Pemphis acidula Fort. (Lythraceae) -- mangrove associate (plant) in the Phillipines. Mangrove-associated seahorses in southeast Asia Bengal tiger (Panthera tigris tigris) in the Sunderbans mangal of India and Bangladesh Proboscis monkey, Malaysia Avicennia marina ssp. australasica considered rare in Australia Mangrove finch Brown pelican (Pelicanus occidentalis caroliniensis and Pelicanus occidentalis occidentalis) http://endangered.fws.gov/i/b/sab2s.html Mangrove orchid (Dendrobium mirbelianum), Queensland, Australia. See http://www.env.qld.gov.au/environment/plant/endangered/wie.html Smalltooth Sawfish (Pristis pectinata) See proposed listing at Federal Register: April 16, 2001 (Volume 66, Number 73) Proposed Rules Page 19414-19420 Mangrove cuckoo endangered on Montserrat Nightingale reed-warbler (Acrocephalus luscinia) in Saipan Mariana common moorhen (Gallinula chloropus) in Saipan Information gaps and caveats Aaron Ellison: The dependence of mangrove species diversity and stand structural characteristics on rainfall and other large-scale climatic factors varies among geographic regions. Ellison (2001, in press) provides some global comparisons. The dependency is relatively weak in Central America (Ellison 2002, in press) but strong in Australasia. Elizabeth Farnsworth: Sea-level rise will soon rival pollution and habitat conversion as a major threat to mangrove ecosystems, especially the shallow carbonate platforms along the Caribbean coast and islands. Recommended priorities for conservation-driven research agenda and next steps Clarify that the site managers should get support from local experts in their region and not think that they can simply apply the information from the formats. Do an easy to use manual (going from basic to complex). Making it digestible (language among technician and ordinary) Example: Explain in a simple way (naturalistic style) giving references what healthy systems and basic indicators are. It is recommended to seek advice from experts in communication or environmental education. [Recommendation of Elizabeth Farnsworth. Consult with Candy Feller, Smithsonian Institution (SERC, Edgewater, Maryland), who has worked in projects of education in Belize.] When taking decisions of management is important to know the tropic network of the system. Consult with local expert (member of the community) When defining buffer zones in protected areas, is important to consider the area that is defined by extreme events. It is recommended to carry out a landscape ecology analysis (historic analysis) through mapping, satellite images, ethnoscience and aerial photography. References Cintrn-Molero, G. and Schaeffer-Novelli, Y. 1981. I. Los manglares de la costa brasilea: revisin de la literature. II. Los manglares de Santa Catarina. Informe Tcnico preparado para la Oficina Regional de Cincia y Tecnocloga para Amrica Latina y el Caribe de UNESCO y la Universidad Federal de Santa Catarina, Brasil. 67p. Cintrn-Molero, G. and Schaeffer-Novelli, Y. 1983. Introduccin a la ecologa del manglar. UNESCO/ROSTLAC, Montevideo. 109p. Cintrn-Molero, G.; Lugo, A.E.; Pool, D.J. & Morris, G. 1985. Mangroves of arid environments in Puerto Rico and adjacent islands. Biotropica, 10: 110-121. Duke, N. C., M. C. Ball, and J. C. Ellison. 1998. Factors influencing biodiversity and distributional gradients in mangroves. Global Ecology and Biogeography Letters 7: 27-48. Ellison, A.M. and E. J. Farnsworth. 1996. Anthropogenic disturbance of Caribbean Mangrove Ecosystems: past impacts, present trends, and future predictions. Biotropica 28: 549-565. Ellison, A. M. and E. J. Farnsworth. 2000. Mangrove communities. In M.D. Bertness, S.D. Gaines, and M. E. Hay (eds.). Marine Community Ecology. Pp. 423-442. Sinauer Associates, Sunderland, MA. Ellison, A. M., E. J. Farnsworth, and R. E. Merkt. 1999. Origins of mangrove ecosystems and the mangrove biodiversity anomaly. Global Ecology and Biogeography 8:95-115. Feller, I. C. 1995. Effects of nutrient enrichment on growth and herbivory of dwarf red mangrove (Rhizophora mangle). Ecological Monographs 65:477-506. Gomez, M. and S. Winkler. 1991. Bromelias en manglares del Pacifico de Guatemala. Rev.Biol.Trop 39:207-214. Graham, A. 1995. Diversification of Gulf Caribbean mangrove communities through Cenozoic time. Biotropica 27 (1):20-27. Hogarth, P. 1999. The biology of mangroves. Oxford University Press, Oxford, United Kingdom. Linacre, E. and Geerts, B. 1997. Climates and weather explained. Routledge, London. Lugo, A.E. and Snedaker, S.C. 1974. The ecology of mangroves. Ann. Rev. Ecol. Syst., 5: 39-64. Murren, C.J. and A. M. Ellison. 1996. Effects of habitat, plant size, and floral display on male and female reproductive success of the neotropical orchid Brassavola nodosa. Biotropica 28:30-41. Pea, G.M. and Vasquez, P.G. 1985. Un relicto de manglar en San Pedro (Piura): estudio preliminary. Boletn de Lima, 7 (42): 27-32. Rico-Gray, V., J. T. Barber, L. B. Thien, E. G. Ellgaard, and J. J. Toney. 1989. An unusual animal-plant interaction: feeding of Schomburgkia tibicinis (Orchidaceae) by ants. Amer.J.Bot 76: 603-608. Schaeffer-Novelli, Y., Cintrn-Molero, G., Adaime, R.R. and Camargo, T.M. 1990. Variability of mangrove ecosystem along the Brazilian coast. Estuaries, 13 (2): 201-219. Schaeffer-Novelli, Y. and G. Cintrn-Molero. 1993. Mangroves of arid environments of Latin America. In: Lieth, H. & Al Masoom, A. (eds.). Towards the rational use of high salinity tolerant plants. Vol. 1. Kluwer Academic, Dordrecht, pp. 107-116. Tomlinson, P. B. 1995. The botany of mangroves. Cambridge University Press, Cambridge, UK. Twilley, R. R. 1995. Properties of mangrove ecosystems and their relation to the energy signature of coastal environments. In: Hall, C. A. S. (ed.). Maximum power . pp. 43-62. University of Colorado Press, Boulder, Colorado, USA. Walter, K. S. and H. J. Gillett (eds.). 1998. 1997 IUCN red list of threatened plants. Compiled by the World Conservation Monitoring Centre. IUCN The World Conservation Union, Gland, Switzerland and Cambridge, UK. 862 pp. Burger, J., K. Cooper, D. J. Gochfield, J. E. Saliva, C. Safina, D. Lipsky, and M. Gochfield. 1992. Dominance of Tilapia mossambica, an introduced fish species, in three Piero Rican estuaries. Estuaries 15: 239-245. Ellison, A.M. 1999. Cumulative effects of oil spills on mangroves. Ecological Applications 9: 1490-1492. Ellison, A.M. 2000. Mangrove restoration: Do we know enough? Restoration Ecology 8: 219-229. Ellison, A.M. and E.J. Farnsworth. 1990. The ecology of Belizean mangrove root-fouling communities. I. Epibenthic fauna are barriers to isopod attach of red mangrove roots. Journal of Experimental Marine Biology and Ecology 142: 91-104. Ellison, A. M. and E. J. Farnsworth. 1993. Seedling survivorship, growth, and response to disturbance in Belizean mangal. American Journal of Botany 80: 1137-1145. Ellison, A.M. and E. J. Farnsworth, 1996a. Spatial and temporal variability in growth of Rhizophora mangle saplings on coral cays: links with variation in insolation, herbivory, and local sedimentation rate. Journal of Ecology 84: 717-731. Ellison, A.M. and E. J. Farnsworth. 1996b. Anthropogenic disturbance of Caribbean Mangrove Ecosystems: past impacts, present trends, and future predictions. Biotropica 28: 549-565. Ellison, A. M. and E. J. Farnsworth. 1997. Simulated seal level change alters anatomy, physiology, growth, and reproduction of red mangrove (Rhizophora mangle L.) Oecologia 112: 435-446. Ellison, A. W. and E. J. Farnsworth. 2000. Mangrove communities. In: Bertness, M. D., S.D. Gaines, and M. E. Hay (eds.) Marine Community Ecology, Pp. 423-442. Sinauer Associates, Sunderland, MA. Ellison, A. M., E. J. Farnsworth, and R. R. Twilley. 1996. Facultative mutualism between red mangroves and root-fouling sponges in Belizean mangal. Ecology 77: 2431-2444. Farnsworth, E.J. and A.M. Ellison. 1991. Patterns of herbivory in Belizean mangrove swamps. Biotropica 23: 555-567. Farnsworth, E.J. and A. M. Ellison. 1996. Scale dependent spatial and temporal variability in biogeography of mangrove epibiont communities. Ecological Monographs 66: 45-66. Farnsworth, E. J. and A. M. Ellison. 1997a. Global patterns of pre-dispersal propagule predation in Mangrove Forests. Biotropica 29: 318-330. Farnsworth, E.J. and A. M. Ellison. 1997b. The global conservation status of mangroves. Ambio 26: 328-334. (An excellent review of global threats to mangroves and their ecological impacts. ) Farnsworth, E.J., A.M. Ellison, and W.K. Gong. 1996. Elevated CO2 alters anatomy, physiology, growth and reproduction of red mangrove (Rhizophora mangle L.). Oecologia 108: 599-609. Feller, I.C. 1993. Effects of nutrient enrichment on growth and herbivory of dwarf red mangrove. Ph.D. Dissertation, Georgetown University, Washington, D.C. Hogarth, P. 1999. The biology of mangroves. Oxford University Press, Oxford, United Kingdom. Kjerfve, B., L. D. de Lacerad and E. H. S. Diop, (editors). 1997. Mangrove ecosystem studies in Latin America and Africa. UNESCO, Paris. Lugo, A.E. and Snedaker, S.C. 1974. The ecology of mangroves. Ann. Rev. Ecol. Syst., 5: 39-64. Myers R. L. and J. J. Ewel, eds. 1990. Ecosystems of Florida. University of Central Florida Press, Gainesville. Rabinowitz, D. 1978. Dispersal properties of mangrove propagules. Biotropica 10: 47-57. Rico-Gray, V., J. T. Barber, L. B. Thien, E. G. Ellgaard, and J. J. Toney. 1989. An unusual animal-plant interaction: feeding of Schomburgkia tibicinis (Orchidaceae) by ants. Amer.J.Bot 76: 603-608. Twilley, R. R. 1995. Properties of mangrove ecosystems and their relation to the energy signature of coastal environments. In: Hall, C. A. S. (ed.). Maximum power . pp. 43-62. University of Colorado Press, Boulder, Colorado, USA. Wolanski, E., Y. Mazda, and P. Ridd. 1992. Mangrove hydrodynamics. Pp. 43-62. In Robertson, A.I. and D.M. Alongi (eds.) 1992. Tropical mangrove ecosystems. American Geophysical Union. Washington, DC Klekowski, E.J., Jr., L. E. Corredor, J. M. Morrell, and C. A. del Castillo. 1994a. Petroleum pollution and mutation in mangroves. Marine Pollution Bulletin 28: 166-169. Klekowski, E.J., Jr., L. E. Corredor, R. Lowenfeld, E. H. Klekowski, and J. M. Morrell. 1994b. Using mangroves to screen for mutagens in tropical marine environments. Marine Pollution Bulletin 28: 346-350. Klekowski, E. J. Jr., R. Lowenfeld, and P. K. Hepler. 1994c. Mangrove genetics. 2. Outcrossing and lower spontaneous mutation rates in Puerto Rican Rhizophora. International Journal of Plant Science 155: 373-381. Klekowski, E. J. Jr., S. A. Temple, A. M. Siung-Chang, and K. Kumarsingh. 1999. An association of mangrove mutation, scarlet ibis, and mercury contamination in Trinidad, West Indies. Environmental Pollution 100: 1-5. Lowenfeld, R. and E. J. Klekowski, Jr. 1992. Mangrove genetics. 1. Mating systems and mutation rates of Rhizophora mangle in Florida and San Salvador Island, Bahamas. International Journal of Plant Science 153: 394-399. (Referencia.??? On p.21. Modelo hidrulico que describe flujo laminar en manglar y mudflats Alfonso Aguirre for references on page 22 . 1996 Australian publication sobre mtodos de monitoreo de ecosistemas tropicales on page 23. Ivan Jose Nieff..on page 23. Proyecto GTZ-VE para la cienaga Sta Marta-Colombia, on page 26. Robertson, A.I. and D.M. Alongi (eds.) 1992. Tropical mangrove ecosystems. American Geophysical Union, Washington, DC. Snedaker, S.C. & Snedaker, J.G. (eds.) 1984. The mangrove ecosystem: research methods. Monographs on Oceanographic Methodology, vol. 8. United Kingdom, UNESCO Publication. Walsh for Introducion al la Ecologia de Manglar, on page 26. Windevoxel for references, on Page 22. Recommended resources Cicin-Sain, B. & Knecht, R.W. 1998. Integrated coastal and ocean management: concepts and practices. Washington, D.C., Island Press. 517 p. Citrn-Molero, G. & Schaeffer-Novelli, Y. 1983. Mangrove forests: ecology and response to natural and man induced stressors. In: Coral Reefs, Seagrass beds and Mangrove: their Interaction in the Coastal Zones of the Caribbean. UNESCO Reports in Marine Science, 23:87-113. Cintrn, G. & Schaeffer-Novelli, Y. 1983. Introduccin a la ecologa del manglar. UNESCO/ROSTLAC, Montevideo. 109p. Cintrn-Molero, G. & Schaeffer-Novelli, Y. 1992. Ecology and management of New World mangroves: 233-258. In: Seeliger, U. (ed.), Coastal Plant Communities of Latin America. California Academic Press. 392 p. Ellison, A. M. 2001. Macroecology of mangroves: large-scale patterns and processes in tropical coastal forests. Trees: Structure & Function (in press to be published in December 2001). Ellison, A. M. 2002. Wetlands of Central America. Wetlands Ecology & Management (in press). Ellison, A. M. and E. J. Farnsworth. 2000. Mangrove communities. In M.D. Bertness, S.D. Gaines, and M. E. Hay (eds.). Marine Community Ecology. Pp. 423-442. Sinauer Associates, Sunderland, MA. FAO forestry paper 117, Mangrove forest management guidelines (Rome, 1994). Field, C. (ed.). 1996. Restoration of mangrove ecosystems. ITTO and ISME, Okinawa, Japan. Hamilton, S.L. & Snedaker, S.C. (eds.). 1984. Handbook for mangrove area management. Hawaii, IUCN, UNEP & East-West Center, Environment and Policy Institute. 123 p. Hogarth, P. 1999. The biology of mangroves. Oxford University Press, Oxford, United Kingdom. Kathiresan, K. and B. L. Bingham. 2001. Biology of mangroves and mangrove ecosystems. Adv. Mar. Biol. 40:84-251. Longhurst, A. 1998. Ecological geography of the sea. Academic Press. 398 p. Mangrove Restoration, special section in Restoration Ecology 8(3): 217-259. September 2000. Schaeffer-Novelli, Y.; Cintrn-Molero, G.; Adaime, R.R. & Camargo, T.M. 1990. Variability of mangrove ecosystem along the Brazilian coast. Estuaries, 13 (2): 201-219. Schaeffer-Novelli, Y. & Cintrn, G. 1990. Status of mangrove research in Latin America and the Caribbean. Boletim Instituto Oceanogrfico, So Paulo, 38 (1): 93-97. Schaeffer-Novelli, Y. & Cintrn-Molero, G. 1993. Mangroves of arid environments of Latin America. In: Lieth, H. & Al Masoom, A. (eds.). Towards the Rational Use of High Salinity Tolerant Plants, vol. 1. Kluwer Academic, Dordrecht, pp. 107-116. Schaeffer-Novelli, Y.; Cintrn-Molero, G.; Soares, M.L.G. & T. De-Rosa, M.M.P. 2000. Brazilian mangroves. Aquatic Ecosystem Health and Management, 3: 561-570. Sherman, K.; Alexander, L.M. & Golg, B.D. (eds.) 1993. Large marine ecosystems: stress, mitigation, and sustainability. Washington, D.C., American Association for the Advancement of Science Press. 376 p. Snedaker, S.C. & Snedaker, J.G. (eds.) 1984. The mangrove ecosystem: research methods. Monographs on Oceanographic Methodology, vol. 8. United Kingdom, UNESCO Publication. Suman, D.O. (ed.). 1994. El ecosistema de manglar en America Latina y la cuenca del Caribe: su manejo y conservacin. Rosensteil School of Marine and Atmospheric Science, University of Miami, Miami, FL Note by Aaron Ellison: Kathiresan and B. L. Bingham (2001) and Ellison & Farnsworth (2000) provide a good review of the mangrove ecosystem. The special feature in Restoration Ecology (September 2000) on mangrove restoration, along with C. Fields 1996 book on mangrove restoration Restoration of mangrove ecosystems provide a wealth of management techniques. All site managers should have the FAO forestry paper 117, Mangrove forest management guidelines (Rome, 1994), on their shelves. A good, relatively recent overview of issues for management and conservation of mangroves in Latin America is provided by D. O. Suman, editor (1994), El ecosistema de manglar en America Latina y la cuenca del Caribe: su manejo y conservacin, available from the Division of Marine Affairs & Policy, Rosensteil School of Marine and Atmospheric Science, University of Miami, 4600 Rickenbacker Causeway, Miami, FL 33149, USA. (Recommended publications are in bold.) Chapter5Freshwater habitat types 5.1. Large wetland systems General description and geographic variation Wetlands include large swampy floodplains and adjacent vegetation. The wetland systems discussed here also include some other non-freshwater systems like coastal wetlands that share many of the same key factors. Participants: Eduardo Asanza, Mark Breyer, Peter Esselman, Katie Frohardt, Wolfgang J. Junk, David Marques, Alonso Ramirez, Carolina da Silva, David Braun (facilitator & recorder), Jonathan Higgins (facilitator), Tarcisio Granizo (TNC staff). Geographic Variation (Describe succinctly any major floristic, structural or biogeographic variation across the Americas): Major forces in floodplain systems are hydrological regime, source of water, nutrient or water chemistry, vegetation cover, lateral connectivity, and biotic components. They vary at different geographical scales. 1. hydrologic regime: 1.1. type of pulse: slow and predictable annual pulse systems: these are the large monomodal floodplain rivers in S. America: Orinoco, Amazon, Negro, Paraguay, Parana, and large sheet of flooded wetlands: Pantanal, Parana, Bananal, Llanos bajos, mojos in Bolivia, savannas in ??, [Experts, please indicate where the "savanas" is. Thanks.] Interrios in Argentina and wetlands in Chaco. These are monomodal systems, dry season wetlands. Study of these systems in absence of long term hydrologic data could be done through tree cores and/or sediment analyses. slow, unpredictable and irregular pulse system: in dry Chaco and Caatinga. fast, predictable, frequently pulsing systems that are tidally influenced for short time: Pacific rivers in Ecuador, and freshwater coastal tidal wetlands. fast, unpredictable pulsing system: all minor low-order and rain driven systems. Aquatic insects are specifically adapted to these systems. 1.2. amplitude: 1.2.1. large amplitude: e.g., up to 15 m in Amazon, because of large basin size and floodplain. 1.2.2. medium amplitude: e.g., about 1 m in Pantanal, relatively large floodplain compared with basin. 1.2.3. stable/low amplitude systems: small fluctuations, e.g., coastal freshwater systems. Typha spp. are good indicators. Terrestrial bromeliad, which sits at high mean flood in Pantanal, is a good indicator. [Experts, you just mentioned that Pantanal is a medium amplitude system (see 1.2.2.). Is the bromeliad considered here an indicator of stable/ low amplitude system? Would you please clarify the roles of bromeliad in these two systems? Thanks. ] 2. source of water: Zonation of plant communities can be associated with changes in water levels. This factor is related to hydro regime or water chemistry. 2.1. rainwater: savannas or Roraima, parts of Bananal, and Pantanal. They have completely different chemistry conditions. 2.2. permanent connection to lagoons/lakes: e.g., freshwater tidal wetlands, Lago Titicaca and associated wetlands. 2.3. permanent connection to large rivers: e.g., Amazon, Rio Paraguay. 3. nutrients/water chemistry: This factor is largely determined by the underlying geomorphology of the basin. 3.1. rich: white water from the Andes, high nutrients with neutral pH, high primary production floodplains with fertile soils. 3.2. intermediate: mostly clear, transparent, low dissolved or suspended solids, lower nutrients. 3.3. low : mostly blackwater some clearwater these have lowest nutrients, low sediment, highly acidic, low pH , low conductivity, inhibits primary production because of humic acids. Exceptions: some black water systems in Ecuadorian Amazon are very productive because soils are eutrophic or mesotrophic. 4. Vegetation cover: related to climate. Vegetation cover type depends on annual rainfall pattern for savanna or forest distincation. Herbaceous vegetation is determined by water level. 4.1. dominated by savanna: with pronounced dry season, 6-8 months, and a combination of flood, fire, and drought stress, e.g., Pantanal and Bananal. Variation on this is Chaco/Caatinga where shrubs or scrubs and scattered trees are found. 4.2. dominated by forest: trees can grow in areas of lower water stresses. 4.3. dominated by herbaceous: These are stable, permanent wetlands, e.g., coastal wetlands thoughout Brazil and Uruguay. Some times are associated with river, e.g., lower Parana. 5. lateral connectivity: describes the connection of floodplains habitats between permanent aquatic and terrestrial systems. Many floodplain systems belong to this type. Connectivity results in systems constantly progressing to steady state and kick back, some moving faster to steady state (high) than others (low connectivty much time to develop stable communities). Species are preferentially selected. It is important to maintain habitat diversity over lateral range of floodplain system. The scale of lateral connectivity increases when moving downstream. This follows the geomorphological changes associated with larger river systems. A lot of variations or gradients exist within each class of connectivity. 5.1. high: permanent exchange of river with floodplain. Water, nutrients and biotic components are always exchanging. 5.2. medium: annual, short exchange during maximum flood. The system is constantly being reset. 5.3. low: connectivity occurs on extreme events at much longer time scale, e.g., 5 to 50 years or more depending on large scale climatic processes. For example, lakes far off main rivers in western Amazonia, exchange may occur at 800 to 1200 year interval during huge flood. 6. biotic components: biota include: resident biota to wetlands (includes endemics), migrants into wetlands from terrestrial (jacar) and deepwater (fishes), long distance migrants (birds and fishes), occasional visitors (these dont care whether wetlands are there but add to biodiversity), endemic invertebrates and vertebrates species living exclusively on floodplain trees. Elimination of any species likely results in direct loss of associated species (e.g., parasites). Paleoclimatic history constrains type of wetlands and possible biota. Any drastic change in climate will result in major shift in biota. Temperate systems have had relatively short time periods for evolutionary process. In some parts of the Tropics, longer periods have existed and allowed much more speciation. Currrent biota is constrained by past history. In Pantanal, biota is relatively poor because of extreme dry period during which extinction rates were high. Open niches for large populations (These species are rare elsewhere, and interspecific competition is low) with less diversity. In Amazon, there has been long period of relatively uninterrupted disturbance which has resulted in smaller populations with much more diversity. In general, nutrients are linked to species richness or abundance depending on group and requirements. This relationship varies depending on the species composition and the scale. For example, in central Amazonia, macrophytes diversity increases with nutrients and pH. Floodplain forest diversity is linked to inundation periodicity. Questions have been raised regarding whether blackwater vs. whitewater floodplain forests are more diverse. Depending on the biotic group, whitewater systems may have more macrophytes while blackwater may have more fishes. There is a dramatic difference in biomass, whitewater systems are high in biomass due to higher nutrient availability. Pantanal is a relatively nutrient limited system, intermediate productivity because of climate stress. Fish migration or spawning is keyed to high water level. In Rio Parana, fish migrate during flood season or high water level. (This is true in many rivers.): Fishes have principal spawning period during increasing flood pulse because more habitats are becoming available. In many places, fishes move and get ready for spawning with rise (the hydro regime May pause) and then another rise comes and fish can spawn very successfully. Many aquatic vertebrates follow this rise in hydro pulse. In Rio Parana and other stable systems that are characterized by relatively low fluctuation, the presence of emergent macrophytes is an indication of physical limit in water level. The size or species composition of macrophytes changes with water level. The fluctuations in the size or species composition of macrophytes can therefore give evidence for changes in mean water level. Community types/zonation and major gradients within the system (patterns) Zonation of plant communities is associated with changes in water level that is related to hydrological regime. In floodplain river system, herbaceous plant communities dominate stable permanent wetlands; savanna vegetation prevails in areas with pronounced dry season and a combination of flood, fire and drought stress; forests are common in areas of lower water stresses. In the Amazon basin, the underlying geomorphology determines the water chemistry of the rivers. Three river types are distinguished: 1) White water rivers originated from the Andes, with rich nutrients, neutral pH, and high production floodplains. 2) Clear water rivers with lower nutrients, low dissolved or suspended solids, and transparent water. 3) Black water rivers with lowest nutrients, low pH, low sediment, low conductivity and low primary production. Examples CENTRAL AMAZON MAINSTEM AROUND MANAUS Key FactorsJustification for Factor SelectionMin. Integrity Threshold(s) Justifi-cation for Threshold Indicators for Field-Based MonitoringNatural hydrologic regime: slow pulse, predictable floodplain systems. note large variation in most rivers as one moves upstream or downstream (true for Amazon, Parana, Paraguay, and Araguay)Important for maintaining habitat diversity and landscape pattern and ecological processes. Annual pulse: key for fish migration, nutrient cycling, and connection to wetlands. Extreme pulse/drought: strong set back effects and export of accumulated organic material. Example: in Amazon, at Manaus, 10 m annual fluctuations in water level (18 28m) Annual pulse regular mean max/min? Extreme pulse/drought these should be set to ensure a maximum minimum flow and minimum maximum flow at some recurrence interval. (Note what are the key species. May need to set hydro expectations for multiple years of extreme dry or wet years to reset system.)1. Use gauges to monitor water level. 2. Annual pulse: measure successful catfish and characid reproduction and balanced age structure. [also indirectly affects caiman, turtles reproduction or habitat structure.] 3. Radar or satellite can be used to measure major vegetation units and flooding extent. 4. Extreme pulse: monitor those species (tree spp.?) that depend on rare events (habitat complexity measure); or monitor reduction in recruitment of tree species. 5. Consultation with local communities on patterns and frequency of flooding regime. 6. Without long term hydrological data sets, it might be useful to take tree ring cores to show historical flooding patterns (This works for about 80% of tree species, e.g., Macrolobium acaciifolium, Genipa spruceana. Need to sample many cores > 100 to remove individual tree influence. Density of floodplain forestReduces energy in system and helps maintain overall system geomorphology.Ages of tree species in mature stands of floodplain forest (up to 400 years old). Natural sedimentation rates.Presence of old floodplain forests (via tree cores). Predominance of Salix spp. which need new sediment to establish. Identify channel changes in western Amazon region via remote sensing. Remote sensing of upstream deforestation. Sediment traps?Age, structure, and composition of floodplain forestDetermines ecosystem function. A good indicator of persistence of biological diversity of the forest. Spectrum of age/size classes: e.g., for some species, need to have high proportion of small size with few older individuals, and no class should be missing. This depends on autecology. Diversity of species (e.g., expect about 200 tree species on islands in central Amazonian region), percent of forest cover; especially need to have older species which can permit degraded system to rebound. In Amazonia, there is a gradient in tree species diversity, increasing from east to west. Locally, highest diversity exists on highest levees, but most human activities occur here as well.Presence of some species of Ficus indicates recent disturbance. [Experts, would you please indicate what these Ficus species are. If you know of any published studies, please cite. Thanks.] Consulting community members (e.g. fishers) who are knowledgeable about forest history.Communities of fruit-eating fishes Dispersal and maintenance of genetic diversity of floodplain tree species that depend on fishes as dispersal agents. Good protein source for humans.About 20% (check #) [Experts, please provide literature supporting this estimate. Thanks.] of fishes are fruit eaters, but some modify feeding habits within its life history. fisherman knowledge landings at major ports Migratory fishesSensitive to large scale connectivity. Good commercial value.Maintain connectivity within system (up to 2000 km for some catfishes) Check literature for possible densities but likely unfruitful, [Experts, if you find any published studies, please cite. Thanks.] estimate production of 900,000 tons in Amazon per year. Over-fishing has depleted large catfish in many areas.fisherman knowledge landings at major portsFreshwater predators (e.g., freshwater dolphins, giant otter, osprey, caiman, ibis and other avifauna.)Control trophic structure and nutrient cycling. Sensitive to toxic bioaccumulation.check literature for densities of predators, e.g., population densities of black caiman vary from 5 to 280 per km in Amazonian Ecuador. [Experts, if you find any published studies, please cite. Thanks.]Consultation with local fisherman/hunter. Diurnal census of birds, dolphins (rookeries). Nocturnal census of caiman. Census of giant otter nests. PANTANAL (extending from 16 to 220S, 53 to 580W) Key FactorsJustification for Factor SelectionMin. Integrity Threshold(s) Justifi-cation for Threshold Indicators for Field-Based MonitoringHydrologic regime :slow, predictable annual pulse; medium amplitude. Variation in timing and duration occurs longitudinally along Paraguay mainstem. More rain in north than south. Rain coincides with flooding in north. Severe drought stress occurs in north due to flood and rains end together; delayed flooding in south, usually 2-3 months after rains.Annual pulse: key for fish or bird movement and migration, nutrient cycling, creation and connection to wetlands and geomorphology. Extreme pulse/drought: important for long-term maintenance of habitat diversity, landscape patterns and ecological processes. May be a multiannual occurrence of extremes.Annual pulse: regular mean max/min??? Extreme pulse/drought: should be set to ensure a maximum minimum flow and minimum maximum flow at some recurrence interval based on historical record. May need to set hydro expectations for multiple years of extreme dry or wet years to reset system. Threats: during multiple drought years more land exposed which leads to more habitat destruction because of increased access for cattle ranching and deforestation.Vegetation cover: In Pantanal, mosaic of savanna that shifts inter-annually. Cattle have effects on this pattern. Variation of vegetation can be observed from east to west and north to south along main rivers.Provides key habitat structure for many species, productivity of organic material, and input of nutrients for system. Mosaic of savana (campo limpo and campo sujos with introduced species) with capoes e cordilheras: Thresholds of overall natural vegetation cover cannot go below x% of original cover. [Experts, please estimate the X value. If you know of any published studies, please cite. Thanks.] Thresholds are the minimum needed to maintain structure, composition and spatial arrangement of the mosaic. The mosaic shifts with wet years (more forest) and dry years (more grassland). It is extremely important not to modify the key habitats capoes and cordilheras, nor refuges for aquatic species, e.g., bahias or deep lakes or permanent water bodies.Remote sensing data. Field work to measure composition of the vegetation.Natural geomorphological process: variation depending on basin characterisitcs, sediment load, and hydraulic geometry. Key to maintain channel process, formation of habitat and vegetation structure, and lateral floodplain access.Maintain natural dynamics of fluvial morphology (sediment load from uplands, lateral connectivity from main channels). Threats: In Rio Taquari, additional erosion from upland land use of many basins forces the formation of a braided system, which is not natural today. In Manso, lower sediment loads from dam increase downstream erosion. Low declivity of region translates into potential for huge spatial effects in flooding area and vegetation. Roads and levee constructions can be devastating because they block lateral connectivity.Sampling: simple traps??. Estimation of natural erosion via basin characteristics? Literature Cited No information available. Recommended resources Angermeier ? and Karr ?. 1983. Title?? Env. Biol. Fishes 9: 117-135. [Experts, please complete the citation. Thanks.] Dugan, P. 1992. Conservacin de humedales. UICN, Suiza Giovannini, S.G.T.; Motta Marques, D.M.L. da.; and B. Irgang. 2000. Sucession in experimental constructed wetlands under different hydroperiods in subtropical climate. Verh. Int. Ver. Limnol. 27, December. In press. Goulding M., M.L. Carvalho & E.G. Ferreira 1988. Rio Negro: Rich life in poor water. The Hague: SPB Academic Publishing. Granizo, T. (compilador). 1997. Uso sostenible de humedales en Amrica del Sur: Una proximacin. UICN-Sur. Quito. 126 p. Hamilton,S.K. 1999. Potential effects of a major navigation project (Paraguay-Parana Hidrovia) on inundation in the Pantanal flood plains. Regulated Rivers: Research and Management 15: 289 299. Junk, W.J. 1993. Wetlands of tropical South America. In: Whigham, D.F., Dyrygova, D., Hejny, S. (eds.) Wtlands of the world: Dordrech: inventory ecology and management. Kluwer Academic, p.679-739. Junk, W.J., da Silva, C. J. 1995. Neotropical floodplains: A comparison between the Pantanal of Mato Grosso and the large Amazonian River floodplains. In: Tundisi, J.G., Bucudo C.E.M., Tundisi T.M. (eds.). Limnology in Brazil. Rio de Janeiro: Brazilian Academy of Sciences, Brazilian Limnological Society p. 195-217. Larson, Adamus and Clairain Larson. 1989. Functional Assessment of Freshwater Wetlands. Univ. of Massachusetts at Amherst & WWF. Miller, R.R. 1982. Pisces. In S.H. Hurlbert and A.Villalobos-Figueroa (eds.). Aquatic biota of Mexico, Central America, and the West Indies. San Diego State University, San Diego, CA. p. 486-501. Mitsch, W.J. and J.G. Gosselink. 2000. Wetlands. John Wiley Motta Marques, D.M.L. da.; and Giovannini, S.G.T. 2000. Hydroperiod influence in the establishment of emergent macrophytes Zizaniopsis bonariensis and Scirpus californicus in experimental constructed wetlands. Verh. Int. Ver. Limnol. 27, December. In press. Padoch, C., J.M. Ayres, M. Pinedo-Vasquez and A. Henderson (eds.) Vrzea: diversity, development, and conservation of Amazonias whitewater floodplains. The New York Botanical Garden. Pringle, CM 1997. Exploring how disturbance is transmitted upstream: going against the flow. Journal of the North American Benthological Society 16:425-438. Pringle, CM, M. Freeman, and B. Freeman. 2000. Regional effects of hydrologic alterations on riverine macrobiota in the New World: tropical - temperate comparisons. BioScience 50: 807-823. Sioli, H. 1984. The Amazon: limnology and landscape ecology of a mighty tropical river and its basin. Dr. J. W. Junk. Dordrecht. Smits, A.J.M., P.H. Nienhuis and R.S.E.W. Leuven (ed.) 2000. New approaches to river management. Backhuys Publishers, Leiden, The Netherlands. Villanueva, Adolfo O. N., David da Motta Marques,and Carlos E. M. Tucci. 2001. The Taim Wetland conflict: a compromise between environment conservation and Irrigation. Water International Journal 25 (4), December. In press. Winemiller, ? and Leslie ?. 1992. Title?? Env. Biol. Fishes 34: 29-50. [Experts, please complete the citation. Thanks.] 5.2. Caribbean Basin small flood plain rivers Compare with other document on Caribbean Basin small flood plain rivers General description and geographic variation Have 1st through 6th order streams; floodplains are not large due to topographic and hydrographic constraints (peak hydrographs are brief, watersheds are small, runoff is highly seasonal, terrain does not provide many places for wide flood retention). Community types/zonation and major gradients within the system (patterns) No information available. Ecological integrity factors for landscape context Table xxx. Ecological integrity factors for Caribbean Basin small flood plain rivers Key FactorsJustification for Factor SelectionEcological Thresholds:(Minimum Integrity Threshold)Justification for Threshold Determination(e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityHydrologyUnimodal annual hydrograph, brief but intense runoff, storm driven rather than persistent rain; some rain events even in dry season; 3-4 m ppt per year, 80% within a 3-5 month season; ground water inputs between storms via slow surface soil drainage, sometimes karst; baseflows much less than 10% of peak flows.Regular 5-year magnitude floods. Larger infrequent floods (e.g., 50 year events).Smaller events allow some maintenance of floodplain and bed texture. Bigger floods needed to maintain floodplain vegetation and morphology.Peak-flow gauging, daily flow gauging.ChemistryHigh solute concentrations (NaCl and P from both geothermal and atmospheric sources), heavily diluted during runoff events, but variable geochemical inputs from one tributary to next. Low natural N inputs (or high uptake and denitrification); wide temperature range depending on channel width and flow. pH highly variable over time and basin due to varying controls.N:P ratio during low flow. Absolute SRP [Experts, please indicate what SRP stands for. Thanks] & NO3-NHighly variable, dependent on geologic context.SRP, inorganic N.ConnectivityLongitudinal matters for diadromous shrimps (Atya spp.) and fishes (catadromous mugilids). Lateral matters for nutrient and sediment and water storage, and for providing refuge and feeding during floods.X% of all headwaters with unimpeded access, distributed evenly across the entire headwater areas of each basin. Free passage of X% of shrimps and fishes past barriers during downstream and upstream migration particularly at night. Lateral connectivity accounted for within hydrology thresholds mentioned aboveDiadromous species need headwaters for feeding, maturation, and reproduction. Organisms must be able to pass major barriers (e.g., dams) longitudinally to complete life cycle Need to ensure accessibility of some minimal proportion of freshwater habitat to account for natural variation in habitat suitability from year to yearTabulation of blockages and diversion structure types from imagery and inventories Censusing Atya spp. and J. pichardi at barriers to confirm free passage during up migration.Biotic zonationAllochthonous, heterotrophic headwaters grade into autotrophic mainstems, Little feeding specialization among fishes except detritivores/grazing plus piscivores; Clear zonation in fish assemblages: headwaters and coastal plains similar, both contrast to high order low gradient (deltaic) fishes that are mostly marine invaders Compositionally, there is a species core and zonal specialistsLess than X% invasive species in total number and total biomass. X% of zonal specialists expected to be present within a given zone at relative abundance that approximates reference conditions. Need to look at possibility of invertebrate zonation and associated thresholds. [Experts, please estimate the value of various X%. If you know of any published studies, please cite. Thanks.]Zonation is a crucial aspect of ecosystem; invasive species such as Tilapia can become high proportion of biomass but it is not clear if there is niche competition from them as well. Need to ensure some minimal richness of fish species as well as ensure minimal representation of particular specialists (mid-reach, high-elevation, and deltaic-marine specialist fishes). Need research to better document invertebrate zonation and see if indexes could be built.Fish and invertebrate census (composition and structure) relative to baseline studies.TurbidityLow during low flow. Peak flows naturally mobilize some sediment (but not a lot). Solute-enriched water has higher natural turbidity; planktonic turbidity only in the lower, flatwater zone.System specific turbidity standards (not greater than X NTUs, [Experts, please estimate the X value, and indicate what NTU stands for. Thanks.] OR X m underwater visibility.Need to prevent low-flow transparency from falling so much that there is a major loss of light penetration to the stream bottom (for photosynthesis and visual feeders)Hydrolab turbidity measurements and/or Secchi disk data.Channel geomorphologyHighest structural diversity in lower-order reaches, maintained by peak events. Woody debris important, increasingly for the higher order streams where it is the major source of habitat structure (whereas in low-order streams geologic structure is also present). High energy flows rework channels and maintain habitat size and structure.Threshold rule would involve characterization of mixture of micro- (substrate and cover) and meso-habitat (riffle, run, pool) types, not just as amounts but their spatial distribution relative to each other.Pool-riffle-run diversity needs to be maintained, as does diversity of microhabitat provided by coarse geologic and woody materials. Different rules for headwater versus middle reach versus low elevation zones.Channel transect data (both lateral and longitudinal). Remote sensing imagery used for mesohabitat analysis.Bed textureSubstrate size or diversity important for invertebrate and fish assemblages in lower order reaches. Sandy or clay bottoms occur in high-order channels. Muddy bottom in some backwaters, near sea, and in lowland small 1st-order streams.Research needed to specify a threshold in terms of particle embeddedness and range of particle sizes (compared to reference levels for different reach types).Fines would be a low proportion of bed materials in headwater and middle reaches, naturally flushed out by peak flows even if there is some natural patchy accumulation during low flows. Pebble counts Sieve samples  Information gaps and caveats Recommended priorities for conservation-driven research agenda and next steps Literature Cited Recommended resources Hurlbert, S.H. and A.Villalobos-Figueroa (eds.) 1982. Aquatic biota of Mexico, Central America, and the West Indies. San Diego State University, San Diego, CA. Whigham, D.F., Dyrygova, D., Hejny, S. (eds.) 1993. Wtlands of the world: Dordrech inventory ecology and management. Kluwer Academic. 5.3. Rain-surface runoff driven riverine system General description and geographic variation This freshwater system type occurs largely in more arid, lower elevation regions where rainfall is the source of stream water and where there is little retention of precipitation on the watershed due to high rates of runoff and high evapotranspiration. The hydrograph in this system type is characterized by runoff events with varying peak discharges, and by low-flows that are significantly lower than these peak flows. Low flows also decline rapidly after peak flows, and base flows are quite low. First order streams may become intermittent during prolonged intervals between rain events; 2nd and higher order streams are perennial. In the Caribbean basin, hurricanes are a significant source of high-magnitude flow events, and have a roughly 3-10-year return interval. In SE Brazil, there are no hurricanes. Other occurrences are likely throughout South America. Presumably species vary regionally, too. In Puerto Rico, the case we discussed in detail, a salt/freshwater shrimp, Atya, plays a key role as primary consumer throughout these rivers. Community types/zonation and major gradients within the system (patterns) There is little differentiation from 1st to larger-order streams in this system type due to the flushing effect of major runoff events, which sweep most aquatic fauna down into refuges either pools or the bottom delta/estuary from which they later re-colonize the entire system. The floodplain plays a crucial role in system function, both as trap (via mechanical filtration and biological uptake) of nutrients and sediment, but is also undifferentiated due to the massive seed dispersal effects of flood events. Microhabitats created in the stream by large woody debris and other major obstacles create the only significant structure differentiating biota along the stream network. Waterfalls in headwaters also present barriers to movements of some aquatic fauna. Ecological integrity factors for landscape context of Rain-Surface Runoff Driven Riverine System (Puerto Rico example) Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold)Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityClimate: specifically temperature, humidity, precipitation regimes. Climate factors are not amenable to direct human control at the site scale but need to be monitored as reference for establishing expectations for other factors, particularly hydrographic factors.N/AN/ABasic weather data on air temperature, air moisture, rainfall (continuous gauging).Hydrologic factors: peak flow magnitude, low flow magnitude, long-term mean flow.The hydrograph is largely rainfall-runoff event driven, with low flows shaped by shallow ground water seepage. Major rainfall events play a crucial role in scouring the system and forcing post-storm recolonization while the persistence of flow between storms provides a continuous river corridor for habitat, feeding, breeding, etc. No intermittent flow other than in extreme headwaters, mean low and high flows not to exceed 10th and 90th percentiles for flow magnitudes, mean flow not to exceed range of middle 50-percentile (25th to 75th) of natural flow. [Experts, please provide literature citation. Thanks.] The biota of this system type are adapted to the predictable occurrence of scouring flow events as well as the persistence of low but continuous flow between such events. Increased size and frequency of peak events might move biota out too often, not allow for full recovery between events, causing simplification of the system. Similarly, loss of continuity during low flows would fragment the system. Lack of gauge data will require putting in gauges, preferably in numerous streams to develop reference data.Peak stage, daily or twice-daily stage.Riparian forest factors, including flooded width, canopy height, species & age compositionFlooding of the riparian zone supports nutrient uptake, sediment filtering, and the release of woody debris and particulate organic matter (POM) back into the stream, where their decomposition provides the crucial base of the heavily heterotrophic aquatic food web.Riparian forest and shrub vegetation must occur across at least the full width of the natural 5-yr flood zone.The mean hurricane return interval in Puerto Rico (the case we used) is roughly 3-10 years. If the riparian forest zone is wide enough that it gets inundated regularly at this return interval, then the system will have its minimum ability to filter nutrients and sediment and generate plant matter for the stream. Note that storms of this size are not sufficient to scour away large amounts of the riparian vegetation, and do not alter the age/size or species composition of this plant assemblage.Measurements from low-expense, low-altitude imagery data.Sediment inputThese systems exhibit naturally low sediment loading due to dense plant cover on their watersheds, despite the high intensity of some rainfall events and the steepness of the watershed slopes in their headwaters. Most fine sediment that is delivered to the stream is also scoured and flushed out by subsequent events. Sediment delivery is associated only with peak flow events and, as noted above, is usually easily flushed from higher gradient reaches by runoff events so long as hydrograph is intact.Instantaneous load not higher than would be associated with a 5-yr storm (e.g., hurricane) event. Annual load (Total Suspended Solids: TSS) does not exceed the Nth percentile of natural variation. [Experts, please estimate the N value. If you know of any published studies, please cite. Thanks.] Such systems should also be managed via setting a threshold that captures the idea that there is no chronic input of sediment outside of higher energy runoff events, but our group could not articulate exactly how this threshold should be stated.Turbidity but only if highly correlated to TSS (varies with hydrogeology, algal activity), rising stage sampling, bed traps.Channel morphology and hydrodynamicsCrucial to habitat structure and hydraulic roughnessVelocity: wetted perimeter relationship does not exceed ? [Experts,please provide examples of ? thresholds from sites you know or from literature. Thanks.]; also pool-riffle-run sequence.Channel transects at varying flows, plus measurements of low altitude imagery.Coarse woody debrisX% of natural mean log density per kilometer. [Experts, please estimate the X value. If you know of any published studies, please cite. Thanks.]Log size and count per distance (sample reaches).Up/downstream connectivityNo barriers to upstream movement of Atya, no uptake into diversions.Count of barriers of different forms. Ecological integrity factors for condition Table xxx. Ecological integrity factors for condition of rain-surface runoff driven riverine system (Puerto Rico example) Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold)Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityBiotic composition: dominant species of Atya, e.g., A. monticolaMinimun population size, access to full migratory range.Nocturnal visual density estimate, trap countsBiotic composition: live-bearing fishesSpecies richness? Biomass?Census methodsBiotic composition: diadromous fishesSpecies richness? Biomass?Census methodsNutrient (N:P, POM) inputLow natural inputs, readily flushed out, diluted, or removed via bio uptake.Chronic (non-storm) input does not exceed natural background by X% [Experts, please estimate the X value. If you know of any published studies, please cite. Thanks.]N, P; Epilithic algal samplers (tile/slide substrate devices) Information gaps and caveats Insects are better studied in these systems than other features. Hydrograph records are rare. Ideas about nutrient dynamics are based on analogies to other systems throughout Central and South America. Life histories of fishes are poorly known. Recommended priorities for conservation-driven research agenda and next steps Literature Cited Recommended resources Angermeier and Karr. 1983. [need to add the title.] Env. Biol. Fishes 9: 117-135. Goulding M., M.L. Carvalho & E.G. Ferreira 1988. Rio Negro: Rich life in poor water. The Hague: SPB Academic Publishing. Hynes, H.B.N. 1975. The stream and its valley. Verh. Int. Ver. Theor. Ang. Limnol. 19:1-15. Miller, R.R. 1982. Pisces. Pages 486-501 in S.H. Hurlbert and A.Villalobos-Figueroa (eds.) Aquatic biota of Mexico, Central America, and the West Indies. San Diego State University, San Diego, CA. Pringle, C. M. 1997. Exploring how disturbance is transmitted upstream: going against the flow. Journal of the North American Benthological Society 16:425-438 Pringle, C. M. 2000. Threats to US public lands from cumulative hydrologic alterations outside of thier boundaires. Ecological Applications 10:971-989 Pringle, C. M., M. Freeman and B. Freeman, 2000. Regional effects of hydrologic alterations on riverine macrobiota in the New World: Tropical - temperate comparisons. BioScience 50:8078-823 Sioli, H. 1984. The Amazon: Limnology and Landscape Ecology of a mighty tropical river and its basin. Dr. J. W. Junk. Dordrecht. Vannote, R. et al. 1980. The river continuum concept. Can. J. Fish. Aq. Sc. 37: 130-137. Winemiller and Leslie. 1992. [need to add the title.]. Env. Biol. Fishes 34: 29-50. 5.4. Karstland subterranean freshwater system General description and geographic variation These systems exhibit high degrees of endemism due to their different histories of colonization; any carbonate system can support an underground freshwater system given appropriate hydrologic conditions and each cave system. [Experts, please add a paragraph that describes the karstland subterranean freshwater system or cite a reference so that I can add the description later. Thanks.] Community types/zonation and major gradients within the system (patterns) No information available. Ecological integrity factors Table xxx. Ecological integrity factors for Karstland subterranean freshwater system Key FactorsJustification for Factor SelectionEcological Thresholds:(Minimum Integrity Threshold)Justification for Threshold Determination(e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityHydrology: magnitude of low water levelsHydrograph has two critical features that define the habitat: Pulse flow from surface precipitation events that drain directly into fracture and sinkhole features, coupled with baseflow from slow infiltration and release of storage from small fractures after storm events. Biota are adapted to hiding to avoid being dislodged during peaks of hydrographs, but are sensitive to reduction in baseflow levels as this causes changes in available habitat.Case-specific low water elevation cutoff based on when water levels fall below some elevation where hydrologic connectivity among caves or between caves and surface is lost.Hydrologic connectivity between caves or from caves to surface depends on water table level establishing these connections.Nutrients and other chemicalsSystem based on inputs of POM and DOM [Experts, please indicate what DOM stands for. Thanks.] from land surface; exact type probably does not matter as microbes and micro-fauna adapted to consuming anything, and rest of biota consume these smaller biota. Simplicity of system does make it especially vulnerable to poisoning. N or P or altered DOM/POM enrichment would alter food web by making system more habitable by non-stygo-fauna that could compete better in less oligotrophic conditions.X% increase in annual input of POM and DOM. [Experts, please estimate the X value. If you know of any published studies, please cite. Thanks.]Systems are very sensitive to pollution from excessive organic matter input and also poisoning. Exactly how much change in POM or DOM will depend on each case.[Experts, please give a few examples. Thanks.]Biota: each macroscopic species as its own factorCave aquatic biota are so rare, and the food web so simple that tracking the higher levels of the food web (fishes, crustaceans, molluscs) provides info on status of entire food web.10% or greater change in census for top level fauna.Given natural stability of inputs, would expect very stable native populations. As a result, any deviation from natural numbers would be suspect.SedimentFauna probably naturally adapted to seeking refuge during high flow events when sediment would be most likely to be present in water. However, sedimentation into sink-holes and other recharge features could alter hydrology. Further, excess accumulation of sediment in caverns and fractures could eliminate habitat.Any accumulation at select points in system would be signal of trouble.Inorganic chemistryEffects of pH of incoming water on carbonate rock define the natural inorganic chemistry of these systems. Alteration of input pH could change saturation levels for Ca and Mg.Assuming high stability, any change in baseflow pH would be suspect. Information gaps and caveats Recommended priorities for conservation-driven research agenda and next steps Literature Cited Recommended resources 5.5. Glacial source streams General description and geographic variation All glacial headwaters in the Andes start at high elevation (5-6,000m). There are three basic types: 1. High gradient eastern draining streams that join larger systems with other water sources at lower elevation. These larger systems are not dominated by glacial source stream characteristics. 2. High gradient, relatively short western draining streams that are glacial dominated throughout, and drain into the Pacific ocean. 3. High gradient streams that merge into larger, low gradient streams in inter-Andean valleys (1,500m elev.). These streams then become high again after leaving the inter-Andean valley. Community types/zonation and major gradients within the system (patterns) Type 1: Major zones are high elevation (>5,000m)? Paramo? Cloud forest? Low elevation? [Experts, please describe other characteristics of Type 1 in addition to "high elevation," and formulate your questions for several "?." Thanks.] Type 2: Major zones are same as type 1? [Experts, please indicate the Type 2 characteristics that are different from Type 1. Thanks.] Type 3: Major zones are: high elevation, high gradient, low gradient, moderate elevation in inter-Andean valleys, and high gradient, lower elevation. Biotic community zones include the high elevation zone which is depauperate, moderate elevation zone which has native catfish and duck species, amphibians, crustaceans, mollusks and other macroinvertebrates. Lower elevation zone has fishes (?) and dipper bird species. Ecological integrity factors for landscape context Table xxx. Ecological integrity factors for landscape context of glacial source streams Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold) Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityHydrologic regimeBiota specialized for low inter and intra-annual flow variation, and diurnal pulses.Minimum and maximum daily mean flows and diurnal mean and maximum values. Must not chronically exceed minimum lows or maximum highs.Variation is minimal.Water level gauge1Turbidity regime Diurnal patterns of daytime milky conditions and nighttime clear conditions limit primary productivity and maintain food web structure.Daily patterns of change, 80% of night clarity, 80% of opaqueness during daytime.Initial estimates (need to evaluate).Secchi disc, or Transmissometer1SedimentationGenerally low, but increased sedimentation levels in valleys.Need to look at daily averages, set maximum chronic levels since natural phenomenon produces acute high sediment episodes.Levels associated with natural range of variation (daily low mean without episodic events).3Solar radiation levels in cloud forest sectionSolar radiation causes temperature increase and also UV exposure. Cloud forest destruction may limit cloud production and cause harm to native biota (e.g., Amphibians).Need to establish natural UV light levels.Indicates how cloud forest destruction is affecting aquatic biota.Film exposure monitoring, or regular photometer readings.3 Ecological integrity factors for condition Table xxx. Ecological integrity factors for condition of glacial source streams Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold) Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityExotic fish The presence of native ducks and fish species is negatively correlated with the presence of brown and rainbow trout.Presence or absence of exotic species.Indicates the highest biological impact.Seining, backpack shocking.1Native biotic compositionNative catfish, invertebrate-eating ducks, dipper birds and amphibians.Critical population densities and ranges need to be established. Indicate most critical conditions.Seines, electroshock, bird counts1Temperature Cold or cool water is necessary for native biota. It also limits the invasibility of exotic species.High elevations: 8-11C0 Cloud forest: 10-13 C0 [Experts, please provide literature citation. Thanks.Natural range of variationTemperature gauge, or regular temperature readings.2Native vegetation in the inter-Andean valley Species rich, open shrublands and/or woodlands, humid or dry depending on the valleyNeed to estimate natural cover types and % of cover. Need to work with vegetation experts who know the areas that have remnant natural vegetation. Other methods to identify potential natural vegetation.3 Information gaps and caveats Need information on lower elevation zones of types 2 and 3. Recommended priorities for conservation-driven research agenda and next steps Identify best remaining inter-Andean systems and identify impact thresholds for biota. Literature Cited Recommended resources Horn, S. P. and K. H. Orvis. 1999. Investigacin Limnologca y Geomorfologca de Lagos Glaciares del Parque Nacional Chirripo, Costa Rica. Revista Informe Semestral, Vol. 35: 95-106. Chapter6Marine habitat types 6.1. Coastal lagoons General description and geographic variation No information generated. Community types/zonation and major gradients within the system (patterns) No information generated. Ecological integrity factors for condition Table xxx. Ecological integrity factors for condition of coastal lagoons Key FactorsJustification for Factor SelectionEcological Thresholds: (Minimum Integrity Threshold) Justification for Threshold Determination (e.g., Natural Range of Variation)Indicators for Field-Based MonitoringFactor PriorityBacteriological quality of the water The general health of a lagoon depends greatly on anthropogenic impacts. These are clearly expressed in the coliform concentration. Review FDA [Experts, please write out the full name of FDA. Thanks.]FDA norm for environmental health. Shellfish Sanitation Program Manual, FDA.Total coliforms Nutrients: N & P An excess of nutrients easily breaks the equilibrium of the system, leading to eutrophication. Varies by siteHigh and persistent nutrient {concentrations} lead to eutrophication, degrading the system. Low {concentrations of} nutrients reduce the carrying capacity of the system.O2 A reduction of the concentration of O2 can break the equilibrium of the system. Associated with over-saturation of the carrying capacity and an elevated time (t) of residence in the water. Varies by site. A prolonged period of less than 50% O2 saturation is risky. A low and persistent O2 saturation is associated with eutrophication of the system. Tends to be associated with a high T (Temperature) and C. [Experts, please indicate what C stands for. Thanks.]% of O2 saturation. (Seasonal; higher t in summer)Coverage of seagrass beds (ej. Zostera) The abundance and condition of the seagrass beds {literally "prairie"} is the essential support of the system. Reflects the general health of the system. Its reduction diminishes the biological richness of the system, {and} the carrying capacity. Is a source of productivity and has homeostatic functions: filter, stabilizer, substrate.negative tendency In San Quintin, Baja California, one % of the 15% indicates good condition. [Experts, please provide literature citation. Thanks.] % of coverage (annual) in space: map or satellite image. Comparative analysis study of diverse lagoons. Geomorphological (conservation)Human alterations (marinas, wave-breakers, ports, piers, etc.) radically modify the hydrology of the system, changing all its equilibrium.Continuos observation of the coastline and monitoring projects.Development in neighboring zones The eternal ?? {A word is missing from the original. It probably was "development"} directly affects the conservation of a lagoon. The proximity of settlements, industry, agriculture, tourism, etc., compromises its conservation.Appropriate distance (buffer zone) between the lagoon and intensive development. The immediate proximity of the system and the disturbance leads to the deterioration of the system.Study: development of models and carrying capacity of particular lagoons, analysis of the trophic web.DischargesThese are point sources of contamination (organic, coliforms, agricultural, industrial, etc.).In F [Experts, please indicate what F stands for. Thanks.] {In function} of the dilution capacity, which tends to be low for organic and none for industrial.Discharges should be avoided.Identification of species, density and biomass, distribution, vegetation maps.Abundance of green algae (Enteromorpha y Ulva)Indicates eutrophication of the system; excess of nutrients due to drainage.Develop an indicator. Information gaps and caveats Recommended priorities for conservation-driven research agenda and next steps Literature Cited Recommended resources Bardach JE (ed). 1998. Sustainable Aquaculture. John Wiley & Sons. New York, USA. Clark JR 1996. Coastal Zone Management Handbook. Lewis Publishers. Boca Raton, USA. [Experts, please add more reference citations. Thanks.] 6.2. Coral reefs General description and geographic variation No information generated. Community types/zonation and major gradients within the system (patterns) No information generated. Ecological integrity factors Tropical marine Sseagrass beds, coral reefs, coastal lagoons, estuaries, mangroves forests, and associated wetlands and coral reefs are functionally connected integrated in many areascreating an ecological mosaic matrix system. There are also functional relationships within the coastal zone that link land and sea. Integrate land and sea ecologically. We identified three types of key factors: First order determinants: Primary key factors (e.g. physiography, hydrologic regime, and currents) These primary key factors must be maintained determine the most vigorous or absolute viability (existence of the system) within their ranges of natural variability. It is not appropriate to discuss thresholds for these factors. We need to understand what maintains the integrity without the physiographic or hydrological thresholds. At first, it is not important to identify thresholds. However, it is necessary to deal that to support the complete integrity without thresholds of the fisiografa and of the system hidrolgico (current). En el primer orden que marca la viabilidad ms rigurosa, no es pertinente hablar de umbrales. Es necesario entender que mantener la integridad completa sin umbrales de la fisiografa y del sistema hidrolgico (corrientes).It is important to understand what maintains the complete integrity Second order determinants Secondary key factors (environmental determinants): Among the many environmental determinants, we analyzed the following factors that are common to the systems. Penetration of light / depth Temperature Salinity It was relatively easy to identify agree upon the scope (variability) of these determinants because they are widely reported in the literature. (No particular citations listed) Third order determinants Tertiary factors (principally biotic factors): The determinants of the third order were principally specific to the scale of site and the surrounding seascape. We identify the following factors that are common to the systems that we analyzed.as important to the analyzed systems: Percent Live Substrate Cover Level of Herbivory Recruitment and biological connectivity Distribution and abundance of associated flora and fauna Nutrient levels in marine waters Degree of Turbidity and Sedimentation Ecological integrity factors for landscape context Table xxx. Ecological integrity factors for landscape context of coral reefs PRIVATEKey factorsJustification for the Selection of the FactorMinimal Integrity ThresholdsJustification for the determination of the threshold (natural Range of variation)Indicators for field monitoringTurbidityTurbidity limits the development coral communities by blocking sunlight needed for metabolic processes within the coral colonies. It will ultimately have ramifications for coral species diversity, abundance, and distribution.Threshold 100 mg/l (check limits in Rogers' manual, 1990) Production of mucus at certain levels (look for details on in Rogers or another work) Thresholds are well documented in scientific literature. It is useful to consider turbidity together with salinity and temperature.Seasonal variability of sedimentation and turbidity;; contribution of sediments using the universal soil loss equation ( To see Hallock, Muller-Karger and You Find, 1993, Natl. Geog. Beast. And Explor.); use of Secchi disk; aerial images, etc.TemperatureTemperature, together is one of the critical factors regulating the survival of coral reef species and the functioning of the coral reeff ecosystem. Temperature affects biological processes including growth, reproduction, symbiosis, metabolic rate and reproductive capacity of individuals. Threshold: 24 30 C (on range extremes there are species that tolerate temperatures up to 20; Temporal extreme changes in temperature occur and if limited species will survive.Thresholds are well documented in scientific literature Also it is known that the warming of the seas is seriously affecting the viability of coral reefs worldwide. Consider the synergystic effects of temperature together with salinity and turbidity.Determination of thermal (spatial and temporal) regimes using a variety of techniques according to resources and specific conditions. SalinitySalinity, alone and in combination with other physical-chemical aspects, is a key factor in the functioning of the coral reef. Salinity affects biological processes (e.g. growth, reproduction, symbiosis, metabolic rate and reproductive capacity of the organisms, etc.). Threshold 26 - 36 ppm [Experts, please provide literature citation. Thanks.] Consider that there are species that tolerate ends of salinity extremes Consider salinity together with temperature and turbidity..Determination of salinity regime of the reef, with a design that considers the spatial and temporal variation and the proximity of sources of fresh water, depth, precipitation monitoring stations and drought, etc. Generally refractometers are used in situ to take measurements, or they can take water samples to the laboratory for analysis.NutrientsCoral reefs ecosystems exist in an oligotrophic environment. Increased contribution of nutrients for medium or long term duration favors the growth of benthic and planktonic algae and diminishes the coverage of live coral through direct and indiect competition.Nitrogen and phosphorus; information exists in Hatcher, 1990 and also Clark, 1996. [Experts, please complete the citation in the References section. Thanks.]Thresholds are documented in scientific literatureMeasure the patterns of spatial and seasonal distribution chlorophyll (concentrations) and nutrients using different methods, including the use of satellite images and measurements in situ.Recruitment and biological connectivityMaintenance of genetic flow among adjacent populations contributes to the maintenance and/or recovery of the structure and function of the reef following disturbances. Due to larval dispersal by the majority of shallow water tropical marine species, the viability of the populations depends on the contribution of larvae and juvenile of nearby or distant sites, that are connected by oceanic currents. There is no a number for this factor Specific information will be needed for each situation. Thresholds ?? Integrety will be determined by healthy source populations & appropriate currents. In marine organisms it is difficult to know the minimal viable size of a population to guarantee a suitable supply of larvae. Nevertheless, information exists for some species (related to the cycle of life for particular taxonomic groups and their habitat requirements through development, and for the capacity larval distribution over different geographic scales) [Experts, please provide literature citation. Thanks.]Large, healthy and widely distributed populations of coral reef flora and fauna provide for the long-term survival of coral reef ecosystems. This occurs on local and regional scales, depending on the particular geographical area. Determine the influence of oceanic currents in the region. Estimate the degree of connectivity by determining the genetic distance of adjacent populations in different species. Maps of reefs of the region with information about coral coverage, diversity of species, and coral health. Comparative analysis of conditions in protected areas v. not protected in each region.Influence of currentsCurrents provide pathways for genetic flow among populations for dispersion of the larvae, and distribute nutrients and pollutants.There is no known threshold. Just known to be a critical energy input in coral reef ecosystems. Develop ad hoc.Knowledge of the general influence of regional and local currents and their role in the transport of larvae of certain species and other factors will help identify the patterns of currents to be maintained or at least monitored. Speed and direction of the currents (especially at important events like seasonal spawning of ish and invertebrate species The methodology is complex and needs special equipment including current meters, satellite images, etc.Note: For monitoring, some of the key factors, it is important to take in account spatial and seasonal variation in order to have representative information. The guidance given by Rogers et al., (1994) is recommended. For monitoring currents, consider Doppler (ADCP), consult Joaqun Buitrago for more information. Ecological integrity factors for condition Table xxx. Ecological integrity factors for condition of coral reefs PRIVATEKey factorsJustification for the Selection of the FactorThreshold (is) of Minimal IntegrityJustification for the determination of the threshold (natural Range of variation)Indicators for field monitoringRelative abundance and diversity of carnivores and key herbivores (total biomass and number of species), i.e. complete or fragmented trophic structure in terms of species and total biomassThe relative proportion of carnivores:herbivores support the health of the system and its diversity/complexity, and make it less vulnerable to changes of conditions (e.g.. fish defecate and fertilize, herbivores change coverage of algae)There is no general formula. Determined ad hoc. Some examples include the species of fish in question presence/absence of some species of limited abundance species (tortoises, and manatees) Evaluate the information by comparing with ecologically similar areas. Consider different states known due to exploitation, and historical trends of population decline and deterioration of the coral community. Composition measures will be will be the basis for comparative analysis that determines the viability of local communities. The assumption is that at some predator:herbivore composition measure degradation in the structure and functioning of the coral communities will occur.Censuses of abundance, biomass and composition of species inside taxonomic groups (e.g. the family Serranidae), herbivorous fish (Scaridae, etc.) fish coralvoros (Chaetodontidae), scrapers (Acanthuridae) and others etc., Counts of tortoises and manatees.Reproduction of the organisms of the reef community.Guarantees the survival of the reef community.Adequate sex ratios and abundance of the breeding population of selected species. The values of these proportions are specific for certain groups of organisms. Consult the literature. [Experts, please provide literature citation. Thanks.]It guarantees the degree of recruitment necessary to assure the maintenance or growth of viable populations.Censuses of abundance and recruitment, population structure and breeders determined by different methods, according to the organisms. (E.g. Vertical composition, seasonal variation of the proportion for sexes and of the proportion of sexually mature individuals, abundance and vertical structure (depths) of the reproductive aggregations; Census of juvenile and recruitment in spawning or nursery areas.) Rate of growth of corals The rate of historical and current growth indicates the form and conditions under which the coral has developed, and the nature of the reef formation.Thresholds for the rate of growth are defined by analyzing rates from previous decades/centuries. Those measurements of can be used to develop natural ranges of variation. The trend of the health of the reef community is reflected in the chronology of growth over a centurys time. The threshold will indicate the pace of natural growth of corals in past decades or centuries. This will present a better picture of the natural variations of growth, and provide a maximum potential growth rate under some environmental conditions. Growth of new mass must exceed rate of bioeroisnInvestigation: Historical analysis of coral growth (sclerochronology). Monitoring: Annual monitoring is recommended with the same methodology if change is expected to occur quickly.Level of herbivoryRegulates the abundance of algae and allows the recruitment of new individuals of the species of attached/sessile fauna of the reef. Excess herbivory can affect the rate of coral reef bioerosion. Suitable abundance of sea urchin Diadem: 30-70/m2. Abundance of herbivorous fish according to species (see literature for estimates of adequate levels)..A balance must exist among the processes of erosion and accretion of the corals and a drastic reduction of the herbivorous organisms provokes the increase of algal coverage and the suffocation of the corals. The system shifts to one dominated by algae. In the Carib, the massive mortality of sea urchins Diadem at the beginning of the 80s allowed an increase of the coverage algae in many reefs. In the Pacific Ocean, the increase of algal biomass because of ENSO [Experts, please write out the full name of ENSO. Thanks.] resulted in an excessive increase in the populations of sea urchins (Diadem) and herbivory resulted in considerable bioerosion of coral reef formations. Censuses of abundance of species of herbivores including both vertebrates and invertebrates.Coverage of living coral Coral are the principal builder of the reef and the base of the ecosystem. Therefore, it is important to maximize the quantity of living coral.For the Caribbean a reef with coverage of more than 40 % is considered in good condition. In the Pacific Ocean, due to the massive mortality of corals caused by the ENSO there is now an estimated average of 25 % live coral coverage. Greater than 60 % coverage of macro algae represents serious degradation of the systems.Information from 1950-60 indicates that in the Caribbean sea a healthy reef had more than 60 % of live coral coverage. Information from 1960 indicates that in the Pacific Ocean the reefs had a coverage of 90-100 %, but now the average is only 25 %. To preserve coral reefs it is recommended to preserve what currently exists without a focus on minimal percentage of coverage. The viability information is insufficient.Measurements of the live coral coverage, sessile fauna and algae (carpet of macro algae) by means of transects in situ. Information gaps and caveats We do not know much about the natural states of coral reefs, because the studies focused on the monitoring of coral reefs began in the 1950s by thenwhich time they had already been altered by human activities. The experts consider it risky to establish thresholds for ecological factors for systems about which we know so little about the range of natural variability. In addition, for some physical-chemical factors, the thresholds can change due to synergistic influences among the factors themselves. The viability of the marine systems depends on large scale spatial and temporal processes. These processes require functioning seascapes at scales far beyond the reef itselfareas at which complex regional process occur. Therefore networks of conservation areas are needed to effectively conserve the coral reef ecological system. More studies about genetics are needed. (e.g. Genetic distance of adjacent populations of some species and mapping the diversity and coral coverage of the reefs of the region to determine the nature and controlling factors for connectivity. More studies of composition and rates of recruitment to understand better aspects of landscape context and the condition of the reefs. We rRecommended to focus on the following coral genera: focal species of coral : Acropora, Montastrea, Porites and Agaricia in the Caribbean; Pocillopora, Porites and Pavona in the Pacfic; Siderastrea and Mossusmilia in coastal Brazil; and fish species of the following ones two families of fish, Serranidae (grouers) and Lutjanidae (snappers), whose abundance in the coral reefs has diminished in many areas due to over-fishing. Recommended priorities for conservation-driven research agenda and next steps Continue to analyze ecological integrity of marine systems in detail. Ensure that conservation area managers or supporters of conservation of coral reefs have the necessary information and capacitybilities necessary to apply the knowledge we assembled at the Workshop.have available now. Develop an analytical process similar to the Herndon viability workshop in different Latin American countries to define strategies focused on decision makersin particular situation we may need to analyses that include economic, political and cultural contexts in addition to environmental aspects. Construct a paradigm framework with a focused vision for coral reef conservation in Latin America that incorporates the regional locally applicable economic, social, political and cultural factors. To aAchieve this goal, we propose by proposing concrete action plans of action. Literature Cited Clark, 1996, on page 5. Hallock, Muller-Karger and You Find, 1993, Natl. Geog. Beast. And Explor., on page 4. Hatcher, 1990, on page 5. Rogers's manual, 1990 on page 3 Production of mucus (to look for number in Rogers or another work), on page 3. us (to look for number in Rogers or another work), on page 3. Antonius, A. y E. Ballesteros. 1998. Epizoism: A new threat to coral health in Caribbean reefs. Revista de Biologa Tropical. 46 (5): 145-156. Clark, 1996, on page 5. Doppler, Monitorear corrientes, ver Acoustic Doppler Current Profiler (ADCP) consultar a Joaqun Buitrago para ms informacin. [Experts, please complete the citation, Thanks. Please see page 6 Notas.]. Hallock, Muller-Karger y Hallas, 1993, Natl. Geog. Res. and Explor., on page 4. Hatcher, 1990, on page 5. Hill, M. S. 1998. Spongivory on Caribbean reefs releases corals from competition with sponges. Oecologia, 117(1-2):143-150. Meylan, A. 1988. Spongivory in hawksbill turtles: A diet of glass. Science, 239:393-395 Rogers C.S. 1985. Degradation of Caribbean and western Atlantic coral reefs and decline of associated fisheries. Proc. 5th Int. Coral Reef Congr 6: 491-496. Rogers, C.S. 1990. Responses of coral reefs and reef organisms to sedimentation. Mar Ecol. Prog. Ser. 62: 185-202. Rogers, C., Garrison, G., Hills, Z., and Franke, M. 1994. Coral Reef Monitoring Manual for the Caribbean and Western Atlantic. U.S. National Park Service. Manual de Rogers, 1990 on page 3 Produccin de mucus (buscar cifra en Rogers 1990 u otro trabajo), on page 3. Recommended resources Birkeland, C. (ed.) 1997. Life and Death of Coral Reefs. Chapman and Hall, USA 536 p. Doppler, Monitorear corrientes, ver Acoustic Doppler Current Profiler (ADCP) consultar a Joaqun Buitrago para ms informacin.( Monitorear currents, to see Doppler (ADCP) - to consult Joaqun Buitrago for more information.) [Experts, please complete the citation, Thanks. Please see page 6 Notas.]. Rogers, C., Garrison, G., Hills, Z., and Franke, M. 1994. Coral Reef Monitoring Manual for the Caribbean and Western Atlantic. U.S. National Park Service. Westmacott, S., K. Teleki, S. Wells and J. West. 2000. Management of Bleached and Severely Damaged Coral Reefs. Information Press, Oxford. IUCN, Gland. 36 p. Wilkinson, C. (ed.) 2000. Status of Coral Reefs of the World: 2000. GCRMN/AIMS, 363 p. 6.3. Seagrass beds General description and geographic variation Caribbean: Sea grasses are widely distributed. Tropical Pacific: Sea grasses exist mainly at the river mouths. Community types/zonation and major gradients within the system (patterns) No information generated. Ecological integrity factors for landscape context Table xxx. Ecological integrity factors for landscape context of seagrass beds Key FactorsJustification for factor selection Minimum Integrity Threshold(s)Justification for determining the threshold (e.g., Natural Range of Variation)Field-based monitoring indicatorsFactor PrioritySunlightA system dominated by phanerogams needing abundant light for photosynthesis.Search information in literature [Experts, please provide literature citation. Thanks.] for grasses dominated by Thalassia, Syringodium, Zostera, Rupia, etc.Plant growth decreases under certain thresholds. The degree of sunlight determines vertical distribution.Water turbidity measured with a Secchi disk, turbidometer, etc.SalinityIt defines community structure and grassland function.Specific thresholds for each community type (Zieman, J.; see the CARICOMP manual in the West Indies University web page.) [Experts, please complete the citation in the References section. Thanks.]Salinities under and above the thresholds affect the structure and function of these systems, causing epiphytism, disease, growth and distribution changes.Determination of medium salinity regimes with refractometer.Sediment and water chemistry (organic matter and nutrient contents, and pH levels in sediments)Maintain structure, function and productivity of the communities. A serious imbalance can cause eutrophication . For references, see Zieman and Hector Guzman, Daisy Garcia (INTECMAR, Venezuela).Values under and above the thresholds affect the structure and function of these systems, causing epiphytism, disease, growth and distribution changes.Determine organic matter contents in sediment (total P, ammonium, total N), pH of the sediment. (see Zieman [Experts, please complete the citation in the References section. Thanks.] for methods to measure factors.)Water dynamics (waves, currents, tides, storms) It determines vertical and horizontal distribution of the seagrass bed and its community structure and biomass.Thresholds for these factors are specific and known for prairies and semi-closed systems (see Hedgepeth, Guadalupe de la Lanza, Alfonso Aguirre will provide information on this). [Experts, please complete the citation in the References section. Thanks.] In general, factors should provide water flow dynamics that ensure systems viability.Time of water permanence in semi-closed places (days) (see Lawrence Mee). [Experts, please complete the citation in the References section. Thanks.] Coastal geomorphology (sediment granulometry, physiography, bathymetry).It determines physical structure of the system, which in turn determines vertical and horizontal distribution of the seagrass bed and its community structure and biomass.There is no threshold for geomorphology. Granulometric composition and dynamics of the sediment are specific for each site.Its recommended to avoid any modification of the original physiographic structure. For instance, avoid dredging, filling, accumulation of excessive sediment load. (see the CARICOMP manual in the West Indies University web page.)Genetic connectivity.Consult J. Zedler, Univ. of California. [Experts, please provide J. Zedlers e-mail address or cite Zedlers publications in the References section. Thanks.] Ecological integrity factors for condition Table xxx. Ecological integrity factors for condition of seagrass beds Key FactorsJustification for factor selection Minimum Integrity Threshold(s)Justification for determining the threshold (e.g., Natural Range of Variation)Field-based monitoring indicatorsFactor PriorityHerbivory Fundamental for maintaining primary and secondary productivity of communities by controlling epiphytism, nutrient recycling and export of material to adjacent systems.Information from experts is not enough to establish thresholds, but its known that high levels of diversity and abundance of herbivores (turtles, fish, manatees, birds, and invertebrates such as sea urchins, crustaceans, mollusks, etc.) SENTENCE INCOMPLETE IN THE SPANISH Thresholds exist for epiphyte cover on phanerogams (see specialized literature: Jackson ? 1997, Bjorndal 1982] [Experts, you are welcome to add more literature citations. Thanks.]The decrease of herbivore density causes an increase of epiphytism, affecting growth and causing death to phanerogams. Herbivory intensity can be measured through leaf covering by epiphytes. Study of herbivore feeding habits through stomach contents analysis, observations of frequency of bites on fish, vegetation clearings caused by turtles, etc. Associated fauna and flora (residents and visitors).Presence and diversity of species and functional groups and abundance of associated fauna and flora ensures ecological system integrity.[Consult Alejandro Arrivallaga, USGS Louisiana; Silvia Ibarra, CICESE, and J. Zieman, at the Univ. of Virginia; J. Zedler, Univ. of California; Jorge Herrara, CINVESTAV, Mrida] [Experts, please complete the citation in the References section. Thanks.] Information gaps and caveats Recommended priorities for conservation-driven research agenda and next steps Literature Cited Alejandro Arrivallaga, USGS Louisiana, on page 5. Bjorndal. K.A. 1982, The consequences of herbivory for the life history pattern of the Caribbean green turtle, Chelonia mydas. In: Bjorndal K.A. (ed) Biology and conservation of sea turtles. Smithsonian Institution Press. Washington, D.C. pp 111-116. Hedgepeth, Guadalupe de la Lanza, Alfonso Aguirre will provide information on this, on page 3. Herrara, Jorge CINVESTAV, Mrida, on page 5. Ibarra, Silvia. CICESE, on page 5. Jackson, ? 1997. Reefs since Columbus Coral reefs 16, Suppl. S23-S32, on page 5. CARICOMP manual on West Indies University web page, on page 2 and page 4. Mee, Lawrence on page 3. Zedler, L. Univ. de California, on page 4. Zieman, J., specific thresholds for each community type, on page 2. Zieman, J., at the Univ. de Virginia; on page 5. See Zieman y Hctor Guzmn, Daisy Garca (INTECMAR, Venezuela) for references, on page 2. See Zieman for methods of factor measurement, on page 2. Recommended resources Clark JR 1996. Coastal Zone Management Handbook. Lewis Publishers. Boca Raton, USA. Dixon, Scura & vant Hof 1993. Ecological and Economic Goals: Marine Parks in the Caribbean. Ambio, vol 22 (2,3): 117-125. Flores Hernandez. et al ..? other authors names? (1997). Anlisis y Diagnstico de los Recursos Pesqueros Crticos del Golfo de Mxico. Publisher? [Experts, please complete the citation. Thanks.] Annex 1 Workshop contributors Lowland moist non-flooded broadleaf forest Rodolfo Dirzo Robin Foster David Oren Wendy Goyert (recorder) Kit Kernan (facilitator) Xiaojun Li (TNC staff) Lowland moist flooded broadleaf forest Eduardo Asanza Peter Esselman Katie Frohardt Wolfgang J. Junk David Marques Alonso Ramirez Carolina da Silva David Braun (facilitator & recorder) Jonathan Higgins (facilitator) Orlando Rangel (review the 1st draft document) Mark Breyer (TNC staff) Tarcisio Granizo (TNC staff) Pine/pine-oak/oak forest Tim Fahey Ricardo Grau Alejandro Hernandez Sally P. Horn Maarten Kappelle Maximina Monasterio Jesus Orlando Rancel Margaret Stern Kenneth R. Young Roger Sayre (recorder) Jerry Touval (recorder) Pat Comer (facilitator) Hugo Arnal (TNC staff). Montane cloud forest Tim Fahey Ricardo Grau Alejandro Hernandez Sally P. Horn Maarten Kappelle Maximina Monasterio Jesus Orlando Rangel Margaret Stern Kenneth R. Young Roger Sayre (Recorder) Jerry Touval (Recorder) Pat Comer (Facilitator) Hugo Arnal (TNC staff). Dry forest Mary T. K. Arroyo Mauricio Cortero Gordon W. Frankie Carmen Josse Darin E. Prado Andrew Taber Guadalupe Morales (recorder) Susan Anderson (facilitator) David Mehlman (TNC staff) Agnes Velloso (TNC staff) Savanna Nelson Agudelo Stephan Beck Jeanine Maria Felfili Otto Huber Ron Myers Vania Pivello Shirley Keel (recorder) Rafael Calderon (lst Day facilitator) Laurenz Pinder (2ed Day facilitator) Montane grassland (pramo and puna) Stephan Beck Tim Fahey Ricardo Grau Alejandro Hernandez Sally P. Horn Maarten Kappelle Maximina Monasterio Jesus Orlando Rancel Margaret Stern Kenneth R. Young Roger Sayre (recorder) Jerry Touval (recorder) Pat Comer (facilitator) Hugo Arnal (TNC staff) Hot xeric systems Mary T. K. Arroyo Mauricio Cortero Gordon W. Frankie Carmen Josse Darin E. Prado Andrew Taber Guadalupe Morales (recorder) Susan Anderson (facilitator) David Mehlman (TNC staff) Agnes Velloso (TNC staff). Mangroves Pendiente Large wetland systems Pendiente Caribbean Basin small flood plain rivers Peter Esselman Alonso Ramirez David Brown (facilitator & recorder) David Benda (recorder) Rain-surface runoff driven riverine system Pendiente Karstland subterranean freshwater system Pendiente Glacial source streams Pendiente Coastal lagoons Alfonso Aguirre Joaqun Buitrago Georgina Bustamante Jorge Corts Domingo Flores Hector Guzmn Patrcia Monteiro Cunningham Yara Schaeffer-Novelli Nestor Windevoxhel-Lora John Tschirky (recorder) Cristina Lasch (facilitator) Coral reefs Alfonso Aguirre Joaqun Buitrago Georgina Bustamante Jorge Corts Domingo Flores Hector Guzmn Patrcia Monteiro Cunningham Yara Schaeffer-Novelli Nestor Windevoxhel-Lora John Tschirky (Recorder) Cristina Lasch (Facilitator) Seagrass beds Alfonso Aguirre Joaqun Buitrago Georgina Bustamante Jorge Corts Domingo Flores Hctor Guzmn Patrcia Monteiro Cunningham Yara Schaeffer-Novelli Nestor Windevoxhel-Lora John Tschirky (recorder) *+1[\    * + , F Y  ) ? 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