The ecology of tropical montane cloud forests: a global synthesis
Research Article
Towards integrated ecological research in tropical montane cloud forests
- Patrick H. Martin, Peter J. Bellingham
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- Published online by Cambridge University Press:
- 13 September 2016, pp. 345-354
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Tropical tropical montane cloud forests (TMCFs) cover a small portion of the Earth, yet they are significant biodiversity hotspots and centres of endemism, and they provide important hydrological and biogeochemical functions that affect human livelihoods. Given their fundamental sensitivity to climate, TMCFs also serve as an early warning system for climate change impacts. This paper outlines a new international initiative, CloudNet, that aims to promote integrated research across TMCFs, and introduces a special issue that reviews emerging themes and topics in the ecology of TMCFs, highlighting knowledge gaps and suggesting new directions for research. CloudNet is helping coordinate several new research projects and protocols: (1) a global repository of TMCF data and meta-analyses across multiple sites; (2) a multi-site study of plant functional traits across TMCFs; (3) a multi-site study of decomposition processes across TMCFs; (4) a protocol for standardizing climate data collection across TMCFs. These studies are intended to evaluate the extent to which general patterns emerge, accounting for biogeographic, phylogenetic and environmental differences among sites. Common data collection across TMCFs should also allow better integration across disciplines, such as linking nutrient limitation, seed production and propagule recruitment, and enable cross-site comparisons of how TMCFs respond to drivers of global change, including rising cloud bases, increasing temperatures, altered disturbance regimes, biological invasions and extinction, and changing human land use.
Tropical montane cloud forest: environmental drivers of vegetation structure and ecosystem function
- Timothy J. Fahey, Ruth E. Sherman, Edmund V.J. Tanner
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- Published online by Cambridge University Press:
- 09 November 2015, pp. 355-367
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Tropical montane cloud forests (TMCF) are characterized by short trees, often twisted with multiple stems, with many stems per ground area, a large stem diameter to height ratio, and small, often thick leaves. These forests exhibit high root to shoot ratio, with a moderate leaf area index, low above-ground production, low leaf nutrient concentrations and often with luxuriant epiphytic growth. These traits of TMCF are caused by climatic conditions not geological substrate, and are particularly associated with frequent or persistent fog and low cloud. There are several reasons why fog might result in these features. Firstly, the fog and clouds reduce the amount of light received per unit area of ground and as closed-canopy forests absorb most of the light that reaches them the reduction in the total amount of light reduces growth. Secondly, the rate of photosynthesis per leaf area declines in comparison with that in the lowlands, which leads to less carbon fixation. Nitrogen supply limits growth in several of the few TMCFs where it has been investigated experimentally. High root : shoot biomass and production ratios are common in TMCF, and soils are often wet which may contribute to N limitation. Further study is needed to clarify the causes of several key features of TMCF ecosystems including high tree diameter : height ratio.
Geographic, environmental and biotic sources of variation in the nutrient relations of tropical montane forests
- James W. Dalling, Katherine Heineman, Grizelle González, Rebecca Ostertag
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- 20 November 2015, pp. 368-383
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Tropical montane forests (TMF) are associated with a widely observed suite of characteristics encompassing forest structure, plant traits and biogeochemistry. With respect to nutrient relations, montane forests are characterized by slow decomposition of organic matter, high investment in below-ground biomass and poor litter quality, relative to tropical lowland forests. However, within TMF there is considerable variation in substrate age, parent material, disturbance and species composition. Here we emphasize that many TMFs are likely to be co-limited by multiple nutrients, and that feedback among soil properties, species traits, microbial communities and environmental conditions drive forest productivity and soil carbon storage. To date, studies of the biogeochemistry of montane forests have been restricted to a few, mostly neotropical, sites and focused mainly on trees while ignoring mycorrhizas, epiphytes and microbial community structure. Incorporating the geographic, environmental and biotic variability in TMF will lead to a greater recognition of plant–soil feedbacks that are critical to understanding constraints on productivity, both under present conditions and under future climate, nitrogen-deposition and land-use scenarios.
Natural disturbance, vegetation patterns and ecological dynamics in tropical montane forests
- Shelley D. Crausbay, Patrick H. Martin
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- 14 July 2016, pp. 384-403
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Disturbance is a central process in forest dynamics, yet the role of natural disturbance in tropical montane forests (TMFs) has not been systematically addressed. We posit that disturbance in TMFs has a wider role than commonly acknowledged and its effects are distinctive because: (1) TMFs often have very low rates of productivity due to low resources, and so recovery from disturbance may be slow, (2) montane forests have marked environmental heterogeneity which interacts with disturbance, (3) a large percentage of TMFs are regularly exposed to high energy windstorms and landslides, and (4) TMFs contain a biogeographically rich mixture of tree species with divergent evolutionary histories that interact differently with different disturbance types. We reviewed the literature on natural disturbance in TMFs and found 119 peer-reviewed papers which met our search criteria. Our review shows that disturbance is widespread in TMFs with pronounced effects on structure, function, composition and dynamics. Disturbance is also evident in the ecology of TMF biota with clear examples of plant life-history traits adapted to disturbance, including disturbance-triggered germination, treefall gap strategies and resprouting ability. Important aspects of TMF disturbances are stochastic and site-specific, but there are broad patterns in disturbance type, frequency and severity along latitudinal, altitudinal and environmental gradients. Compared with the lowland tropics, TMF disturbances are more spatially structured, TMFs experience more disturbance types in a given area due to environmental complexity, and TMFs are much more prone to small-scale yet severe landslides as well the large and potentially catastrophic disturbances of cyclones, forest die-back and fire. On the whole, natural disturbance should assume a larger role in models of ecosystem processes and vegetation patterns in TMFs. An improved understanding of what creates variation in disturbance severity and post-disturbance recovery rates, how composition and diversity feedback on disturbance type and likelihood, and how global change will alter these dynamics are important priorities in future TMF ecology research.
Plant carbon and water fluxes in tropical montane cloud forests
- Sybil G. Gotsch, Heidi Asbjornsen, Gregory R. Goldsmith
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- 15 July 2016, pp. 404-420
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Tropical montane cloud forests (TMCFs) are dynamic ecosystems defined by frequent, but intermittent, contact with fog. The resultant microclimate can vary considerably over short spatial and temporal scales, affecting the ecophysiology of TMCF plants. We synthesized research to date on TMCF carbon and water fluxes at the scale of the leaf, plant and ecosystem and then contextualized this synthesis with tropical lowland forest ecosystems. Mean light-saturated photosynthesis was lower than that of lowland forests, probably due to the effects of persistent reduced radiation leading to shade acclimation. Scaled to the ecosystem, measures of annual net primary productivity were also lower. Mean rates of transpiration, from the scale of the leaf to the ecosystem, were also lower than in lowland sites, likely due to lower atmospheric water demand, although there was considerable overlap in range. Lastly, although carbon use efficiency appears relatively invariant, limited evidence indicates that water use efficiency generally increases with altitude, perhaps due to increased cloudiness exerting a stronger effect on vapour pressure deficit than photosynthesis. The results reveal clear differences in carbon and water balance between TMCFs and their lowland counterparts and suggest many outstanding questions for understanding TMCF ecophysiology now and in the future.
Biodiversity and metacommunity structure of animals along altitudinal gradients in tropical montane forests
- Michael R. Willig, Steven J. Presley
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- 09 November 2015, pp. 421-436
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The study of altitudinal gradients has made enduring contributions to the theoretical and empirical bases of modern biology. Unfortunately, the persistence of these systems and the species that compose them is threatened by land-use change at lower altitudes and by climate change throughout the gradients, but especially at higher altitudes. In this review, we focus on two broad themes that are inspired by altitudinal variation in tropical montane regions: (1) dimensions of biodiversity and (2) metacommunity structure. Species richness generally decreased with increasing altitude, although not always in a linear fashion. Mid-altitudinal peaks in richness were less common than monotonic declines, and altitudinal increases in richness were restricted to amphibian faunas. Moreover, gradients of biodiversity differed among dimensions (taxonomic, phylogenetic and functional) as well as among faunas (bats, rodents, birds) in the tropical Andes, suggesting that species richness is not a good surrogate for dimensions that reflect differences in the function or evolutionary history of species. Tropical montane metacommunities evinced a variety of structures, including nested (bats), Clementsian (rodents, bats, gastropods), quasi-Clementsian (reptiles, amphibians, passerines) and quasi-Gleasonian (gastropods) patterns. Nonetheless, compositional changes were always associated with the ecotones between rain forest and cloud forest, regardless of fauna.
Seed-dispersal ecology of tropical montane forests
- Hazel Chapman, Norbert J. Cordeiro, Paul Dutton, Dan Wenny, Shumpei Kitamura, Beth Kaplin, Felipe P. L. Melo, Michael J. Lawes
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- 03 August 2016, pp. 437-454
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Seed-dispersal ecology in tropical montane forests (TMF) differs in some predictable ways from tropical lowland forests (TLF). Environmental, biogeographic and biotic factors together shape dispersal syndromes which in turn influence forest structure and community composition. Data on diaspore traits along five elevational gradients from forests in Thailand, the Philippines, Tanzania, Malawi and Nigeria showed that diaspore size decreases with increasing altitude, fleshy fruits remain the most common fruit type but the relative proportion of wind-dispersed diaspores increases with altitude. Probably corresponding to diaspore size decreasing with increasing elevation, we also provide evidence that avian body size and gape width decrease with increasing altitude. Among other notable changes in the frugivorous fauna across elevational gradients, we found quantitative evidence illustrating that the proportion of bird versus mammalian frugivores increases with altitude, while TMF primates decrease in diversity and density, and switch diets to include less fruit and more leaf proportionately. A paucity of studies on dispersal distance and seed shadows, the dispersal/predation balance and density-dependent mortality thwart much-needed conclusive comparisons of seed dispersal ecology between TMF and TLF, especially from understudied Asian forests. We examine the available evidence, reveal knowledge gaps and recommend research to enhance our understanding of seed dispersal ecology in tropical forests. This review demonstrates that seed dispersal is a more deterministic and important process in tropical montane forests than has been previously appreciated.
The functional roles of epiphytes and arboreal soils in tropical montane cloud forests
- Sybil G. Gotsch, Nalini Nadkarni, Autumn Amici
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- Published online by Cambridge University Press:
- 13 July 2016, pp. 455-468
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Epiphytes and their associated decomposing litter and arboreal soils (herein, epiphytic material, EM) are ubiquitous features of tropical montane cloud forests (TMCF) and play important roles in ecosystem function. EM intercepts water and nutrients from the atmosphere and from intercepted host tree sources, and may contribute significant inputs of these resources to the forest floor. Despite the importance of EM in the TMCF, a systematic review of the ecosystem roles of EM has not been compiled before. We have synthesized the literature that documents functions of EM in undisturbed TMCFs and discuss how these roles may be affected by disturbances, including changes in climate and land use. The range of EM biomass and water storage in the TMCF varies greatly across sites, with different amounts associated with stand age and microclimate. EM is important as habitat and food for birds and mammals, with over 200 species of birds documented as using EM in the Neotropics. Given its sensitivity to moisture, projected shifts in cloud base heights or precipitation due to changes in climate will likely have a large impact on this community and changes in EM diversity or abundance may have cascading impacts on the ecosystem function of the TMCF.