Vertical distribution and soil organic matter composition in a montane cloud forest, Oaxaca, Mexico

In montane cloud forests (MCF), the main soil organic carbon (SOC) pool is believed to be constituted by organic debris accumulated on soil surface and, to a lesser extent, by the organic fraction associated with the mineral matrix. The vertical distribution of SOC within soil has strong implications on the composition, stabilization and turnover of the soil organic matter (SOM). In ecosystems like MCF, where the climatic and edaphic conditions varied with altitude, the SOM accumulation and stabilization mechanisms possibly respond to these changes. For that reason, we studied the vertical distribution, accumulation and chemical composition of SOM in five montane cloud forest communities located between 1,500 and 2,500?m a.s.l. Two main SOC accumulation patterns were found: one at 1,500, 1,950 and 2,400?m a.s.l., with SOC content gradually decreasing with depth (cumulative); and another at 2,050 and 2,500?m a.s.l. where SOC had a strong maximum in the surface horizon and a less pronounced increase the spodic horizon (eluviation–illuviation pattern). The total SOC pool in soil decreased in inverse relation to altitude from 227?C?ha^?1 at 1,500?m a.s.l. down to 143?mg?C?ha^?1 at 2,500?m a.s.l. About 40–60?% of total SOC content corresponded to the surficial organic horizon. The chemical fractionation of the SOM denoted in general predominance of the fulvic acid fraction, and high content of humin and humic acid fractions. We considered that the main SOC vertical distribution processes were related to the raw humus accumulation, decomposition in situ, podzolization in the eluviation–illuviation pattern soils mainly.     

Rainfall interception and fog precipitation in a tropical montane cloud forest of Guatemala

Tropical montane cloud forest hydrology is complex because of the presence of epiphytic life-forms that increase canopy surfaces and fog persistency. Fog precipitation is a hydrological input common to cloud forests, and forms when fog droplets are intercepted by the canopy and fall to the forest floor. Interception and fog precipitation was determined for a 2100 m site and a 2550 m site in a first-order tributary of the Sierra de las Minas Biosphere Reserve, Guatemala by calculating the difference between throughfall and gross precipitation for a 44-week period. Both sites were situated within closed-canopy cloud forests. The 2100 m site was on the windward slope of Montaña de Miranda near the lower boundary of the cloud forest and the 2550 m site was at the summit. Fog precipitation was found during periods in which throughfall exceeds gross precipitation. Fog precipitation was greater at 2550 m than at 2100 m. Data collected by precipitation and throughfall gauges demonstrate the existence of seasonal fog precipitation with the greatest fog precipitation occurring in the dry season (November–April). Fog precipitation contributes approximately 1 mm per day to the hydrological budget of the cloud forest at 2550 m during the dry season, and 0.5 mm per day during the rainy season (May–October).     

Fungal diversity in coffee plantation systems and in a tropical montane cloud forest in Veracruz,Mexico

We compared the abundance, species richness and diversity of saprobic filamentous microfungi in the forest and in coffee plantation systems (with different biophysical structures of the vegetation and agricultural management) and evaluated the degree of similarity in species composition among these sites. Soil washing was used to isolate the saprobic filamentous microfungi. Emerging colonies were quantified and transferred to tubes with culture medium and then mounted on semi-permanent slides for identification under the microscope. From 90 samples and 4,500 inoculated soils particles, 415 species were distinguished. The genera Trichoderma, Penicillium, Fusarium, Chaetomium and Humicola were the most frequent in all study sites. The transformation of the tropical montane cloud forest in coffee plantation had no significant effect on the abundance, species richness and diversity of the saprobic filamentous microfungi. Effects of the biophysical structures of the vegetation and the agricultural management of the sites were only detected at the level of dominant genera (Fusarium, Trichoderma and Penicillium) and on the evenness. The low degree of similarity among the six sites suggests the existence of a high exchange of species.     

Spatial and temporal variations in soil respiration in relation to stand structure and soil parameters in an unmanaged beech forest

Soil CO2 efflux (soil respiration) plays a crucial role in the global carbon cycle and efflux rates may be strongly altered by climate change. We investigated the spatial patterns of soil respiration rates in 144 measurement locations in a 0.5-ha plot and the temporal patterns along a 300-m transect in the 0.5-ha plot. Measurements were made in an unmanaged, highly heterogeneous beech forest during 2000 and 2001. We investigated the effects of soil, roots and forest stand structure on soil respiration, and we also assessed the stability of these spatial patterns over time. Soil temperature alone explained between 68 and 95% of the temporal variation in soil respiration; however, pronounced spatial scatter of respiration rates was not explained by soil temperature. The observed spatial patterns stayed remarkably stable throughout the growing season and over 2 years. The most important structural parameter of the stand was the mean diameter at breast height of trees within a distance of 4 m of the measurement locations (m-dbh4), which explained 10-19% of the variation in soil respiration throughout the growing season. Among the soil chemical parameters, carbon content (bulk as well as dissolved) and magnesium content explained 62% of the spatial variation in soil respiration. The final best model combining soil, root and stand structural parameters (fine root biomass, soil carbon content, m-dbh4 and soil water content) explained 79% of the variation in soil respiration, illustrating the importance of both biotic and abiotic factors.     

13C-NMR spectroscopy studies of humic substances in subtropical perhumid montane forest soil

We investigated soil organic matter in a forest of natural Hinoki cypress (Chamaecyparis obtusa) under perhumid weather conditions in north central Taiwan. Humic substances along the transect from the summit and footslope to lakeshore were characterized by use of solid-state cross-polarization, magic-angle-spinning ¹³C nuclear magnetic resonance spectroscopy (CP/MAS ¹³C-NMR). The major components of soil organic carbon in whole soil and humic substances were alkyl-C, O-alkyl-C, and di-O-alkyl-C, ranging from 60.6% to 80.7%, then aromatic-C, 7.5% to 9.8%. The degree of humification of soil organic matter, both O-alkyl-C/alkyl-C ratio and aromaticity, decreased slightly from the summit to lakeshore. The content of functional groups of polar and acidic groups, including O-alkyl-C, di-O-alkyl-C, and carboxyl-C, corresponded with the topographical effect, increasing slightly from the summit to lakeshore. However, the relatively low degree of humification in soils of this perhumid forest and low aromaticity were due to high precipitation and acidity, which appeared to hinder organic matter decomposition with topography change.     

Key attributes to the disturbance response of montane cloud forest trees: shade tolerance,dispersal mode and the capacity to form a seed bank

? Context

Understanding the biological mechanisms related to plant response to disturbance is essential for predicting the changes in the structure of plant communities resulting from disturbance.

? Aims

The aims of this study are to identify the regeneration traits linked to the response to disturbance of tree species of a montane cloud forest in order to define regeneration types and to explore whether disturbance intensity acts to select specific regeneration types that can colonize disturbed plots.

? Methods

A theoretical model was developed and evaluated using structural equation modeling to link two latent variables: regeneration type and vulnerability to disturbance. We studied two plots that were subjected to different intensities of disturbance and one plot of mature forest.

? Results

Shade tolerance, dispersal mode, and the capacity to form a seed bank were the traits that determine regeneration type. There was a trend for re-sprouting capacity to be associated with the regeneration types dominant in disturbed plots. Regeneration types were differentially distributed among the sites.

? Conclusion

There was evidence that disturbance intensity has a filtering effect on the colonization of disturbed plots. Species vulnerability to disturbance depends on specific combinations of regeneration traits. Structural equation modeling is useful for exploring plant response to disturbance using key parameters linked to plant persistence.     

Transpiration of silver Fir (Abies alba mill.) during and after drought in relation to soil properties in a Mediterranean mountain area

Context

Silver fir is declining and dying at its southern margin on the Mediterranean area, where climate is expected to be warmer and drier. At a regional scale, silver fir seems to be vulnerable to drought, and at a forest stand scale, tree death seems to be distributed according to soil water availability.

Aim

To understand the vulnerability of silver fir to drought, factors involved in the regulation of transpiration were assessed with respect to soil properties in order to document the spatial distribution of death rates.

Methods

Soil properties were characterized by electric resistivity measurements. Sap flow density and predawn needle water potential were recorded on sampled trees during several years, and crown specific transpiration was estimated. In addition, the vulnerability of coarse roots and branches to cavitation was quantified.

Results and conclusion

Trees growing on soils with a large water storage capacity were the most vulnerable to drought induced soil water deficits. Transpiration was down-regulated as soon as predawn water potential decreased. The vulnerability to cavitation was low, which protected the trees from run-away xylem embolism. Severe soil water deficits led to a rapid decrease of transpiration, which was still visible the following year. The drop-off in transpiration was mainly due to inner sap flow that almost ceased after the drought on all monitored trees. Our results suggest that root dynamic and the ability of roots to take up water were modified by soil water deficit over several years. Such a regulation scheme needs to be better documented and included in models to address silver fir forest responses to drought.     

Response of water and nutrient fluxes to improvement fellings in a tropical montane forest in Ecuador

Management of natural forests might be one option to reduce the high deforestation rate in Ecuador. We therefore evaluated the response of water and nutrient cycles in a natural tropical montane forest to improvement fellings with the aim of favoring economically valuable target trees which will later be harvested with additional ecosystem impacts not considered here.     

Lichen diversity in temperate montane forests is influenced by forest structure more than climate

Although the effect of forest management on lichens in temperate forests has been widely examined, little is known about the influence of management-related factors on their biodiversity relative to factors that cannot be altered by management. Here we determined whether forest structure or climate determines lichen diversity in the Bavarian Forest National Park in southeastern Germany, taking spatial variables into account. We investigated 517 single tree stems along 4 transects in 113 pre-stratified plots (8 m in diameter) in this montane forest. We grouped environmental variables into three sets: climate (macroclimate, non-manageable), forest structure (manageable), and space. The explanatory powers of these sets of variables for lichen diversity were compared using variance partitioning for the lichen community, species density, and threatened species density. The relationships of single characteristics of forest structure with lichen species diversity were analyzed using generalized linear models (GLM). Lichen diversity was better explained by stand structures than by climate. Spatial effects influenced the number of species per plot. Among the structural features, the availability of dead wood and sycamore maple as well as forest continuity were most important for the enhancement of lichen diversity. Open canopy structures affected the total diversity positively. Although the availability of large trees was not an influential factor in the GLM at the plot level, high diversity levels were generally associated with large stem diameters at the level of single stems. We provide recommendations for sustainable forest-management practices that aim at specifically enhancing lichen diversity in temperate areas experiencing low levels of air pollution.     

Implications of fires on carbon budgets in Andean cloud montane forest: The importance of peat soils and tree resprouting

Fire in tropical montane cloud forests (TMCFs) is not as rare as once believed. Andean TMCFs sit immediately below highly flammable, high-altitude grasslands (Puna/Páramo) that suffer from recurrent anthropogenic fire. This treeline is a zone of climatic tension where substantial future warming is likely to force upward tree migrations, while increased fire presence and fire impacts are likely to force it downwards. TMCFs contain large carbon stocks in their peat soils and their loss through fire is a currently unaccounted for regional source of CO₂. This study, conducted in the southern Peruvian Andes (>2800 m), documents differences in live tree biomass, fine root biomass, fallen and standing dead wood, and soil organic carbon in 4 paired-sample plots (burned versus control) following the severe ground fires that occurred during the 2005 Andean drought. Peat soils contributed the most to biomass burning emissions, with lower values corresponding to an 89% mean stock difference compared to the controls (mean ± SE) (54.1 ± 22.3 vs. 5.8 ± 5.3 MgC ha⁻¹). Contrastingly, carbon stocks from live standing trees differed by a non-significant 37% lower value in the burned plots compared to the controls, largely compensated by vigorous resprouting (45.5 ± 17.4 vs. 69.2 ± 13.4 MgC ha⁻¹). Both standing dead trees and fallen dead wood were significantly higher in the burned plots with a three-fold difference from the controls: dead Trees 45.2 ± 9.4 vs. 16.4 ± 4.4 MgC ha⁻¹, and ca. a 2 fold difference for the fallen dead wood: 11.2 ± 5 vs. 6.7 ± 3.2 MgC ha⁻¹ for the burned plots versus their controls. A preliminary estimate of the regional contribution of biomass burning emissions from Andean TMCFs for the period 2000-2008, resulted in mean carbon emission rates of 1.3 TgC yr⁻¹ (max-min: 1.8-0.8 TgC yr⁻¹). This value is in the same order of magnitude than South American annual fire emissions (300 TgC yr⁻¹) suggesting the need for further research on Andean forest fires. On-going projects on the region are working on the promotion of landowner participation in TMCFs conservation through REDD+ mechanism. The heart of the proposed initiative is reforestation of degraded lands with green fire breaks enriched with economically valuable Andean plant species. The cultivation of these species may contribute to reduce deforestation pressure on the Amazonian cloud forest by providing an alternative income to local communities, at the same time that they prevent the spread of fire into Manu National Park and adjacent community-held forests, protecting forest and reducing CO₂ emissions.     

Natural development and regeneration of a Central European montane spruce forest

Montane Norway spruce forests of Central Europe have a very long tradition of use for timber production; however, recently there has been increasing concern for their role in maintaining biological diversity. This concern, coupled with recent severe windstorms that led to wide-spread bark beetle outbreaks, has brought the management of montane spruce forests to the forefront of public policy discussions in Central Europe. In order to shed light on the natural development and current structure of mature montane spruce forests, we established four 0.25 ha research plots in a semi-natural montane spruce forest in the Šumava Mountains (The Bohemian Forest), Czech Republic. We mapped all trees, extracted increment cores for age and growth-pattern analyses, and inventoried all current tree regeneration, including the substrates on which it was found. Stands were characterized by uni-modal tree diameter distributions and high basal areas (56.6 m² ha⁻¹ on average), indicating a natural transition from the stem exclusion phase towards the understory reinitiation phase. The stands showed largely single-cohort recruitment age structures, however, with recruitment spanning seven decades. Our analyses suggest that this cohort existed as advance regeneration prior to major disturbances in the late 1800s, which included post-bark beetle salvage logging. Spatial pattern analyses of living and dead stems combined, showed an increase in uniformity of living trees, pointing to the role of natural density-dependent mortality. However, past growth patterns and historical documentation suggest that low intensity canopy disturbances (wind and snow) also caused mortality and diversified canopy structure. Because the stands developed naturally over the past 120+ years and thus escaped thinning operations, high volumes of coarse woody debris (94 m³ ha⁻¹) and snag densities (546 stems ha⁻¹) have accrued. Advance spruce regeneration was quite abundant and existed primarily on deadwood substrates, even though these occupied only a small percent of stand area. Because of salvage logging in the late 1880s, these stands do not qualify, according to the traditional paradigm, as natural spruce forests. As a result, they are recently subject to active management practices including salvage logging that remove dead and dying trees. Given the importance of deadwood for forest regeneration and recovery from disturbance, as demonstrated in this study, we argue that dead wood removal may limit future natural regeneration in these stands. Thus, the purported benefits of removing dead and dying trees from semi-natural forests must be carefully weighed against the potential detrimental impacts on natural spruce forest regeneration and biodiversity.     

Diversity and abundance of arbuscular mycorrhizal fungi spores under different coffee production systems and in a tropical montane cloud forest patch in Veracruz, Mexico

We evaluate the arbuscular mycorrhizal fungi (AMF) community as measured by spores in different coffee production systems (at the depth of 0?C15?cm). In addition, we analyze the similarities between the AMF communities in coffee production systems and those that occur in a tropical montane cloud forest patch in order to evaluate the capacity of coffee production systems to preserve the native AMF community. We carried out four samplings in five coffee production systems representative of a vegetation structure gradient, and in a forest. From 120 soil samples, 33 morphospecies were detected. In all the sites, the dominant morphospecies were Glomus clarum and Glomus sp. 3. We found no significant difference in AMF spore richness between sites. Diversity was similar in most of the coffee production systems. Significant differences were only detected in spore abundance; during the dry season the forest, shaded traditional rustic system and shaded simple system presented the highest spore abundance. With the exception of one species exclusive to the forest, the coffee production systems all share the same AMF species as the forest. The coffee production systems with the greatest similarity to cloud forest were the shaded traditional rustic system and the shaded simple system. It is suggested that control of weeds and fertilization could be important factors influencing the composition and abundance of AMF spores in coffee production systems.     

Early seedling establishment of two tropical montane cloud forest tree species: The role of native and exotic grasses

Tropical montane cloud forest has been undergoing a drastic reduction because of its widespread conversion to pastures. Once these forests have been cleared exotic grasses are deliberately introduced for forage production. Exotic grass species commonly form monodominant stands and produce more biomass than native grass species, resulting in the inhibition of secondary succession and tree regeneration. The purpose of this study was to assess the effect of native vs. exotic grass species on the early establishment of two native tree seedlings (Mexican alder, Alnus acuminata and Jalapa oak, Quercus xalapensis) on an abandoned farm in central Veracruz, Mexico. Seedling survival and growth were monitored (over 46 weeks) in relation to grass cover and height, and available photosynthetic active radiation (PAR). More seedlings survived in the presence of the native grass Panicum glutinosum than those growing with the exotic grass Cynodon plectostachyus (92% vs. 48%). The causes of seedling mortality varied between species; Q. xalapensis was affected by herbivory by voles but mainly in the exotic grass-dominated stands, whereas A. acuminata seedlings died due to competition with the exotic grass. A. acuminata seedlings increased more in height in the exotic grass-dominated stands (102 ± 7.8 cm) compared to native grass-dominated stands (51 ± 4.7 cm). Grass layer height, cover and available PAR were correlated (Pearson; p < 0.05). In the exotic grass dominated plots, grass layer height was correlated with the relative height growth rates of Q. xalapensis (Pearson; p < 0.05). These results indicate that the exotic grass may be affecting tree regeneration directly (grass competition) and indirectly (higher herbivory). Passive restoration may occur once P. glutinosum dominated pastures are abandoned. However, when C. plectostachyus dominates, introduction of early and mid successional tree seedlings protected against vole damage is needed.     

Contribution of forest floor fractions to carbon storage and abundance patterns of arbuscular mycorrhizal fungal colonisation in a tropical montane forest

Forest floor carbon stocks, which include different components of litter, hemic and sapric materials, have not been empirically quantified in tropical montane forest, although they influence soil carbon (C) pools. To date, the contribution of arbuscular mycorrhizae in C sequestration potentials in tropical montane forests have not been clearly investigated. This study determined the amount of C stocks in the different decomposing layers of forest floor, mainly litter, hemic and sapric materials. The abundance of arbuscular mycorrhizal root colonisation differed among forest floor fractions. Forest floor was measured for depth, area density, dry mass and carbon fraction separately in Sungai Kial Forest Reserve, Pahang, Malaysia to calculate C stocks. Percentages of root colonisation in the hemic and sapric materials were investigated. The results showed that forest floor C stocks were significantly higher in hemic (5 Mg C ha^?1) and sapric (7.7 Mg C ha^?1) compared with the litter fragments (1.5 Mg C ha^?1). Mycorrhizal root colonisation was significantly higher (75%) in the toeslope compared with the summit area in the hemic materials. Segregation of forest floor layers provided greater accuracy in forest floor C stocks reporting.     

Direct seeding of late-successional trees to restore tropical montane forest

Natural regeneration of large-seeded, late-successional trees in fragmented tropical landscapes can be strongly limited by a lack of seed dispersal resulting in the need for more intensive restoration approaches, such as enrichment planting, to include these species in future forests. Direct seeding may be an alternative low-cost approach to planting nursery-raised tree seedlings, but there is minimal information on its efficacy or when in the successional process this technique will be most successful. We tested directly seeding five native tree species into habitats representing passive and active restoration approaches: (1) recently abandoned pasture; (2) naturally establishing, young secondary forests; and (3) young, mixed-species (fast-growing N-fixers and commercially valuable species) tree plantations established to facilitate montane forest recovery in southern Costa Rica. We monitored germination, survival, growth, and above- and below-ground biomass over a 2-year period. Germination in pastures, secondary forests, and tree plantations was similar (∼43%). Seedling survival after one and two years was significantly higher under tree plantations (91% year 1, 75% year 2) compared to secondary forests (76, 44%) or pastures (74, 41%). Moreover, seedlings had greater total biomass and lower root:shoot ratios in the plantations, suggesting higher nutrient availability in that treatment. Costs for direct seeding were 10- to 30-fold less per 100 seedlings after 2-year compared to nursery-raised seedlings planted at the same sites; however, there are important trade-offs to the two restoration approaches. Planting nursery-raised seedling is a more effective but higher cost approach for rapidly establishing canopy cover and restoring large areas whereas direct seeding is a more efficient way to enrich an existing system. We particularly recommend using direct seeding as a complimentary measure to the more intensive restoration approach of planting fast-growing and N-fixing trees.     

Response of vascular epiphyte diversity to different land-use intensities in a neotropical montane wet forest

Although vascular epiphytes contribute substantially to the biodiversity of tropical montane forests, it is unclear how their diversity and community composition is affected by forest alteration. We studied the response of vascular epiphyte assemblages to different intensities of land-use in a montane wet forest of northeastern Ecuador: (1) unmanaged mature forest; (2) mature forest with mid- and understorey opened for cattle grazing; and (3) isolated remnant trees in cattle pastures. The numbers of individuals and species of epiphytes per host tree did not differ significantly between land-use types, neither did total epiphyte species richness (n = 30 trees). However, total species richness of pteridophytes was significantly lower on isolated remnant trees compared to unmanaged forest, whereas several taxa rich in xerotolerant species (Bromeliaceae, Orchidaceae, Piperaceae) exhibited the opposite trend. An analysis of floristic composition using ordination (NMS) and randomisation techniques (MRPP) showed that epiphyte assemblages on isolated remnant trees were significantly distinct from unmanaged forest while managed forest was intermediate between those two vegetation types. Ordination analysis further indicated reduced floristic heterogeneity in disturbed habitats. These results suggest considerable, rapid species turnover since land-use change 6 years prior to study, with pteridophytes being replaced by more xerotolerant taxa. We attribute this floristic turnover primarily to changes in microclimate towards higher levels of light and desiccation stress associated with forest disturbance. Our results support the notion that community composition offers a more sensitive indicator of human disturbance than species richness.     

Change in forest structure in the boreal montane ecosystem of Mount Mitchell,North Carolina1

Surveys were undertaken on Mt. Mitchell, N. C. (USA) from Spring 1984 through 1987 to determine the extent and rate of decline and mortality of red spruce and Fraser fir ecosystems. Data indicate that regardless of cause and effect mechanisms the physical structure and integrity of the boreal montane forest has greatly deteriorated over the past 4 years. Along ridges, and particularly on west facing aspects greater than 80% of all mature stands are dead. No definitive conclusions can be made concerning etiological agents; however, current hypotheses include biotic factors, climatic abberations, and/or the acute or chronic stress from anthropogenic pollutants.