Diversity and ecology of lianas in tropical dry evergreen forests on the Coromandel Coast of India under various disturbance regimes

In this study we attempted to explore patterns of diversity, abundance, climbing and dispersal mode of lianas in relation to disturbance in 40 Indian subtropical dry forests. The sites were selected to represent four disturbance categories: relatively undisturbed, moderately disturbed, much disturbed and heavily disturbed. All lianas ≥1 cm dbh were counted, which resulted in a total amount of 5689 individuals of lianas, representing 77 species in 62 genera and 32 families. Liana species richness and abundance increased with forest disturbance, but the liana basal area values showed an opposite trend, with high scores in undisturbed sites. Twining was the main climbing mechanism (61.3%) and zoochory (59.6%) was the main dispersal mode in all the four forest categories. Application of Bray–Curtis cluster analysis produced three distinct clusters in which the much disturbed category was more distant from the others. High abundance of large lianas in undisturbed sites and that of the invasive Lantana camara in heavily disturbed site signals the conservation significance of the less disturbed study sites. The predominance of zoochorous dispersal indicates the faunal dependence of lianas, besides of host trees, thus underlining the need for a holistic approach in biodiversity conservation of this and similar tropical forests.     

Liana diversity and distribution in four tropical dry evergreen forests on the Coromandel coast of south India

Liana diversity was inventoried in four tropical dry evergreen forest sites that are characterized by numerous trees, of short stature and small diameter, and a varying degree of anthropogenic disturbance, on the Coromandel coast of south India. A 1-ha plot was established in each of the four sites and was subdivided into 100 quadrats of 10 m× 10 m. All lianas 1 cm diameter at breast height (dbh) rooted within the plot were enumerated. The species richness and density of lianas, with respect to site disturbance and forest stature, varied across the sites. Liana density totaled 3307 individuals (range 497–1163 individuals ha^–1) and species richness totaled 39 species (range 24–29 species ha^–1) representing 34 genera and 24 families. Combretaceae, Asclepiadaceae, Capparaceae and Vitaceae were the well-represented families. The top five species Strychnos minor, Combretum albidum, Derris ovalifolia, Jasminum angustifolium and Reissantia indica contributed 55% of total density. The slopes of the species–area curves were different for each of the four sites and the curve stabilized in only one site. Of the four climbing modes recognized among the total 39 species, 18 were twiners (56% of the total density). Eight species (24% of density) were tendril climbers and 12 species (16% of density) were scramblers. Hugonia mystax was the only hook climber. All the 39 species and 88% of liana density were encountered within a category of 6 cm dbh or less, and a similar pattern prevailed in the individual sites. Of the three diaspore dispersal modes found among the 39 liana species, animal (64%) and wind (23%) dispersal were predominant over the autochorous mode (13%). Liana diversity and distribution in dry forest communities appear to be influenced by forest stature and site disturbance levels. In the light of the extent of liana diversity and sacred grove status of the study sites, the need for forest conservation, involving local people, is emphasized.     

Tropical dry forests in New Caledonia

Tropical dry forest is the most endangered major vegetation type in the New Caledonia biodiversity hotspot. Vegetation surveys following a transect method used by Gentry were undertaken in two tropical dry forest sites, Ouen-Toro and Pindai, in order to compare species richness, floristic composition, and structure. Pindai contained significantly higher species richness than Ouen-Toro, although there was little difference in forest structure. Tropical dry forest sites in New Caledonia were compared to seven other biodiversity hotspots with tropical dry forest where Gentry's transect method was employed. New Caledonia and other tropical dry forests on islands contain significantly lower species richness than mainland tropical dry forests in biodiversity hotspots. However, New Caledonia contained the highest number of threatened species based on IUCN global conservation categories. Tropical dry forest in New Caledonia appears to be the world's most endangered tropical dry forest based on the extent of forest, number of reserves, and threatened species. Management of tropical dry forests on private and community lands is absolutely imperative to the long-term persistence of this ecosystem.     

Regeneration of tropical dry forests in Central America,with examples from Nicaragua

Abstract. This paper focuses on the management potential of natural forests in the tropical dry zones of Central America, particularly Nicaragua. Distribution and status of dry forest formations are reviewed. Two case studies from the Pacific coast of Nicaragua are presented to illustrate the disturbance created by traditional utilization, and to show the relatively high potential for natural forest management, with many of the tree species having economic value. Natural forest management, together with conservation and reforestation, is viewed as the main management option for the tropical dry forest areas. In spite of its importance, this option has received little attention or promotion from the governmental forestry sector. Four complementary options which can be devised for silvicultural work in production forests are: encouragement of advanced growth of desirable tree species, inducement of natural regeneration, coppice management, and compensatory planting. These options form different strategies for rehabilitation of the production and conservation functions of the forest and may well be combined in the same area. Research applied to better utilization of the forest resources should form an intrinsic part of rural development programs. Research work in tropical dry forest areas should focus on ecological, as well as socio-cultural and economical aspects related to the management options. Experimental sites are important enabling and securing long-term basic and applied biological research. They may also serve as units for demonstration, training and extension.     

Species Richness,Spatial Variation,and Abundance of the Soil Seed Bank of a Secondary Tropical Rain Forest1

Despite the importance of the soil seed bank in tropical forest regeneration, little is known about spatial variability in species composition and abundance of seeds stored in the soil. To develop sampling methods for comparative studies, we examined species richness, spatial variation, and abundance of germinants from the soil seed bank in a 16 year old secondary, tropical wet forest at La Selva Biological Station, Costa Rica. Surface soil (10 cm deep, 4.7 cm diameter) was collected at the intersection points of a gridded 1 ha plot (10 × 10-m grid, 121 samples) and in a nested 100 m² subplot (2 × 2-m grid, 36 samples). The 1 ha plot had a density of 4535 seeds/m² with 34 species observed. Based on a series of 100 randomized species accumulation curves, a Michaelis-Menten fit predicted a mean species richness of 36.3 species; the number of observed species was close to the predicted asymptote. A nonparametric, first-order jackknife species richness estimator predicted a species richness of 37.0 species. Eighty-five and 95 percent of the observed species richness is contained, on average, within 41 and 74 pooled samples, respectively. Within the 100 m² nested subplot, a density of 5476 seeds/m² was observed, comprising 26 species with an estimated species richness (Michaelis-Menten fit) of 29.1 species. The jackknife species richness estimator predicted a species richness of 36.7 species. For species richness and abundance of both plots, spatial autocorrelation statistics (Moran's I) were not significantly different from zero at lag distances from 2 to 100 m, indicating a random distribution at these spatial scales. For this site, accurate estimates of species composition depend upon the number of samples collected as well as the spatial distribution of sampling effort. Many small samples distributed over a large area provide greater accuracy and precision for estimating species richness of the soil seed bank.     

Diversity of understory plants in undisturbed and disturbed tropical lowland forests of Little Andaman Island, India

Species richness and density of understory plants were investigated in eight 1 ha plots, distributed one each in undisturbed and disturbed tropical evergreen, semi-evergreen, deciduous and littoral forests of Little Andaman island, India, which falls under one of the eight hottest hotspots of Biodiversity in the world viz. the Indo-Burma. One hundred 1 m⁻² quadrats were established in each 1 ha plot, in which all the understory plants (that include herbs, undershrubs, shrubs and herbaceous climbers) were enumerated. The total density of understory plants was 6,812 individuals (851 ha⁻¹) and species richness was 108 species, representing 104 genera and 50 families. Across the four forest types and eight study plots, the species richness ranged from 10 to 39 species ha⁻¹. All the disturbed sites harbored greater number of species than their undisturbed counterparts. Herbs dominated by species (63%) and density (4,259 individuals). The grass Eragrostis tenella (1,860 individuals; IVI 40), the invasive climber Mikania cordata (803; IVI 20) and the shrub Anaxagorea luzonensis (481; IVI 17.5) were the most abundant species. Poaceae, Asteraceae, Acanthaceae, Orchidaceae and Euphorbiaceae constituted the species-rich families represented by 6 species each. The species-area curves attained an asymptote at 0.8 ha level except in sites DD and DL, indicating 1 ha plot is not sufficient to capture all the understory species in disturbed forests. The alien weeds formed about one-fourth of the species richness (31 species; 28%) and density (1,926 individuals; 28.3%) in the study sites, indicating the extent of weed invasion and the attention required for effective conservation of the native biodiversity of the fragile island forest ecosystem.     

Tree allometry and improved estimation of carbon stocks and balance in tropical forests

Tropical forests hold large stores of carbon, yet uncertainty remains regarding their quantitative contribution to the global carbon cycle. One approach to quantifying carbon biomass stores consists in inferring changes from long-term forest inventory plots. Regression models are used to convert inventory data into an estimate of aboveground biomass (AGB). We provide a critical reassessment of the quality and the robustness of these models across tropical forest types, using a large dataset of 2,410 trees ≥ 5 cm diameter, directly harvested in 27 study sites across the tropics. Proportional relationships between aboveground biomass and the product of wood density, trunk cross-sectional area, and total height are constructed. We also develop a regression model involving wood density and stem diameter only. Our models were tested for secondary and old-growth forests, for dry, moist and wet forests, for lowland and montane forests, and for mangrove forests. The most important predictors of AGB of a tree were, in decreasing order of importance, its trunk diameter, wood specific gravity, total height, and forest type (dry, moist, or wet). Overestimates prevailed, giving a bias of 0.5–6.5% when errors were averaged across all stands. Our regression models can be used reliably to predict aboveground tree biomass across a broad range of tropical forests. Because they are based on an unprecedented dataset, these models should improve the quality of tropical biomass estimates, and bring consensus about the contribution of the tropical forest biome and tropical deforestation to the global carbon cycle. Electronic Supplementary Material Supplementary material is available for this article at     

Effects of environmental heterogeneity on species diversity and composition of terrestrial bryophyte assemblages in tropical montane forests of southern Ecuador

Background: Most studies on tropical bryophytes deal with epiphytic species. This is the first ecological study of tropical forests that focuses specifically on terrestrial bryophytes.

Aim: To investigate the differences between slope and ridge environments in upper montane forests of southern Ecuador in terms of species diversity (richness, abundance), species composition and life forms of terrestrial bryophytes.

Methods: We used Non-metric Multidimensional Scaling (NMDS) to group bryophyte relevés by study location, habitat type and exposure class. Species indicator values were calculated and compared for different habitats.

Results: In total, 140 species were recorded, the majority being liverworts. NMDS analyses and Mantel correlations clearly separated between slope and ridge relevés, and between sunny and shaded microhabitats on ridges. Bryophyte life forms also showed different distribution patterns in slope and in ridge habitats. Mosses were more prominent in sunny than in shaded microhabitats.

Conclusions: Environmental differentiation between ridges and slopes, and small-scale variation in microclimatic conditions caused by differences in exposure, were stronger predictors of species richness and composition than geographical distance between study sites.     

Titimbera,a new genus of Orthocladiinae from South and Central America (Diptera: Chironomidae)

Titimbera n. gen. is erected based on the males of three new species from South and Central America: T. amazonica n. sp. from the Amazon region, Brazil; T. titi n. sp. from Venezuela and T. laselvensis n. sp. from Costa Rica. The combination of bare eyes and wing membrane; antenna without strong apical seta; scalpellate acrostichals in mid scutum; costa strongly extended; R₄₊₅ ending opposite to M₃₊₄; Cu₁ strongly curved to slightly sinuous; anal point sitting high on tergite IX, nearly parallel-sided with bluntly rounded apex; and club-shaped to subtriangular gonostylus with distinct heel will separate the genus from all other orthoclads.     

Dynamics of viable nitrifier community, N-mineralization and nitrification in seasonally dry tropical forests and savanna

The study was conducted in Vindhyan region, to assess the N-mineralization, nitrification and size of viable community of ammonium- and nitrite-oxidizing bacteria as affected by different sites and seasons. Six different ecosystems (four forests and two savannas), which differ in terms of topography, vegetation and moisture status, were selected for the present study. The soils of the study sites differ significantly in its physico-chemical properties. The savanna site had significantly higher pH (7.2), bulk density (1.37 g cm(-3)) and silt content (67.80%) but lower water holding capacity (1.37%), total-C (16,356 microg g(-1) dry soil), N (1090 microg g(-1) dry soil) and P (213 microg g(-1) dry soil) than forest sites. The soil moisture content, N-mineralization, nitrification rates and numbers of ammonium- and nitrite-oxidizing bacteria were highest in the wet season and lowest in dry season, while the size of mineral-N (NH4(+)-N and NO3(-)-N) showed a reverse trend at the sites. The N-mineralization, nitrification and nitrifier population size differ significantly across the site and season. The numbers of free-living cells of ammonium- and nitrite-oxidizing bacteria were significantly related to each other and to N-mineralization, nitrification, soil moisture and mineral-N components. The N-mineralization, nitrification and the viable number of nitrifying cells were consistently higher for forest soils compared to savanna sites. It was concluded that soil microbial process (N-mineralization and nitrification) and nitrifier population size were dependent on site topography, vegetation cover and soil moisture status.     

Vegetation analysis and tree population structure of tropical wet evergreen forests in and around Namdapha National Park,northeast India

Species composition, diversity and tree population structure were studied in three stands of the tropical wet evergreen forest in and around Namdapha National Park, Arunachal Pradesh, India. Three study stands exposed to different intensities of disturbances were identified, viz., undisturbed (2.4 ha) in the core zone of the park, moderately disturbed (2.1 ha) in the periphery of the park and highly disturbed (2.7 ha) outside the park area. In total 200 plant species belonging to 73 families were recorded in three stands. Tree density and basal area showed a declining trend with the increase in disturbance intensity. The densities of tree saplings and seedlings were lower in the disturbed stands than in the undisturbed stand. Species like Altingia excelsa, Olea dioica, Terminalia chebula, Mesua ferrea and Shorea assamica in the undisturbed stand and Albizia procera alone in the moderately disturbed stand contributed more than 50% of the total tree density in respective stands. The undisturbed stand contained young tree population. In the highly disturbed stand, the tree density was scarce, but had uncut trees of higher girth class (>210 cm GBH). Low shrub density was recorded in both disturbed stands due to frequent human disturbances; the broken canopy and direct sunlight enhanced the abundance of herbs in these stands. With a species rarity (species having <2 individuals) of ca. 50%, the tropical wet evergreenforests of the Namdapha National Park and its adjacent areas warrant more protection from human intervention and also eco-development to meet the livelihood requirements of the local inhabitants in the peripheral areas of the Namdapha National Park in order to reduce the anthropogenic pressure on the natural resources of the park.     

Forest structure and carbon dynamics in Amazonian tropical rain forests

Living trees constitute one of the major stocks of carbon in tropical forests. A better understanding of variations in the dynamics and structure of tropical forests is necessary for predicting the potential for these ecosystems to lose or store carbon, and for understanding how they recover from disturbance. Amazonian tropical forests occur over a vast area that encompasses differences in topography, climate, and geologic substrate. We observed large differences in forest structure, biomass, and tree growth rates in permanent plots situated in the eastern (near Santarém, Pará), central (near Manaus, Amazonas) and southwestern (near Rio Branco, Acre) Amazon, which differed in dry season length, as well as other factors. Forests at the two sites experiencing longer dry seasons, near Rio Branco and Santarém, had lower stem frequencies (460 and 466 ha^–1 respectively), less biodiversity (Shannon–Wiener diversity index), and smaller aboveground C stocks (140.6 and 122.1 Mg C ha^–1) than the Manaus site (626 trees ha^–1, 180.1 Mg C ha^–1), which had less seasonal variation in rainfall. The forests experiencing longer dry seasons also stored a greater proportion of the total biomass in trees with >50 cm diameter (41–45 vs 30% in Manaus). Rates of annual addition of C to living trees calculated from monthly dendrometer band measurements were 1.9 (Manaus), 2.8 (Santarém), and 2.6 (Rio Branco) Mg C ha^–1 year^–1. At all sites, trees in the 10–30 cm diameter class accounted for the highest proportion of annual growth (38, 55 and 56% in Manaus, Rio Branco and Santarém, respectively). Growth showed marked seasonality, with largest stem diameter increment in the wet season and smallest in the dry season, though this may be confounded by seasonal variation in wood water content. Year-to-year variations in C allocated to stem growth ranged from nearly zero in Rio Branco, to 0.8 Mg C ha^–1 year^–1 in Manaus (40% of annual mean) and 0.9 Mg C ha^–1 year^–1 (33% of annual mean) in Santarém, though this variability showed no significant relation with precipitation among years. Initial estimates of the C balance of live wood including recruitment and mortality as well as growth suggests that live wood biomass is at near steady-state in Manaus, but accumulating at about 1.5 Mg C ha^–1 at the other two sites. The causes of C imbalance in living wood pools in Santarém and Rio Branco sites are unknown, but may be related to previous disturbance at these sites. Based on size distribution and growth rate differences in the three sites, we predict that trees in the Manaus forest have greater mean age (~240 years) than those of the other two forests (~140 years).     

Soluble reactive phosphorus (SRP) transport and retention in tropical, rain forest streams draining a volcanic landscape in Costa Rica: in situ SRP amendment to streams and laboratory studies

Soluble reactive phosphorus (SRP) transport/retention was determined in two rain forest streams (Salto, Pantano) draining La Selva Biological Station, Costa Rica. There, SRP levels can be naturally high due to groundwater enriched by geothermal activity within the surfically dormant volcanic landscape, and subsequently discharged at ambient temperature. Combined field and laboratory approaches simulated high but natural geothermal SRP input with the objective of estimating the magnitude of amended SRP retention within high and low SRP settings and determining the underlying mechanisms of SRP retention. First, we examined short-term SRP retention/transport using combined SRP-conservative tracer additions at high natural in situ concentrations. Second, we attempted to observe a DIN response during SRP amendment as an indicator of biological uptake. Third, we determined SRP release/retention using laboratory sediment assays under control and biologically inhibited conditions. Short-term in situ tracer-SRP additions indicated retention in both naturally high and low SRP reaches. Retention of added SRP mass in Upper Salto (low SRP) was 17% (7.5 mg-P m⁻² h⁻¹), and 20% (10.9 mg-P m⁻² h⁻¹) in Lower Salto (high SRP). No DIN response in either nitrate or ammonium was observed. Laboratory assays using fresh Lower Salto sediments indicated SRP release (15.4 ± 5.9 μg-P g dry wt.⁻¹ h⁻¹), when incubated in filter sterilized Salto water at ambient P concentration, but retention when incubated in filter sterilized river water amended to 2.0 mg SRP l⁻¹ (233.2 ± 5.8 μg-P g dry wt.⁻¹ h⁻¹). SRP uptake/release was similar in both control- and biocide-treated sediments indicating predominantly abiotic retention. High SRP retention even under biologically saturated conditions, absence of a DIN response to amendment, patterns of desorption following amendment, and similar patterns of retention and release under control and biologically inhibited conditions all indicated predominantly abiotic P flux.     

Woody plants diversity,floristic composition and land use history in the Amazonian rain forests of Madidi National Park,Bolivia

A floristic inventory of woody plants was carried out to analyse the relationships between floristic similarity and geographical distance, and to compare the effect of land use history on the floristic composition between sites. Three lowland and two submontane sites were studied in Madidi, Bolivia. In one site, there is evidence of an Inca ruin. A total of 877 species and 12,822 individuals of woody plants with a diameter at breast height ≥2.5 cm were recorded in 44 0.1–ha plots. Fisher’s Alpha index values were slightly higher for the lowlands than for the submontane. Floristic similarity was higher within sites than between sites as measured by both Sørensen and Steinhaus indexes. The fact that the 30 most important species per site (totalling 94 species) accounted for 61.7% of total individuals, support the hypothesis that Amazonian plant communities are dominated by a limited set of species, genera and families. On the other hand, 18 out of the 94 species were reported in a single site, suggesting that some species are patchy in distribution and may be environmentally determined. Both the oligarchy and environmental-determinism hypotheses can be complementary in order to understand floristic patterns of this region. The Ruins submontane site is floristically the most distinct, and past human disturbance is likely to be the main reason. Since species diversity (ranging from 53 to 122 species per plot) and density (ranging from 157 to 503 per plot) are highly variable in Madidi, to characterize the diversity of a site, it is necessary to quantify an average of 10 0.1-ha plots in a relatively small geographical area.     

Bat assemblage structure in two dry forests of Colombia: Composition, species richness, and relative abundance

We studied the composition, species richness, and relative abundance of bat assemblages in the Colombian dry forests of Chicamocha and Patía. In Chicamocha, 11 bats of the family Phyllostomidae were captured with mist-nets, corresponding to 85–100% of the potential phyllostomids species in the area. Two bats of the family Vespertilionidae were also captured in Chicamocha. In Patía, 12 species were captured with mist-nets, all Phyllostomidae, representing 72–100% of the estimated total number of species in the zone. Minor differences in number of species and composition were detected among sampling periods in Chicamocha. The most common species in this dry forest were Glossophaga longirostris and Sturnira lilium. In Patía, notable differences in the number of species and composition were observed among sampling periods, and the most common species were Artibeus jamaicensis, Carollia perspicillata and Phyllostomus discolor. Arid-zone dwelling bats were absent in Patía and we suggest that this absence may be associated with the isolation of Patía from other northern dry zones of Colombia since Quaternary times. There was also low abundance of bats in Patía, which appears to be related to human disturbance. The most abundant phyllostomid bat species in the two dry forests studied are those that include fruit and/or nectar-pollen from columnar cacti as an important proportion of their diets.     

Dew and its effect on two heliophile understorey species of a tropical dry deciduous forest in Mexico

 A series of horizontal and vertical measurements of dew deposition and dew duration were carried out in a tropical dry deciduous forest in western Mexico (19° 30′ N, 105° 03′ W). The effect of dew on transpiration in heliophile species was also investigated. The amount of dew was very variable with no temporal or spatial pattern. The amount of dew measured at two horizontal transects (maximum and minimum) (at heights of 0.20 m and 1.30 m above ground level) was from 0.014 to 0.203 mm and from 0.013 to 0.061 mm in the middle and at the end of the dry season, respectively. Dew deposition at different vertical levels (0.50–12.5 m height) ranged from 0.04 to 0.36 mm. The duration of dew formation ranged between 60 and 129 min after sunrise above the canopy (a height of 11 m), and between 259 and 290 min after sunrise at a height of 2 m. Daily transpiration rates were 883 and 632 g m^–2 d^–1 in Coccoloba liebmannii and 538 and 864 g m^–2 d^–1 in Jacquinia pungens in January and April, respectively. Transpiration was restricted from sunrise to early afternoon in April, as a result of the pronounced midday closure of stomata in both species. The reduction of transpiration by dew ranged from 13.2 to 50.1 g m^–2 d^–1 and from 4.5 to 77.7 g m^–2 d^–1 for C. liebmannii and J. pungens, respectively. Dew can play an important role in enhancing the survival of heliophile species in the dry season by reducing transpiration rates during the morning. Received: 30 April 1998 / Accepted: 14 December 1998     

The soil C pool in different agroecosystems derived from the dry tropical forest of Guanacaste, Costa Rica

Inventories of soil C pools are still lacking from tropical sites. Our objective was to assess total C and N concentrations in the different mineral soil fractions down to 50 cm depth in relation to selected physical and chemical properties of 5 ecosystems at La Flor Sustainable Center in Guanacaste, Costa Rica. The ecosystems studied were a derived savanna with scattered trees, a gallery forest, an abandoned Mango indigofera L. plantation, a Citrus sp. plantation, and a Saccharum officinarum L. (sugarcane) plantation. Significant differences were found for the main fixed factor ecosystem for all variables analyzed (ANOVA). The TSC concentration was significantly higher in the sugarcane plantation compared to the rest of land use systems. The TSC concentration decreased significantly with increase in depth in all ecosystems and ranged from 20.3–38.3 to 4.3–20.9 g kg⁻¹ in the 0–10 and 40–50 cm depth, respectively. In all cases, the clay + silt fraction (<50 μm) contained the highest C concentration. N concentration (0–10 cm depth) at La Flor ranged from 0.32 to 0.19%, and decreased in the order sugarcane > Curatella savanna > Mango and Citrus plantations > gallery forest. A principal component analysis (PCA) performed with all variables studied showed that the ordination of land uses (ecosystems) in the factorial plane defined by the first two axes was significant (Monte Carlo permutation test, P < 0.0001). The highest TSC pool down to 50 cm depth was obtained in the sugarcane plantation (160 Mg C ha⁻¹) while less C was found in the rest of ecosystems, i.e. from 66 (gallery forest) to 80 Mg C ha⁻¹ (Curatella savanna). The TSC concentration obtained in the sugarcane plot is likely the result of the incorporation of surface residues into the soil that would have otherwise been lost through burning, which is the current practice in the region. Further studies on C stabilization in the clay fraction are thus needed to test the hypothesis of soil C enrichment due to residue management. Finally, trade-offs are to be considered for both preservation of the fragile dTf and the productivity of derived land uses that increases soil C at the same time.