An assessment of the variation of soil properties with landscape attributes in the highlands of Cameroon
Abstract
Soil properties are useful for assessing the potential of landscapes to provide terrestrial ecosystem services, but they are affected by anthropogenic activities and environmental factors including landscape attributes. This study assessed how soil properties are influenced by landscape attributes and their interactions in the highlands of Cameroon using the Land Degradation Surveillance Framework as a data collection tool. Soil properties (soil organic carbon [SOC], clay content, exchangeable bases [ExBas], electric conductivity [EC], boron [B], manganese [Mn], phosphorus [P], pH) were quantified within classes of landscape attributes. Soil samples were collected on 160 (1,000 m2) plots randomly located in a sentinel site of 100 km2 and were analyzed using a combination of conventional laboratory methods and mid-infrared spectroscopy. Soil properties were highly affected by soil depths, land use types, slope gradients, and topographic positions, but less by their interactions. Significant interactions existed between land use types and topographic positions for SOC, EC, ExBas, and pH, and between slope gradients and topographic positions for pH, whereas Mn was influenced by the interaction between land use types and slope gradients. Most soil properties were higher in low altitude plots and those with higher vegetation cover but decreased in the upslope direction. The pH and clay contents were less affected by slope gradient confirming the inherent nature of the properties. These results are useful for site-specific implementation of ecological intensification in areas with complex topography such as the highlands of Cameroon, offering a reference for future ecological policies and landscape restoration.
1 INTRODUCTION
The soil's potential to supply ecosystem services is generally assessed and expressed in relation to its functional properties, which are determined based on the interactions between various soil forming factors and processes (Bockheim, Gennadiyev, Hartemink, & Brevik, 2014; Józefowska, Woś, & Pietrzykowski, 2016). Soil properties are influenced by multiple physical and biochemical interactions driven by anthropogenic activities and biophysical characteristics of the landscape known as landscape attributes including land use types, topographic positions, slope gradients, and soil depths (Fernández-Romero, Lozano-García, & Parras-Alcántara, 2014; Mora, Guerra, Armas-Herrera, Arbelo, & Rodríguez-Rodríguez, 2014). Various landscape attributes differed in their effects on soil properties useful to assess the productive potential of a landscape (Sun, Zhu, & Guo, 2015). These attributes influence the processes and intensities of erosion and sediment redistribution (Dessalegn, Beyene, Ram, Walley, & Gala, 2014; Sun, Shao, Liu, & Zhai, 2014) and alter the production and decomposition of plant litter due to variation in hydrological processes and weather conditions (Fernández-Romero et al., 2014; Wang, Wang, Cao, Bai, & Qin, 2016). Furthermore, landscape attributes impact organic fractions and soil properties at various topographic positions (Ritchie, McCarty, Venteris, & Kaspar, 2007; Sun et al., 2014), even when the soils are derived from the same parent materials and have a single climate regime (Ata Rezaei & Gilkes, 2005).
Several studies have examined how soil properties are influenced by various landscape attributes and have reported significant variations (Arnhold et al., 2015; Huang, Liu, & An, 2015; Lybrand & Rasmussen, 2015; Takoutsing, Weber, Tchoundjeu, & Shepherd, 2015). A slight increase in slope gradient affects soil properties, leads to changes in land cover and vegetation types (Griffiths, Madritch, & Swanson, 2009; Ritchie et al., 2007; Sun et al., 2014), and affects crop yields (Ladoni, Basir, Robertson, & Kravchenko, 2016; Muñoz, Steibel, Snapp, & Kravchenko, 2014). Despite the reported influence of landscape parameters and the resulting consequences on crop yields and ecosystem services, detailed assessment of the variation of soil properties with landscape attributes has not been extensively investigated in the highlands of Cameroon. Thorough statistical bases for the management of soil properties in heterogeneous agricultural and mosaic landscapes are still rarely available in tropical areas (Wang, Fu, Qiu, & Chen, 2001). Demand for agricultural products will increase to feed the ever increasing human population, accommodate the changes in diets and the increase in bioenergy needs (Krausmann et al., 2013). Ecological intensification has been endorsed as part of the solution, but if not well implemented, it may put more pressure on the soil's capacity to fulfill its functions, leading to the degradation of ecosystems and decrease in productivity in the long term (Trivedi, Delgado-Baquerizo, Anderson, & Singh, 2016).
There is actually no single generalizable model of ecological intensification due to variation in soil properties. Any generalization in practice would be contrary to the context-specific and ecosystem-based approaches of ecological intensification (Doré et al., 2011). For complex topographic landscape such as the highlands of Cameroon, models of ecological intensification may include practices such as agroecology, organic agriculture, diversified farming systems, climate smart agriculture, nature mimicry, evergreen agriculture, agroforestry, and conservation agriculture (Tittonell, 2014). These practices will definitely differ in their implementation depending on the influence of landscape attributes. In addition, local knowledge on land management and traditional farming systems in areas with complex topography is essential in designing site-specific and contextualized components of ecological intensification (Lahmar, Bationo, Dan Lamso, Guéro, & Tittonell, 2012). Therefore, there is a need to comprehend the degree of variations in soil properties related to landscape attributes. The objectives of the current study were to (a) assess the effects of landscape attributes (soil depths, land use types, slope gradients, and topographic positions) on selected soil properties (soil organic carbon [SOC], clay content, electrical conductivity, exchangeable bases [ExBas], boron, manganese, phosphorous, and soil reaction [pH]) and (b) measure the interactive effects of the attributes for evidence-based and informed land management decisions for complex topographic landscapes. We hypothesized that mean values for soil properties would be higher in plots with higher vegetation cover and lower altitudes. Our hypothesis is based on the premises that plots with higher vegetation cover have higher nutrient inputs and a higher content of soil organic matter, and nutrients in the upland plots are eroded by runoff and deposited in the lower plots.
2 MATERIALS AND METHODS
2.1 Study site and experimental design
The study site is located in the western highlands of Cameroon (Figure 1). The area is characterized by diverse and mosaic landscape with complex topography, varying vegetation patterns, heterogeneous attributes and is dominated by intense agricultural and livestock activities. The elevation of the site ranged from 1,300 to 1,700 m, and mean annual precipitation ranged from 1,500 to 1,800 mm. The vegetation is savannah type with montane forests and patches of montane grassland, and subalpine grasslands and shrublands located in waterways, watersheds, and protected areas. The cropping systems can best be described as multiple cropping with mixed inter-cropping of crops and trees, and crop yields vary greatly with management practices and the location of the plot along the toposequence. Data were collected using the Land Degradation Surveillance Framework, which is designed to provide a biophysical baseline at the landscape level, and a monitoring and evaluation framework for assessing land degradation and recovery over time (Vågen, Winowiecki, Tamene, & Tondoh, 2010). The experimental design was based on a hierarchical survey approach in a site of 10 × 10 km (100 km2). The site was subdivided into 16 clusters, and 10 sampling plots (1,000 m2 each) were randomly distributed in each cluster, giving a total of 160 sampling plots. Each sampling plot contained four subplots (100 m2 each; Takoutsing et al., 2016; Vågen et al., 2010).

Four landscape attributes were selected for this study: soil depth, land use type, slope gradient, and topographic position. These attributes were selected because previous research has demonstrated that they can significantly affect soil properties (Fernández-Romero et al., 2014; Mora et al., 2014; Ritchie et al., 2007; Sun et al., 2014; Takoutsing et al., 2015; Takoutsing et al., 2017). Data were collected at the levels of plots and subplots. The 160 sampling plots were characterized in terms of geographical location (longitude, latitude, and elevation using a GPS), land use type (classes = cropland, fallow, forest, grassland, or pasture), slope gradient (classes = level [<10%], sloping [10–20%], or steep [>20%]), and topographic position (classes = bottomland, footslope, midslope, ridge/crest, or upland). Data for trees (woody plants with height > 3 m) and shrubs (woody plants with height 1.5–3 m) were recorded at the subplot level. Tree and shrub densities were estimated by counting all stems in the subplot. Height and diameter at breast height were measured on trees, and height, crown length, and width were measured on shrubs. Percent cover of woody and herbaceous vegetation was rated on a 6-point scale: absent, <4%, 4–15%, 15–40%, 40–65%, and > 65%. Soil samples were collected at two depths (topsoil [0–20 cm] and subsoil [20–50 cm]) in the subplots using an auger. For each depth, the samples from the four subplots were pooled together and thoroughly mixed to obtain the homogenous composite sample for each of the 160 sampling plots. Three subsoil samples were not obtained because of depth restriction, so the total number of soil samples was 317 (160 for topsoil and 157 for subsoil).
2.2 Soil analytical procedures
Conventional laboratory analyses methods were used to analyze 10% of the samples for SOC, texture, ExBas, electric conductivity (EC), boron (B), manganese (Mn), phosphorus (P), and soil reaction (pH). These were previously selected as important site-specific soil properties for the study site, and they can be used to rapidly assess the quality of soils to guide land management decisions and indirect measures of soil functions (Takoutsing et al., 2016). The pH was analyzed in a 1:1 H2O solution. Exchangeable bases, phosphorous, manganese, and boron were analyzed using a Melich-3 extraction procedures, whereas EC was determined using a 1:2 soil : water extract at the Crop Nutrition Laboratory (www.cropnuts.com) in Nairobi, Kenya. Soil organic carbon and total carbon (%) were determined by chromic acid digestion and spectrophotometric analysis. Soil texture was measured with a laser diffraction particle size analyzer at the World Agroforestry Centre Plant and Soil Spectroscopy Laboratory in Nairobi, Kenya. Then all samples were analyzed using mid-infrared reflectance based on methods described in Terhoeven-Urselmans, Vagen, Spaargaren, and Shepherd (2010) and Hengl et al. (2015). Infrared spectroscopy has been demonstrated to be a rapid, affordable, and non-destructive method for analyzing a range of materials (Reeves & Smith, 2009; Shepherd & Walsh, 2002; Viscarra Rossel, DJJ, AB, Janik, & Skjemstad, 2006).
2.3 Statistical analyses
Descriptive statistics (mean, median, standard deviation, etc.) were calculated for the soil properties. The general linear model was used to test the effects of landscape attributes on soil properties. The general linear model included the following sources of variation: soil depths, land use types, slope gradients, and topographic positions as main effects (treated as fixed factors), and two-way and three-way interactions between the main effects. Significant differences in means of soil properties between the classes of each landscape attribute were assessed by Tukey's honestly significant difference test. These analyses were done using R software version 2.15.0 (R Development Core Team, 2013).
Relationships between soil properties and landscape attributes were further explored using redundancy analysis (RDA). Developed by Hotelling (1936), RDA has been considerably used in ecology (Bowles, Acosta-Martínez, Calderón, & Jackson, 2014; Campos-Herrera, El-Borai, Rodríguez Martín, & Duncan, 2016; Gutiérrez et al., 2016; Rodríguez Martín et al., 2014; Rousseau, Rioux, & Dostaler, 2006) to assess relationships between a set of dependent variables and a set of explanatory variables. One of the advantages of RDA is that it integrates the techniques of ordination and multiple regression (Borcard, 2006; Legendre & Legendre, 1998; van den Wollenberg, 1977). In this study, RDA was used to identify the combination of landscape attributes that best explains the variation in soil properties. Canonical ordinations are usually carried out with standardized explanatory variables, which are not always dimensionally homogeneous (Borcard, 2006). In this context, the soil properties were transformed to standard normal variables with mean = 0 and variance = 1 (transformed variable = [untransformed variable-mean]/standard deviation). The results of RDA were graphically presented with a bi-plot of the dependent variables (soil properties) and the explanatory variables (landscape attributes). RDA was done using XLSTAT software (Addinsoft Version 2012.2.02).
3 RESULTS AND DISCUSSION
3.1 Ecological characterization
The study site is a mosaic landscape with diverse physical features and ecological conditions that influence soil properties. Land suitable and favorable for agricultural production based on slope classification (slope < 10%) accounts for 44% of the site. The distribution of land use types based on the 160 observations plots illustrates that 88% of the area is either cultivated or under fallow, suggesting that farmers exploit nearly every hectare of land, including those with steep slopes (slope > 20%). Uncultivated plots representing 12% of the area are either unsuitable for agriculture, protected areas, have very steep slopes, or are not easily accessible to farmers. The annual herbaceous cover including crops varied between 15% and 40% for cultivated plots and between 40% and 65% for uncultivated plots. Mean tree and shrub densities were estimated at 140 trees ha−1 and 190 shrubs ha−1 whereas the median values were lower (134 trees ha−1 and 159 shrubs ha−1), indicating variation among land use types. Higher tree and shrub densities were found in uncultivated plots. The high woody cover observed across the area is due to the fact that trees are maintained or planted for a range of products and services including food, farm demarcation, land ownership, and as a legacy for future generations.
3.2 Analysis of variance of soil properties
Analysis of variance (Table 1) indicated that the majority of soil properties were highly affected by all the landscape attributes, but less affected by their interactions. The effects of landscape attributes on soil properties were not significant in only four cases: soil depths on pH, slope gradients on EC and pH, and topographic positions on B. Significant interactions existed between land use types and topographic positions for SOC, EC, ExBas, and pH, between slope gradients and topographic positions for pH, and between land use types and slope gradients for Mn. These interactions mean that the effects of some landscape attributes on these soil properties differed depending on the other landscape attributes. For example, pastures at midslope positions had lower values of SOC compared with upland plots probably due to the effects of erosion, and croplands in the bottomlands and footslopes had higher values of SOC compared with croplands located in the ridge/crest. Fallows had higher values of SOC at bottomland and footslope positions and lower values at upland positions (Figure 2).
Sources of variationa | dfb | Soil propertiesc, probability of F ratioe, and F valuesd | |||||||
---|---|---|---|---|---|---|---|---|---|
SOC (%) | Clay (%) | EC (dS m−1) | ExBas (cmol kg−1) | B (mg kg−1) | Mn (mg kg−1) | P (mg kg−1) | pH | ||
Soil depths (D) | 1 | *** (30.16) | *** (86.87) | *** (69.45) | *** (24.67) | *** (16.66) | *** (11.68) | *** (40.84) | ns (0.56) |
Land use types (L) | 4 | *** (11.02) | *** (6.64) | *** (17.76) | *** (14.04) | *** (16.66) | ** (4.14) | *** (12.37) | *** (11.74) |
Slope gradients (S) | 2 | *** (8.56) | * (3.25) | ns (0.88) | * (4.04) | * (3.40) | *** (34.01) | *** (7.40) | ns (1.06) |
Topographic positions (T) | 4 | *** (9.35) | ** (4.58) | *** (5.16) | ** (4.22) | ns (1.56) | ** (4.28) | * (2.45) | * (3.03) |
D × L | 4 | ns (0.09) | ns (0.25) | ns (0.12) | ns (0.16) | ns (0.18) | ns (0.39) | ns (0.15) | ns (0.45) |
D × S | 2 | ns (0.10) | ns (2.24) | ns (0.47) | ns (0.24) | ns (0.22) | ns (0.40) | ns (0.39) | ns (0.50) |
D × T | 4 | ns (0.53) | ns (0.26) | ns (0.24) | ns (0.84) | ns (0.43) | ns (0.47) | ns (0.34) | ns (0.63) |
L × S | 5 | ns (0.79) | ns (1.56) | ns (2.21) | ns (1.26) | ns (0.13) | *** (6.82) | ns (1.89) | ns (0.49) |
L × T | 5 | *** (4.46) | ns (0.91) | * (2.46) | ** (3.47) | ns (0.39) | ns (0.86) | ns (0.62) | ** (3.71) |
S × T | 3 | ns (0.95) | ns (2.20) | ns (1.36) | ns (0.82) | ns (0.88) | ns (0.79) | ns (0.42) | * (3.51) |
D × L × S | 5 | ns (0.17) | ns (0.27) | ns (0.34) | ns (0.38) | ns (0.35) | ns (0.15) | ns (0.15) | ns (0.46) |
D × L × T | 5 | ns (0.29) | ns (1.87) | ns (0.77) | ns (0.44) | ns (0.37) | ns (0.20) | ns (0.26) | ns (0.32) |
D × S × T | 3 | ns (0.08) | ns (0.80) | ns (0.16) | ns (0.40) | ns (0.11) | ns (0.18) | ns (0.16) | ns (0.65) |
- a Sources of variation—Soil depths (D): topsoil (0–20 cm) and subsoil (20–50 cm). Land use types (L): cropland, fallow, forest, grassland, and pasture. Topographic positions (T): bottomland, footslope, midslope, ridge/crest, and upland. Slope gradients (S): level, sloping, and steep slope; D × L, D × S, L × S, L × T = interactions.
- b df: degrees of freedom for numerator of F ratio (some combinations of treatment levels were not observed, so the numerator degrees of freedom for some interactions was lower than expected); df for denominator of F ratio = 289.
- c Soil properties—SOC: soil organic carbon, EC: electrical conductivity, ExBas: exchangeable bases, B: boron, Mn: manganese, P: phosphorous, pH: pH; values for all soil properties analyzed on original untransformed scales.
- d Number in parentheses are F-values.
- e Probability of F ratio for testing source of variation.
- * p < 0.05, ns p > 0.05.
- ** p < 0.01.
- *** p < 0.001.

In general, a consistent trend was observed for soil depths (Table 2), as all indicators with the exception of clay content were significantly higher in the topsoil. This is probably the result of higher organic matter content (not measured in the study) or the fine texture of the clay-rich soils that slows percolation and reduces leaching (Yimer, Ledin, & Abdelkadir, 2007). In addition, the influence of biological activities (fauna and flora) and management (additional inputs such as organic and inorganic fertilizers) are more pronounced on the upper soil layers, resulting in higher values for most nutrients.
Landscape attributes | Classes | Soil propertiesa | |||||||
---|---|---|---|---|---|---|---|---|---|
SOC (%) | Clay (%) | EC (dS m−1) | ExBas (cmol kg−1) | B (mg kg−1) | Mn (mg kg−1) | P (mg kg−1) | pH | ||
Soil depths | Topsoil (n = 160) | 3.35 b | 67.67 a | 0.13 b | 9.60 b | 0.04 b | 17.34 b | 6.47 b | 5.78 a |
Subsoil (n = 157) | 2.84 a | 73.82 b | 0.09 a | 7.95 a | 0.03 a | 14.81 a | 4.70 a | 5.77 a | |
Land use types | Cropland (n = 156) | 3.09 b | 72.29 b | 0.10 b | 8.96 b | 0.04 a | 14.86 a | 5.43 b | 5.82 b |
Fallow (n = 132) | 3.23 b | 69.47 a | 0.11 b | 9.03 b | 0.04 a | 16.06 a | 5.66 b | 5.80 b | |
Forest (n = 8) | 3.57 b | 65.09 a | 0.17 c | 10.58 b | 0.06 b | 24.55 b | 8.88 c | 5.75 b | |
Grassland (n = 4) | 4.17 b | 62.76 a | 0.21 c | 12.96 b | 0.08 b | 24.49 b | 11.24 c | 5.84 b | |
Pasture (n = 17) | 1.64 a | 70.53 ab | 0.06 a | 3.44 a | 0.03 a | 21.60 ab | 3.62 a | 5.38 a | |
Slope gradients | Level (n = 46) | 3.38 b | 68.90 a | 0.12 a | 10.17 b | 0.043 ab | 16.37 a | 6.43 b | 5.86 a |
Sloping (n = 242) | 3.12 b | 71.33 a | 0.11 a | 8.66 a | 0.038 a | 14.63 a | 5.33 a | 5.78 a | |
Steep (n = 29) | 2.43 a | 68.52 a | 0.10 a | 7.60 a | 0.051 b | 27.76 b | 6.46 b | 5.59 a | |
Topographic positions | Bottomland (n = 52) | 3.44 c | 68.71 a | 0.12 b | 10.01 c | 0.04 a | 15.67 b | 6.10 b | 5.85 b |
Footslope (n = 34) | 3.67 c | 69.01 a | 0.12 b | 9.97 bc | 0.03 a | 10.74 a | 5.50 ab | 5.88 b | |
Midslope (n = 189) | 2.99 b | 70.93 a | 0.10 ab | 8.44 ab | 0.04 a | 17.51 b | 5.67 ab | 5.74 a | |
Ridge/Crest (n = 26) | 2.42 a | 74.95 b | 0.08 a | 7.16 a | 0.03 a | 13.83 ab | 4.21 a | 5.75 ab | |
Upland (n = 16) | 3.05 abc | 71.46 ab | 0.11 ab | 8.96 ab | 0.03 a | 15.64 ab | 5.50 ab | 5.80 ab |
- a Soil properties—SOC: soil organic carbon, EC: electrical conductivity, ExBas: exchangeable bases, B: boron, Mn: manganese, P: phosphorous, pH: pH; values for all soil properties analyzed on original untransformed scales; means with the same letter are not significantly different (p > 0.05) and those with different letters are significantly different (p < 0.05) based on Tukey's honestly significant difference test.
Mean values of most soil properties with the exception of SOC tended to be higher in forests and grasslands compared with other land use types and in plots located in bottomlands and footslopes compared with other topographic positions (Table 2). The lower values observed in croplands and pasture are attributed to the influence of anthropogenic activities such as farming and intensive grazing that reduce organic matter inputs into the soils and reduce vegetation cover (Takoutsing et al., 2013). Higher clay content observed in croplands and pastures is due to reduced vegetation cover caused by anthropogenic activities including agriculture and livestock rearing (Liu, Zhang, Chang, Kan, & Lin, 2012). Clay content did not differ significantly between bottomland and midslope positions, but was higher in ridge/crest and upland positions, probably as the result of the effects of water erosion.
Differences observed in soil properties with topographic positions may be attributed to the combined effects of topography on soil moisture content, depth of soil formation, erosion, and deposition processes. In the study area, different topographic positions have similar vegetation cover, so variation in the values of soil properties at different positions is attributed to differences in soil erosion rates (not quantified in this study), or soil development and deposition processes. Nutrients in the ridge/crest plots are eroded by runoff and deposited in the footslopes and bottomlands, therefore influencing the nutrient concentrations (Figure 3).

The effects of slope gradients on soil properties were inconsistent and did not agree with our hypothesis. Some soil properties were higher on steep plots (e.g., B and Mn) whereas others were higher on level plots (e.g., SOC and ExBas). From observations, slope gradient classes have similar vegetation cover, and land use types are distributed irrespective of the slope. It was generally observed that the intensity and type of management applied to a plot depended on its accessibility by farmers or animals and its proximity to the households. Perhaps the accessibility and proximity to households account for the inconsistent results, but we did not quantify or test these factors in this study.
3.3 Relationships between soil properties and landscape attributes
The RDA revealed several sets of significant relationships between soil properties and landscape attributes (Figure 4). The first two ordination axes (F1 and F2) explained about 92% of the effects of landscape attributes on soil properties (66% from the first axis and 26% from the second axis). The black arrows represent soil properties, and the different color points represent the landscape attributes. The arrows indicate the direction of the change in soil properties, and the lengths of the arrows indicate the extent to which the soil parameters are affected by the landscape attributes. In RDA, most of the soil properties were related to the first axis in quadrants Q3 and Q4, whereas clay content was associated with quadrant Q2 and negatively related to the rest of the indicators (Figure 4). This can be interpreted by the fact that clay content is influenced by soil depth with high values in the subsoil and by the local-scale influences associated with land use (cropland). According to the groupings of RDA, soil properties were divided to assess the effects of landscape attributes: These groups are discussed below.

3.3.1 Soil organic carbon, electric conductivity, and exchangeable bases
SOC, EC, and ExBas were higher in forests and grasslands than in the other land use types (Table 2). These results were expected because most of the soil properties are coupled with organic matter content and are consistent with other studies that assessed the effects of explanatory variables on soil properties (Arnhold et al., 2015; Huang et al., 2015; Lybrand & Rasmussen, 2015; Takoutsing et al., 2015). The higher concentration of SOC, ExBas, and also P in the topsoil, level, and steep plots, bottomlands/footslopes, and grasslands (Table 2) is an indication of higher organic matter content, dominance of roots, humus, and associated organisms responsible for biological activities. The accumulation of organic matter at the bottomland positions is favored by runoff and erosion coupled with wetter conditions. The significantly higher concentration of ExBas in the topsoil probably reflects three factors: Productivity of some plots is improved by fertilizer inputs; vegetation can pump ExBas from subsoil to topsoil layers; ExBas is almost immobile and loss through leaching should be relatively low (Yimer et al., 2007). The highest values for EC observed in grasslands and forests, bottomlands, and footslopes, as well as topsoil (Table 2), implied that EC is related to soil organic matter content. In contrast, the effect of slope gradients on EC was not significant (Table 2), indicating that there is no major difference in cumulative salt accumulation along the slope and that this property is not influenced by the effects of erosion and leaching processes. The low value for EC in pastures is probably due to less return of organic matter inputs and low tree densities caused by intensive grazing (Nesbitt & Adl, 2014). Soil organic matter is estimated by the determination of SOC content, which refers to the carbon component of organic compounds. Because soil organic matter presents more challenges for direct measurement, most laboratories prefer to focus their reports on SOC (Nelson & Sommer, 1982; Schumacher, 2002).
3.3.2 Phosphorous, manganese, and boron
Phosphorous is an important nutrient for plant development, and P deficiency is a limiting factor for crop production. Phosphorous concentration in agricultural soils is improved through fertilizer applications. However, in the highlands of Cameroon, higher concentrations of P are instead found in forests and grasslands (Table 2 and Figure 4). The low concentration and P in croplands can be attributed to the low application of fertilizers in farming systems (Takoutsing et al., 2016). The low application is generally attributed to the high cost of fertilizers and the low purchasing power of the smallholder farmers. In the highlands of Cameroon, P losses from soil are generally through erosion, leaching, and the uptake by plants and removal with harvests. Erosive P losses at one site may result in P inputs at other sites due to topography. Typically, this leads to P-depleted upslope and summit positions in fields and P-enriched downslope positions and valleys, the latter often close to vulnerable watercourses. On the other hand, P fixation tends to be higher in clay-rich soils (Dessalegn et al., 2014; Havlin, Beaton, Tisdale, & Nelson, 1999), and most P is bound either in nonliving organic matter or in the soil microbial biomass, as shown by its association with SOC in F1 of the RDA (Figure 4). Phosphorous added to the soil as a fertilizer will considerably increase the concentration and the readily available pool available for crop uptake. However, higher P applications can precipitate out of solution through reactions with charged molecules and soil particles, becoming part of the fixed pool and therefore much less available.
Positive correlations were observed between P, Mn, and B as shown in Q3 of the RDA (Figure 4). Just like phosphorous, boron was found to be higher in forests and grasslands, whereas manganese was higher in forests and pastures. Though Mn is one of the essential micronutrients for plants, it can be toxic to plants if available in excess in the soil. We observed an inverse relationship between Mn concentration and soil pH in the RDA (Figure 4). The toxicity levels of Mn vary from one soil type to another, and a threshold level for Mn is not available for most African soils. Boron is known to percolate and leach into the lower profile of the soil; therefore, one would have expected a higher concentration in the subsoil (Zia, Ahmad, Khaliq, Ahmad, & Irshad, 2006). In contrast, B concentration was instead higher in the topsoil. This is consistent with previous studies that attributed the higher B concentration in the upper soil layer to the higher organic matter content (Miwa & Fujiwara, 2010). In addition, clay-rich soils limit percolation and reduce B leaching into the subsoil. On the other hand, B was not influenced by topographic position (Table 2), suggesting that it is relatively stable in the soil and does not accumulate in the bottomlands and footslopes due to erosion.
3.3.3 Soil reaction and clay content
Soil pH and clay content are probably among the most informative measurements that can be made to determine soil characteristics. Though not grouped together in Figure 4, both are less affected by anthropogenic activities compared with other soil properties. The availability of essential nutrients is greatly affected by pH. Pastures had significantly lower pH than the other land use types (Table 2), probably due to the removal of basic cations caused by animal grazing (Ahmed, Biondini, & Grygiel, 1994). The pH and SOC were observed to be associated in Q4 of the RDA (Figure 4) and are linked to organic matter content. Higher acidity results in lower biomass production and, consequently, lower inputs of organic matter into the soil. Soil acidity, by influencing the availability of nutrients to plants, also regulates indirectly the production of biomass and soil biota activities (Takoutsing et al., 2016). In contrast, no significant differences in pH were observed with slope gradients (Table 2), attesting the general acidity of the soils of the study area. Clay content was significantly affected by soil depths (higher in subsoil), land use types (highest in cropland), and topographic positions (highest in ridge/crest; Table 2 and Figure 4). Clay content affects soil texture, and one of the main problems with clay soils is their slow permeability. As discussed above, the agricultural soils are associated with higher clay content, and soil texture, though it is of great importance to plant growth, cannot be modified through management practices or cropping systems.
3.5.Implication for ecological intensification in complex topographic landscapes
A worldwide concern is how to meet the increased demand for ecosystem services caused by the rapidly increasing human population. Because there is little if any scope for agricultural land expansion while also preserving natural forests, meeting the demand can only be achieved through the optimal utilization of available ecosystem services during the agricultural production processes, referred to as ecological intensification (Bommarco, Kleijn, & Potts, 2012). Ecological intensification of agricultural practices enhances productivity through sustainable management practices including intercropping, crop rotations, system diversification, and low chemical inputs (Kovács-Hostyánszki et al., 2017). Quantifying the effects of natural processes on the implementation of ecological intensification presents challenges that are more pronounced in areas with complex topography such as the highlands of Cameroon, where agricultural practices and land management interventions are influenced by climate, soil types, and topography. A holistic intensification cannot be achieved without considering the potential of a given soil to fulfill its functions.
The results of our study revealed that some of the soil properties responded significantly to the types of field management practices, location of the plot along the toposequence and vegetation cover, whereas others such as clay content and pH were less affected by these landscape attributes. Previous studies have reported similar and mixed results and attributed the results to the physical features of the landscapes (Gao, He, Yu, Chen, & Wang, 2014; Sun et al., 2015; Takoutsing et al., 2015). Another study in the same site demonstrated significant spatial variation and well-defined patterns of higher values for key soil functional properties in lowlands and valleys, as well as in zones with high and permanent vegetation cover (Takoutsing et al., 2017). In order to understand this variation and address the challenges faced by ecological intensification, further studies are needed at various scales to develop and test site-specific options for sustainable intensification (Takoutsing et al., 2017; Winowiecki et al., 2016). In addition, there is also a need for systematic assessment of the spatial variability of soil properties within landscapes to generate information relevant for agricultural policy and evidence-informed land management at both plot and landscape levels (Shepherd, Shepherd, & Walsh, 2015; Winowiecki et al., 2016). Soil scientists need to take into account local knowledge in formulating hypotheses on soil functioning that determine soil health, and then set up an approach that is integrative, systems-oriented, and multidisciplinary to ensure sustainability of soil health. In this context, it is fundamental for all stakeholders promoting agricultural intensification to apply low-cost and environmental friendly methods that mitigate nutrient loss through leaching and runoff, soil and water contamination, water erosion, and minimize the production of greenhouse gases. Researchers should investigate how to enhance terrestrial ecosystem services through ecological processes. In addition, efforts are also required to set up effective capacity building programs at various levels to ensure accumulation of necessary baseline data and information on land health necessary to design appropriate interventions as soil health is the first requirement for agricultural development and environmental sustainability (Allen, Singh, & Dalal, 2011). Because the definition of soil health cannot be generalized due to site specific contexts, key soil functional properties to assess the quality for a given soil should be selected based on both anthropogenic and environmental factors. The selection based on one-time field measurement does not suffice, and multiyear sampling is required. If possible, biological properties related to the microbial and faunal activities in soils should be incorporated in the set of properties. Another important challenge for researchers is how to make the research findings relevant to small-scale farmers. Emphasis has been given to influencing policy and decision makers with the hope that there would be positive impacts on smallholder farming systems (Ramisch, 2005; Scoones & Toulmin, 1998). None of the previous approaches has targeted farmers directly and is readily applicable in the farmers' context. This is a limitation for most of the results generated by this and other studies.
4 CONCLUSION
Landscape attributes significantly affected values for most of the selected soil properties in the highlands of Cameroon. The use of RDA allowed us to identify the determinant landscape attributes that affect soil properties. Significant interactions existed between land use types and topographic positions for SOC, EC, ExBas, and pH, and between slope gradients and topographic positions for soil pH, whereas Mn was affected by the interaction between land use types and slope gradients. These interactions indicate that the effects of land use types on these indicators were not the same for all slope gradients and topographic positions. The results show either an overall decrease in soil properties as a result of anthropogenic activities or a decrease in the upslope direction due to natural factors such as runoff. In general, the effects of landscape attributes on soil properties provided mixed results: Some responded more to the types of field management practices, plot location along the toposequence and vegetation cover, whereas some were less affected by anthropogenic activities leading to the conclusion that soil properties are affected by a combination of both management practices and environmental factors. These results are useful for the implementation of ecological intensification in areas with complex topography such as the highlands of Cameroon and other tropical areas with similar environmental conditions, offering a valuable reference to guide the development of future intensification policies.
ACKNOWLEDGMENTS
The authors are thankful to the entire staff of ICRAF Soil-Plant Spectral Diagnostic Laboratory in Nairobi for their assistance during the analyses of soil samples. We are also grateful to all donors who supported this research through their contributions to the CGIAR Research Program on Water, Land and Ecosystems (WLE) http://www.cgiar.org/who-we-are/cgiar-fund/fund-donors-2. This study was undertaken as part of the activities of the land health and management project implemented in the Western Highlands of Cameroon.