Root foraging strategies and soil patchiness in a humid savanna

In Lamto (Côte d'Ivoire), the savanna is a patchy environment as far as soil is concerned: tree clumps and termite mounds lead to higher nutrient contents than in the surrounding savanna. Mature Borassus aethiopum (Mart.) specimens are tall palm trees dominating the community, with aerial parts located out of these nutrient-rich patches.Palm root densities were compared under tree clumps and in the surrounding savanna, and were also sampled along transects between palm trees and nutrient-rich patches (two clumps and one mound). Palm root densities were far higher (up to 10 times) in the nitrogen-rich soil of both clumps and termite mounds than in the surrounding savanna. Evidence is given that palm trees are able to extend their root system as far as 20 m towards these nutrient-rich patches where they proliferate. These results point out a particular root foraging strategy, which is one of the first known for a woody perennial. They also provide new insights for understanding nitrogen cycling and savannas high rate of primary production.     

Temporal and spatial partitioning of water resources among eight woody species in a Hawaiian dry forest

Lowland dry forests are unique in Hawaii for their high diversity of tree species compared with wet forests. We characterized spatial and temporal partitioning of soil water resources among seven indigenous and one invasive dry forest species to determine whether the degree of partitioning was consistent with the relatively high species richness in these forests. Patterns of water utilization were inferred from stable hydrogen isotope ratios (δD) of soil and xylem water, zones of soil water depletion, plant water status, leaf phenology, and spatial patterns of species distribution. Soil water δD values ranged from –20‰ near the surface to –48‰ at 130 cm depth. Metrosideros polymorpha, an evergreen species, and Reynoldsia sandwicensis, a drought-deciduous species, had xylem sap δD values of about –52‰, and appeared to obtain their water largely from deeper soil layers. The remaining six species had xylem δD values ranging from –33 to –42‰, and apparently obtained water from shallower soil layers. Xylem water δD values were negatively correlated with minimum annual leaf water potential and positively correlated with leaf solute content, an integrated measure of leaf water deficit. Seasonal patterns of leaf production ranged from dry season deciduous at one extreme to evergreen with near constant leaf expansion rates at the other. Species tapping water more actively from deeper soil layers tended to exhibit larger seasonality of leaf production than species relying on shallower soil water sources. Individuals of Myoporum sandwicense were more spatially isolated than would be expected by chance. Even though this species apparently extracted water primarily from shallow soil layers, as indicated by its xylem δD values, its nearly constant growth rates across all seasons may have been the result of a larger volume of soil water available per individual. The two dominant species, Diospyros sandwicensis and Nestegis sandwicensis, exhibited low leaf water potentials during the dry season and apparently drew water mostly from the upper portion of the soil profile, which may have allowed them to exploit light precipitation events more effectively than the more deeply rooted species. Character displacement in spatial and temporal patterns of soil water uptake was consistent with the relatively high diversity of woody species in Hawaiian dry forests. Received: 20 May 1999 / Accepted: 2 March 2000     

Water resource partitioning,stem xylem hydraulic properties,and plant water use strategies in a seasonally dry riparian tropical rainforest

This study investigated seasonal variation in the origin of water used by plants in a riparian tropical rainforest community and explored linkages between plant water source, plant xylem hydraulic conductivity and response to the onset of dry conditions. The study focused on five co-dominant canopy species, comprising three tree species (Doryphora aromatica, Argyrodendron trifoliolatum, Castanospora alphandii) and two climbing palms (Calamus australis and Calamus caryotoides). Stable isotope ratios of oxygen in water (¹⁸O) from soil, groundwater, stream water and plant xylem measured in the wet season and the subsequent dry season revealed water resource partitioning between species in the dry season. Measurement of stem-area-specific hydraulic conductivity (K_S) in the wet season and subsequent dry season showed a significant dry-season loss of K_S in three of the five species (Castanospora alphandii, Calamus australis and C. caryotoides) and a decrease in mean K_S for all species. This loss of hydraulic conductivity was positively correlated with the difference between wet-season and dry-season midday leaf water potentials and with leaf carbon isotope discrimination, indicating that plants that were less susceptible to loss of conductivity had greater control over transpiration rate and were more water-use efficient.     

Complexity in water and carbon dioxide fluxes following rain pulses in an African savanna

The idea that many processes in arid and semi-arid ecosystems are dormant until activated by a pulse of rainfall, and then decay from a maximum rate as the soil dries, is widely used as a conceptual and mathematical model, but has rarely been evaluated with data. This paper examines soil water, evapotranspiration (ET), and net ecosystem CO₂ exchange measured for 5 years at an eddy covariance tower sited in an Acacia–Combretum savanna near Skukuza in the Kruger National Park, South Africa. The analysis characterizes ecosystem flux responses to discrete rain events and evaluates the skill of increasingly complex “pulse models”. Rainfall pulses exert strong control over ecosystem-scale water and CO₂ fluxes at this site, but the simplest pulse models do a poor job of characterizing the dynamics of the response. Successful models need to include the time lag between the wetting event and the process peak, which differ for evaporation, photosynthesis and respiration. Adding further complexity, the time lag depends on the prior duration and degree of water stress. ET response is well characterized by a linear function of potential ET and a logistic function of profile-total soil water content, with remaining seasonal variation correlating with vegetation phenological dynamics (leaf area). A 1- to 3-day lag to maximal ET following wetting is a source of hysteresis in the ET response to soil water. Respiration responds to wetting within days, while photosynthesis takes a week or longer to reach its peak if the rainfall was preceded by a long dry spell. Both processes exhibit nonlinear functional responses that vary seasonally. We conclude that a more mechanistic approach than simple pulse modeling is needed to represent daily ecosystem C processes in semiarid savannas. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Seasonal selection of soil types and grass swards by roan antelope in a South African savanna

Roan antelope are distributed mainly in regions characterized by infertile soils, offering food of low quality. We hypothesized that roan may select localities with higher soil nutrient levels and/or grass swards with more favourable properties in terms of food abundance or quality than generally available in those regions. Roan antelope were observed in a savanna region in South Africa where soils of widely varying nutrient status occurred. Roan favoured open grassland over wooded savanna areas. During the wet season, roan preferred sites with felsite‐derived soil of intermediate soil nutrient status. Grasslands growing on nutrient enriched alluvial soils were preferred outside of the early wet season, although most of the favourable sward characteristics were present in other landscape units. Food quantity, rather than quality appeared to attract roan to foraging sites in the late wet and early dry seasons. Food quality appeared more important in the early wet and late dry seasons. The higher degree of clustering of leafy material within foraging swards seemed to be an additional discriminating factor. The factors governing the selection of foraging sites by roan did not seem notably different from those influencing other species of grazing ruminant, but roan nevertheless seemed tolerant of stemmy grasslands growing on nutrient richer substrates.     

Spatial variation in vegetation structure coupled to plant available water determined by two-dimensional soil resistivity profiling in a Brazilian savanna

Tropical savannas commonly exhibit large spatial heterogeneity in vegetation structure. Fine-scale patterns of soil moisture, particularly in the deeper soil layers, have not been well investigated as factors possibly influencing vegetation patterns in savannas. Here we investigate the role of soil water availability and heterogeneity related to vegetation structure in an area of the Brazilian savanna (Cerrado). Our objective was to determine whether horizontal spatial variations of soil water are coupled with patterns of vegetation structure across tens of meters. We applied a novel methodological approach to convert soil electrical resistivity measurements along three 275-m transects to volumetric water content and then to estimates of plant available water (PAW). Structural attributes of the woody vegetation, including plant position, height, basal circumference, crown dimensions, and leaf area index, were surveyed within twenty-two 100-m² plots along the same transects, where no obvious vegetation gradients had been apparent. Spatial heterogeneity was evaluated through measurements of spatial autocorrelation in both PAW and vegetation structure. Comparisons with null models suggest that plants were randomly distributed over the transect with the greatest mean PAW and lowest PAW heterogeneity, and clustered in the driest and most heterogeneous transect. Plant density was positively related with PAW in the top 4 m of soil. The density-dependent vegetation attributes that are related to plot biomass, such as sum of tree heights per plot, exhibited spatial variation patterns that were remarkably similar to spatial variation of PAW in the top 4 m of soil. For PAW below 4 m depth, mean vegetation attributes, such as mean height, were negatively correlated with PAW, suggesting greater water uptake from the deep soil by plants of larger stature. These results are consistent with PAW heterogeneity being an important structuring factor in the plant distribution at the scale of tens of meters in this ecosystem.     

The water relations of two evergreen tree species in a karst savanna

The ecohydrology of karst has not received much attention, despite the disproportionally large contribution of karst aquifers to freshwater supplies. Karst savannas, like many savannas elsewhere, are encroached by woody plants, with possibly negative consequences on aquifer recharge. However, the role of savanna tree species in hydrological processes remains unclear, not least because the location and water absorption zones of tree roots in the spatially complex subsurface strata are unknown. This study examined the water sources and water relations of two savanna trees, Quercus fusiformis (Small) and Juniperus ashei (Buchholz) in the karst region of the eastern Edwards Plateau, Texas (USA). Stable isotope analysis of stem water revealed that both species took up evaporatively enriched water during the warm season, suggesting a relatively shallow water source in the epikarst, the transition zone between soil and bedrock. Q. fusiformis had consistently higher predawn water potentials than J. ashei during drought, and thus was probably deeper-rooted and less capable of maintaining gas exchange at low water potentials. Although the water potential of both species recovered after drought-breaking spring and summer rain events, associated shifts in stem water isotope ratios did not indicate significant uptake of rainwater from the shallow soil. A hypothesis is developed to explain this phenomenon invoking a piston-flow mechanism that pushes water stored in macropores into the active root zones of the trees. Epikarst structure varied greatly with parent material and topography, and had strong effects on seasonal fluctuations in plant water status. The study suggests that tree species of the Edwards Plateau do not commonly reduce aquifer recharge by tapping directly into perched water tables, but more likely by reducing water storage in the epikarst. A more general conclusion is that models of savanna water relations based on Walter's two-layer model may not apply unequivocally to karst savannas.     

Density, survival and dispersal of Anopheles gambiae complex mosquitoes in a West African Sudan savanna village

Abstract. To obtain information on adult populations of Afrotropical malaria vector mosquitoes, mark-release-recapture experiments were performed with Anopheles females collected from indoor resting-sites in a savanna area near Ouagadougou, Burkina Faso, during September 1991 and 1992. Results were used to estimate the absolute population densities, daily survival rates, and dispersal parameters of malaria vectors in that area.
In 1991 a total of 7260 female Anopheles were marked and released, of which 106 were recaptured in the release village and 6 in the neighbouring villages, a total recapture rate of 1.5%. The following year 13, 854 female Anopheles were released and 116 recaptured in Goundri and 8 in the neighbouring villages, a total recapture rate of 0.9%. Recaptures were found in three of eight villages near Goundri. Nearly all of the recaptured mosquitoes were An.gambiae s.l. Of these, molecular determination revealed that An.gambiae s.s. and An.arabiensis were present in a ratio of -2:3.
Two simple random models of dispersal were simulated and the parameters of the models determined by searching for the least-squared fit between simulated and observed distributions. The mean distance moved by individual mosquitoes, estimated in this way, ranged 350–650 m day^-1, depending on die model and the year considered. Population densities were estimated using the Lincoln Index, Fisher-Ford and Jolly's methods. The estimates of population size had high standard errors and were not particularly consistent. A 'consensus' value of 150,000–350,000 mosquitoes is believed to apply for ht An.gambiae s.l. female population. Survival was estimated to be 80–88% per day.     

Carbon fluxes to the soil in a mature temperate forest assessed by <Superscript>13</Superscript>C isotope tracing

Photosynthetic carbon uptake and respiratory C release from soil are major components of the global carbon balance. The use of ¹³C depleted CO₂ (¹³C = –30) in a free air CO₂ enrichment experiment in a mature deciduous forest permitted us to trace the carbon transfer from tree crowns to the rhizosphere of 100–120 years old trees. During the first season of CO₂ enrichment the CO₂ released from soil originated substantially from concurrent assimilation. The small contribution of recent carbon in fine roots suggests a much slower fine root turnover than is often assumed.¹³C abundance in soil air correlated best with temperature data taken from 4 to 10 days before air sampling time and is thus rapidly available for root and rhizosphere respiration. The spatial variability of ¹³C in soil air showed relationships to above ground tree types such as conifers versus broad-leaved trees. Considering the complexity and strong overlap of roots from different individuals in a forest, this finding opens an exciting new possibility of associating respiration with different species. What might be seen as signal noise does in fact contain valuable information on the spatial heterogeneity of tree-soil interaction.     

Leaf gas exchange characteristics and water- and nitrogen-use efficiencies of dominant grass and tree species in a West African savanna

Whereas leaf gas exchange properties are important to assess carbon and water fluxes in ecosystems worldwide, information of this type is scarce for savanna species. In this study, gas exchange characteristics of 2 C₄ grass species (Andropogon canaliculatus and Hyparrhenia diplandra) and 2 C₃ tree species (Crossopteryx febrifuga and Cussonia arborea) from the West-African savanna of Lamto (Ivory Coast) were investigated in the field. Measurements were done in order to provide data to allow the parameterization of biochemically-based models of photosynthesis (for C₄ and C₃ plant metabolic types) and stomatal conductance ; and to compare gas exchange characteristics of coexisting species. No systematic difference was found between grass and tree species for reference stomatal conductance, under standard environmental conditions, or stomatal response to incident light or vapour pressure deficit at leaf surface. Conversely, grass species displayed higher water (1.5-2 fold) and nitrogen (2-5 fold) photosynthetic use efficiencies (WUE and NUE, ratio of net photosynthesis to transpiration and leaf nitrogen, respectively). These contrasts were attributed to the CO₂ concentrating mechanism of C₄ plants. When looking within plant life forms, no important difference was found between grass species. However, significant contrasts were found between tree species, Cussonia showing higher NUE and reference stomatal conductance than Crossopteryx. These results stress the need to account for functional diversity when estimating ecosystem carbon and water fluxes. In particular, our results suggest that the tree/grass ratio, and also the composition of the tree layer, could strongly affect WUE and NUE at the ecosystem scale in West African savannas.     

The influence of soil texture on the soil water dynamics and vegetation structure of a shortgrass steppe ecosystem

We analyzed soil water data from three sites with different soil textures in the shortgrass steppe of northeastern Colorado, USA. Our objective was to evaluate the relationship between the occurrence of plant functional types and the effect of soil texture on soil water availability. Soil water availability was greatest in the upper soil layers at all three sites, but the loamy sand site had significantly greater soil water availability than the sandy clay loam and sandy clay sites in wetter years at depths below 60 cm. Calculations of proportional water availability by layer using both field data and fifty-year soil water model simulations, showed that the sandy clay loam and sandy clay soils on average had greater water availability in layers 30 cm and above, but that the loamy sand had the greatest water availability in layers beneath this, particularly at 105 cm. This observation can be linked to the occurrence of a fine textured subsoil at this site. The textural pattern in the loamy sand profile effectively creates two water resources: a shallow pool accessible to all plants; and a deep pool accessible only to deep-rooted plants. This is offered as an explanation for the co-dominance of the two main plant functional types at the loamy sand site. At the other two sites, shallow-rooted shortgrass vegetation dominated, being more consistent with the general pattern for the area. Thus the patterns of vegetation structure at the three sites were consistent with the hypothesis. Aboveground net primary production data for the three sites, along with transpiration estimates from the model simulations, indicated that the additional water availability in the coarse textured soil was associated with higher overall plant productivity.Nomenclature: Taxonomic nomenclature follows R. L. McGregor & T. M. Barkley (1986) Flora of the Great Plains. Great Plains Flora Association. University Press of Kansas, Lawrence.     

A hydrological tracer study of water uptake depth in a Scots pine forest under two different water regimes

Little is known about the vertical distribution of water uptake by trees under different water supply regimes, the subject of this study, conducted in a Scots pine stand on sandy loam in northern Sweden. The objective was to determine the water uptake distribution in pines under two different water regimes, desiccation (no precipitation) and irrigation (2?mm day^?1 in July and 1?mm day^?1 in August), and to relate the uptake to water content, root and soil texture distributions. The natural ¹⁸O gradient in soil water was exploited, in combination with two added tracers, ²H at 10?cm and ³H at 20?cm depth. Extraction of xylem sap and water from the soil profile then enabled evaluation of relative water uptake from four different soil depths (humus layer, 0–10, 10–25 and 25–55?cm) in each of two 50-m² plots per treatment. In addition, water content, root biomass and soil texture were determined. There were differences in vertical water uptake distribution between treatments. In July, the pines at the irrigated and desiccated plots took up 50% and 30%, respectively, of their water from the upper layers, down to 25?cm depth. In August, the pines on the irrigated plots took up a greater proportion of their water from layers below 25?cm deep than they did in July. In a linear regression, the mean hydraulic conductivity for each mineral soil horizon explained a large part of the variation in relative water uptake. No systematic variation in the residual water uptake correlated to the root distribution. It was therefore concluded that the distribution of water uptake by the pines at Åheden was not a function of root density in the mineral soil, but was largely determined by the unsaturated hydraulic conductivity.     

Spatial scaling of ecosystem C and N in a subtropical savanna landscape

Widely occurred woody encroachment in grass‐dominated ecosystems has the potential to influence soil organic carbon (SOC) and total nitrogen (TN) pools at local, regional, and global scales. Evaluation of this potential requires assessment of both pool sizes and their spatial patterns. We quantified SOC and TN, their relationships with soil and vegetation attributes, and their spatial scaling along a catena (hill‐slope) gradient in the southern Great Plains, USA where woody cover has increased substantially over the past 100 years. Quadrat variance analysis revealed spatial variation in SOC and TN at two scales. The larger scale variation (40–45 m) was approximately the distance between centers of woody plant communities and their adjoining herbaceous patches. The smaller scale variation (10 m) appeared to reflect the local influence of shrubs on SOC and TN. Litter, root biomass, shrub, and tree basal area (a proxy for plant age) exhibited not only similar spatial scales, but also strong correlations with SOC and TN, suggesting invasive woody plants alter both the storage and spatial scaling of SOC and TN through ecological processes related primarily to root turnover and, to a lesser extent litter production, as mediated by time of occupancy. Forb and grass biomass were not significantly correlated with SOC and TN suggesting that changes in herbaceous vegetation have not been the driving force for the observed changes in SOC and TN. Because SOC and TN varied at two scales, it would be inappropriate to estimate SOC and TN pools at broad scales by extrapolating from point sampling at fine scales. Sampling designs that capture variation at multiple scales are required to estimate SOC and TN pools at broader scales. Knowledge of spatial scaling and correlations will be necessary to design field sampling protocols to quantify the biogeochemical consequences of woody plant encroachment at broad scales.     

Precipitation pulse use by an invasive woody legume: the role of soil texture and pulse size

Plant metabolic activity in arid and semi-arid environments is largely tied to episodic precipitation events or “pulses”. The ability of plants to take up and utilize rain pulses during the growing season in these water-limited ecosystems is determined in part by pulse timing, intensity and amount, and by hydrological properties of the soil that translate precipitation into plant-available soil moisture. We assessed the sensitivity of an invasive woody plant, velvet mesquite (Prosopis velutina Woot.), to large (35 mm) and small (10 mm) isotopically labeled irrigation pulses on two contrasting soil textures (sandy-loam vs. loamy-clay) in semi-desert grassland in southeastern Arizona, USA. Predawn leaf water potential (Ψ_pd), the isotopic abundance of deuterium in stem water (δD), the abundance of ¹³C in soluble leaf sugar (δ¹³C), and percent volumetric soil water content (θ_v) were measured prior to irrigation and repeatedly for 2 weeks following irrigation. Plant water potential and the percent of pulse water present in the stem xylem indicated that although mesquite trees on both coarse- and fine-textured soils quickly responded to the large irrigation pulse, the magnitude and duration of this response substantially differed between soil textures. After reaching a maximum 4 days after the irrigation, the fraction of pulse water in stem xylem decreased more rapidly on the loamy-clay soil than the sandy-loam soil. Similarly, on both soil textures mesquite significantly responded to the 10-mm pulse. However, the magnitude of this response was substantially greater for mesquite on the sandy-loam soil compared to loamy-clay soil. The relationship between Ψ_pd and δ¹³C of leaf-soluble carbohydrates over the pulse period did not differ between plants at the two sites, indicating that differences in photosynthetic response of mesquite trees to the moisture pulses was a function of soil water availability within the rooting zone rather than differences in plant biochemical or physiological constraints. Patterns of resource acquisition by mesquite during the dynamic wetting–drying cycle following rainfall pulses is controlled by a complex interaction between pulse size and soil hydraulic properties. A better understanding of how this interaction affects plant water availability and photosynthetic response is needed to predict how grassland structure and function will respond to climate change.     

Effect of soil type on the damping-off of tomato seedlings caused bySclerotium rolfsii in the Nigerian savanna

Summary The conductiveness of tropical savanna soils in Nigeria to the damping-off of tomato seedlings induced bySclerotium rolfsii was studied using soil from different localities in the savanna zones of the country. The results showed that soils low in percentage clay content were more conducive to the disease than those with high clay contents. Statistical analysis also revealed a high negative correlation (r=−0.91) between disease conduciveness indices and percentage clay contents of the various soils, indicating that as clay content of soil increased, disease decreased.     

Comparison of APRI and Hydrus-2D models to simulate soil water dynamics in a vineyard under alternate partial root zone drip irrigation

Alternate partial root zone irrigation (APRI) is a new water-saving irrigation technique. It can reduce irrigation water and transpiration without reduction in crop yield, thus increase water and nutrient use efficiency. Understanding of soil moisture distribution and dynamic under the alternate partial root zone drip irrigation (APDI) can help to develop the efficient irrigation schemes. In this paper, a two-dimensional (2D) root water uptake model was proposed based on soil water dynamic and root distribution of grape vine, and a function of soil evaporation related to soil water content was defined under the APDI. Then the soil water dynamic model of APDI (APRI-model) was developed based on the 2D root water uptake model and soil evaporation function combined with average measured soil moisture content at 0–10 cm soil layer. Soil water dynamic in APDI was respectively simulated by Hydrus-2D model and APRI-model. The simulated soil water contents by two models were compared with the measured value. The results showed that the values of root-mean-square-error (RMSE) range from 0.01 to 0.022 cm³/cm³ for APRI-model, and from 0.012 to 0.031 cm³/cm³ for Hydrus-2D model. The average relative error between the simulated and measured soil water content is about 10% for APRI-model, and from 11% to 29% for Hydrus-2D model, indicating that two models perform well in simulating soil moisture dynamic under the APDI, but the APRI-model is more suitable for modeling the soil water dynamic in the arid region with greater soil evaporation and uneven root distribution.     

Soil water dynamics and vegetation patterns in a semiarid grassland

Long-term (1985–1992) dynamics and spatial variations in soil water below the evaporative zone were evaluated for a shortgrass steppe with a low and variable precipitation regime. Each of a sandy loam, clay loam, and two sandy clay loam sites comprised a toposequence with upland, midslope and lowland positions. Soil water was monitored at 15 cm intervals providing estimates covering 22.5 to 97.5 cm depths.Soil water throughout the profile was highest in the clay loam site and lowest in the sandy loam site. However, stored soil water did not vary systematically among slope positions. Total vegetation cover was highest on the lowland in two sites, but was greatest on the midslope position in the other two. Total vegetation cover was greatest on the CL site, which was the wettest in terms of soil water. Soil water depletion was related to the depth-distribution of roots. There was an inverse relationship between aboveground production and soil water content of the 30, 45 and 60 cm layers during the growth period. Root distributions through the profile did not, however, vary with soil texture or with different soil water profiles controlled by texture. The less variable water content of deeper soil layers is a resource which potentially buffers the impact of pronounced variability in precipitation and thus contributes to vegetation stability of the shortgrass community.     

Seasonal abundance of soil-surface arthropods in relation to some meteorological and edaphic variables of the grassland and tree-planted areas in a tropical semi-arid savanna

Seasonality of relative population abundance in different groups of soil-surface arthropods was investigated monthly by pit-fall traps during a 2-year period in the grassland and tree-planted areas of a tropical semi-arid savanna at Warangal (south India). Densities of most groups were lowest during summer and highest during the rainy season. They were less abundant during winter. Arthropods were recorded in higher numbers in tree-planted compared to grassland areas. Certain arthropods that were found only during part of the year were recorded for a longer period in the tree-planted area. Formicidae,Monomorium indicum Forel,Crematogaster sp. andPachycondyla? tesserinoda (Emery), and Coleoptera,Pachycera sp. reached maximum densities in the rainy season and minimum numbers during winter and summer in the grassland area. However, these species had lower densities during the rainy season and reached maximum densities during winter and summer in the tree-planted area. The seasonal abundance of arthropods showed significant linear correlations with different abiotic environmental variables such as rainfall, soil moisture, organic matter, soil and air temperatures, soil pH, relative humidity at the soil surface, and potassium and phosphorus of surface soil. Soil moisture and rainfall were generally the strongest correlates with densities, particularly in the grassland area.