Sandblasting promotes shrub encroachment in arid grasslands.

Shrub encroachment is a common ecological state transition in global drylands and has myriad adverse effects on grasslands and the services they provide. This physiognomic shift is often ascribed to changes in climate (e.g. precipitation) and disturbance regimes (e.g. grazing and fire), but this remains debated. Aeolian processes are known to impact resource distribution in drylands, but their potential role in grassland-to-shrubland state changes has received little attention. We quantified the effects of 'sandblasting' (abrasive damage by wind-blown soil) on the ecophysiology of dryland grass vs shrub functional types using a portable wind tunnel to test the hypothesis that grasses would be more susceptible to sandblasting than shrubs and, thus, reinforce transitions to shrub dominance in wind-erodible grasslands when climate- or disturbance-induced reductions in ground cover occur. Grasses and shrubs responded differently to sandblasting, wherein water-use efficiency declined substantially in grasses, but only slightly in shrubs, owing to grasses having greater increases in day/nighttime leaf conductance and transpiration. The differential ecophysiological response to sandblasting exhibited by grass and shrub functional types could consequently alter the vegetation dynamics in dryland grasslands in favour of the xerophytic shrubs. Sandblasting could thus be an overlooked driver of shrub encroachment in wind-erodible grasslands.

Woody-plant encroachment: Precipitation, herbivory, and grass-competition interact to affect shrub recruitment.

Woody-plant encroachment is a global phenomenon that has been affecting the southwestern United States since the late 1800s. Drought, overgrazing, herbivory, and competition between grasses and shrub seedlings have been hypothesized as the main drivers of shrub establishment. However, there is limited knowledge about the interactions among these drivers. Using a rainfall manipulation system and various herbivore exclosures, we tested hypotheses about how precipitation (PPT), competition between grasses and shrub seedlings, and predation affect the germination and first-year survival of mesquite (Prosopis glandulosa), a shrub that has encroached in Southern Great Plains and Chihuahuan Desert grasslands. We found that mesquite germination and survival (1) increased with increasing PPT, then saturated at about the mean growing season PPT level, (2) that competition between grasses and shrub seedlings had no effect on either germination or survival, and (3) that herbivory by small mammals decreased seedling establishment and survival, while ant granivory showed no effect. In addition to its direct positive effect on survival, PPT had an indirect negative effect via increasing small mammal activity. Current models predict a decrease in PPT in the southwestern United States with increased frequency of extreme events. The non-linear nature of PPT effects on Mesquite recruitment suggests asymmetric responses, wherein drought has a relatively greater negative effect than the positive effect of wet years. Indirect effects of PPT, through its effects on small mammal abundance, highlight the importance of accounting for interactions between biotic and abiotic drivers of shrub encroachment. This study provides quantitative basis for developing tools that can inform effective shrub management strategies in grasslands and savannas.

Soil organic carbon in drylands: shrub encroachment and vegetation management effects dwarf those of livestock grazing.

Dryland ecosystems occur worldwide and play a prominent, but potentially shifting, role in global biogeochemical cycling. Widespread woody plant proliferation, often associated with declines in palatable grasses, has jeopardized livestock production in drylands and prompted attempts to reduce woody cover by chemical or mechanical means. Woody encroachment also has the potential to significantly alter terrestrial carbon storage. However, little is known of the long-term biogeochemical consequences of woody encroachment in the broader context of its interaction with common dryland land uses, including "brush management" (woody plant clearing) and livestock grazing. Present assessments exhibit considerable variation in the consequences of these land use/land cover changes, with evidence that brush management may counteract sizeable impacts of shrub encroachment on soil biogeochemical pools. A challenge to assessing the net effects of brush management in shrub-encroached grasslands on soil organic carbon (SOC) and total nitrogen (N) pools is that land management practices are typically considered in isolation, when they are co-occurring phenomena. Furthermore, few studies have assessed spatial patterns in brush management and how these are affected in decades following treatment on sites with contrasting grazing histories. To address these uncertainties and interactions, we quantified the impacts of shrub encroachment and their subsequent mortality resulting from brush management (herbicide application) on SOC and N pools in a Sonoran Desert grassland where long-term grazing manipulations (>100 yr) co-occur with shrub encroachment and brush management. Pools of SOC and N associated with herbicided shrubs declined markedly over ~40 yr, offsetting 66% of the increases from shrub encroachment. However, spatial patterns in SOC induced by shrubs persisted over the decades following brush management. Century-long protection from grazing did little to change SOC and N pools. Accordingly, shrub encroachment and shrub mortality from brush management each far outweighed livestock grazing impacts. Consideration of the patterns of SOC and N through space (e.g., bole-to-dripline gradients), time (e.g., shrub age/size), land use (e.g., livestock grazing and brush management), and their interactions will position us to improve predictions of SOC and N responses to land use/land cover change, inform C-based management decisions, and objectively evaluate trade-offs with other ecosystem services.

Spatial patterns of grassland­shrubland state transitions: a 74­year record on grazed and protected areas.

Tree and shrub abundance has increased in many grasslands causing changes in ecosystem carbon and nitrogen pools that are related to patterns of woody plant distribution. However, with regard to spatial patterns of shrub proliferation, little is known about how they are influenced by grazing or the extent to which they are influenced by intraspecific interactions. We addressed these questions by quantifying changes in the spatial distribution of Prosopis velutina (mesquite) shrubs over 74 years on grazed and protected grasslands. Livestock are effective agents of mesquite dispersal and mesquite plants have lateral roots extending well beyond the canopy. We therefore hypothesized that mesquite distributions would be random on grazed areas mainly due to cattle dispersion and clustered on protected areas due to decreased dispersal and interspecific interference with grasses; and that clustered or random distributions at early stages of encroachment would give way to regular distributions as stands matured and density-dependent interactions intensified. Assessments in 1932, 1948, and 2006 supported the first hypothesis, but we found no support for the second. In fact, clustering intensified with time on the protected area and the pattern remained random on the grazed site. Although shrub density increased on both areas between 1932 and 2006, we saw no progression toward a regular distribution indicative of density-dependent interactions. We propose that processes related to seed dispersal, grass­shrub seedling interactions, and hydrological constraints on shrub size interact to determine vegetation structure in grassland-to-shrubland state changes with implications for ecosystem function and management.

Long-term functional plasticity in plant hydraulic architecture in response to supplemental moisture.

BACKGROUND AND AIMS: Plasticity in structural and functional traits related to water balance may determine plant performance and survival in ecosystems characterized by water limitation or high levels of rainfall variability, particularly in perennial herbaceous species with long generation cycles. This paper addresses whether and the extent to which several such seasonal to long-term traits respond to changes in moisture availability. METHODS: Using a novel approach that integrates ecology, physiology and anatomy, a comparison was made of lifetime functional traits in the root xylem of a long-lived perennial herb (Potentilla diversifolia, Rosaceae) growing in dry habitats with those of nearby individuals growing where soil moisture had been supplemented for 14 years. Traditional parameters such as specific leaf area (SLA) and above-ground growth were also assessed. KEY RESULTS: Individuals from the site receiving supplemental moisture consistently showed significant responses in all considered traits related to water balance: SLA was greater by 24 %; roots developed 19 % less starch storing tissue, an indicator for drought-stress tolerance; and vessel size distributions shifted towards wider elements that collectively conducted water 54 % more efficiently - but only during the years for which moisture was supplemented. In contrast, above-ground growth parameters showed insignificant or inconsistent responses. CONCLUSIONS: The phenotypic changes documented represent consistent, dynamic responses to increased moisture availability that should increase plant competitive ability. The functional plasticity of xylem anatomy quantified in this study constitutes a mechanistic basis for anticipating the differential success of plant species in response to climate variability and change, particularly where water limitation occurs.

Response of dominant grass and shrub species to water manipulation: an ecophysiological basis for shrub invasion in a Chihuahuan Desert grassland.

Increases in woody vegetation and declines in grasses in arid and semi-arid ecosystems have occurred globally since the 1800s, but the mechanisms driving this major land-cover change remain uncertain and controversial. Working in a shrub-encroached grassland in the northern Chihuahuan Desert where grasses and shrubs typically differ in leaf-level nitrogen allocation, photosynthetic pathway, and root distribution, we asked if differences in leaf-level ecophysiology could help explain shrub proliferation. We predicted that the relative performance of grasses and shrubs would vary with soil moisture due to the different morphological and physiological characteristics of the two life-forms. In a 2-year experiment with ambient, reduced, and enhanced precipitation during the monsoon season, respectively, the encroaching C(3) shrub (honey mesquite Prosopis glandulosa) consistently and substantially outperformed the historically dominant C(4) grass (black grama Bouteloua eriopoda) in terms of photosynthetic rates while also maintaining a more favorable leaf water status. These differences persisted across a wide range of soil moisture conditions, across which mesquite photosynthesis was decoupled from leaf water status and moisture in the upper 50 cm of the soil profile. Mesquite's ability to maintain physiologically active leaves for a greater fraction of the growing season than black grama potentially amplifies and extends the importance of physiological differences. These physiological and phenological differences may help account for grass displacement by shrubs in drylands. Furthermore, the greater sensitivity of the grass to low soil moisture suggests that grasslands may be increasingly susceptible to shrub encroachment in the face of the predicted increases in drought intensity and frequency in the desert of the southwestern USA.

Protection from livestock fails to deter shrub proliferation in a desert landscape with a history of heavy grazing.

Desertification is often characterized by the replacement of mesophytic grasses with xerophytic shrubs. Livestock grazing is considered a key driver of shrub encroachment, although most evidence is anecdotal or confounded by other factors. Mapping of velvet mesquite (Prosopis velutina) shrubs in and out of exclosures in 1932, 1948, and 2006 in semiarid grasslands of southeastern Arizona, USA, afforded the opportunity to quantify livestock grazing effects on mesquite proliferation over 74 years in the absence of fire to test the widespread assumption that livestock grazing promotes shrub proliferation. In 1932, shrub cover, density, and aboveground biomass were compared on grazed (12%, 173 plants/ha, 4182 kg/ha) and newly protected areas (8%, 203 plants/ha, 3119 kg/ha). By 1948, cover on both areas increased to 18%; yet, density on the protected area increased 300% (to 620 plants/ha), nearly twice that of the grazed area (325 plants/ha). From 1932 to 1948, differences in recruitment of new plants and growth of existing plants were reflected in biomass, which was higher on the protected area (415 plants/ha, 8788 kg/ha) relative to the grazed area (155 plants/ha, 7085 kg/ha), although mortality was equally low ( 0.06%). In 2006, 42 years after an herbicide application reset mesquite cover to 10% on both areas, aboveground mesquite mass was comparable on both areas ( 4700 kg/ha), but cover and density on the protected area (22%, 960 plants/ha) exceeded that on the grazed area (15%, 433 plants/ha). Mesquite mass in 2006 was substantially below 1948 levels, so continued accrual is likely. That shrub recovery from herbicides on a biomass basis was much less than recovery on a cover basis suggests that remotely sensed biomass estimates should integrate land management history. Contrary to widely held assumptions, protection from livestock since 1932 not only failed to deter woody-plant proliferation, but actually promoted it relative to grazed areas. Results suggest (1) that thresholds for grassland resistance to shrub encroachment had been crossed by the 1930s, and (2) fire management rather than grazing management may be key to maintaining grassland physiognomy in this bioclimatic region.

Spatial variation of the stable nitrogen isotope ratio of woody plants along a topoedaphic gradient in a subtropical savanna.

Variation in the stable N isotope ratio (delta15N) of plants and soils often reflects the influence of environment on the N cycle. We measured leaf delta15N and N concentration ([N]) on all individuals of Prosopis glandulosa (deciduous tree legume), Condalia hookeri (evergreen shrub), and Zanthoxylum fagara (evergreen shrub) present within a belt transect 308 m long x 12 m wide in a subtropical savanna ecosystem in southern Texas, USA in April and August 2005. Soil texture, gravimetric water content (GWC), total N and delta15N were also measured along the transect. At the landscape scale, leaf delta15N was negatively related to elevation for all the three species along this topoedaphic sequence. Changes in soil delta15N, total N, and GWC appeared to contribute to this spatial pattern of leaf delta15N. In lower portions of the landscape, greater soil N availability and GWC are associated with relatively high rates of both N mineralization and nitrification. Both soil delta15N and leaf [N] were positively correlated with leaf delta15N of non-N2 fixing plants. Leaf delta15N of P. glandulosa, an N2-fixing legume, did not correlate with leaf [N]; the delta15N of P. glandulosa's leaves were closer to atmospheric N2 and significantly lower than those of C. hookeri and Z. fagara. Additionally, at smaller spatial scales, a proximity index (which reflected the density and distance of surrounding P. glandulosa trees) was negatively correlated with leaf delta15N of C. hookeri and Z. fagara, indicating the N2-fixing P. glandulosa may be important to the N nutrition of nearby non-N2-fixing species. Our results indicate plant 15N natural abundance can reflect the extent of N retention and help us better understand N dynamics and plant-soil interactions at ecosystem and landscape scales.

Woody plants in grasslands: post-encroachment stand dynamics.

Woody plant abundance is widely recognized to have increased in savannas and grasslands worldwide. The lack of information on the rates, dynamics, and extent of increases in shrub abundance is a major source of uncertainty in assessing how this vegetation change has influenced biogeochemical cycles. Projecting future consequences of woody cover change on ecosystem function will require knowledge of where shrub cover in present-day stands lies relative to the realizable maximum for a given soil type within a bioclimatic region. We used time-series aerial photography (1936, 1966, and 1996) and field studies to quantify cover and biomass of velvet mesquite (Prosopis velutina Woot.) following its proliferation in a semidesert grassland of Arizona. Mapping of individual shrubs indicated an encroachment phase characterized by high rates of bare patch colonization. Upon entering a stabilization phase, shrub cover increases associated with recruitment and canopy expansion were largely offset by contractions in canopy area of other shrub patches. Instances of shrub disappearance coincided with a period of below-average rainfall (1936-1966). Overall, shrub cover (mean +/- SE) on sandy uplands with few and widely scattered shrubs in 1902 was dynamically stable over the 1936-1996 period averaging approximately 35% +/- 5%. Shrub cover on clayey uplands in 1936 was 17% +/- 2% but subsequently increased twofold to levels comparable to those on sandy uplands by 1966 (36% +/- 7%). Cover on both soils then decreased slightly between 1966 and 1996 to 28% +/- 3%. Thus, soil properties influenced the rate at which landscapes reached a dynamic equilibrium, but not the apparent endpoint. Although sandy and clayey landscapes appear to have stabilized at comparable levels of cover, shrub biomass was 1.4 times greater on clayey soils. Declines in shrub cover between 1966 and 1996 were accompanied by a shift to smaller patch sizes on both sandy and clayey landscapes. Dynamics observed during the stabilization phase suggest that density-dependent regulation may be in play. If woody cover has transitioned from directional increases to a dynamic equilibrium, biomass projections will require monitoring and modeling patch dynamics and stand structure rather than simply changes in total cover.

Shrub encroachment into grasslands: end of an era?

Shifts in the abundance of grasses and woody plants in drylands have occurred several times during the Holocene. However, our understanding of the rates and dynamics of this state-change in recent decades is limited to scattered studies conducted at disparate spatial and temporal scales; the potential misperceptions of shrub cover change could be remedied using cross spatiotemporal scale analyses that link field observations, repeat ground-level photography and remote sensing perspectives. The study was conducted across a semi-arid landscape in southern Arizona. Local data from long-term transects revealed three distinct chronological phases of shrub cover change: expansion (1961-1991, 0.7% y-1), decline (1992-1997, -2.3% y-1) and stabilization (1998-2012, 22-25% with no net cover change). Twenty-eight years (1984-2011) of broad-scale Landsat Thematic Mapper assessments confirm that shrub cover has been relatively stable in recent decades regardless of grazing regimes and landforms with the exception of the proliferation of succulents at lower elevations (verified by repeat photography acquired in 1987 and 2015) where the physical environment is the harshest, reflecting elevated temperature and winter precipitation deficit. Warmer, drier future climates are predicted to reduce woody plant carrying capacity and promote a shift to xerophytic succulents.

Interrelationships among shrub encroachment, land management, and litter decomposition in a semidesert grassland.

Encroachment of woody plants into grasslands, and subsequent brush management, are among the most prominent changes to occur in arid and semiarid systems over the past century. Despite the resulting widespread changes in landcover, substantial uncertainty about the biogeochemical impacts of woody proliferation and brush management exists. We explored the role of shrub encroachment and brush management on leaf litter decomposition in a semidesert grassland where velvet mesquite (Prosopis velutina) abundance has increased over the past 100 years. This change in physiognomy may affect decomposition directly, through altered litter quality or quantity, and indirectly through altered canopy structure. To assess the direct and indirect impacts of shrubs on decomposition, we quantified changes in mass, nitrogen, and carbon in litterbags deployed under mesquite canopies and in intercanopy zones. Litterbags contained foliage from mesquite and Lehmann lovegrass (Eragrostis lehmanniana), a widespread, nonnative grass in southern Arizona. To explore short- and long-term influences of brush management on the initial stages of decomposition, litterbags were deployed at sites where mesquite canopies were removed three weeks, 45 years, or 70 years prior to study initiation. Mesquite litter decomposed more rapidly than lovegrass, but negative indirect influences of mesquite canopies counteracted positive direct effects. Decomposition was positively correlated with soil infiltration into litterbags, which varied with microsite placement, and was lowest under canopies. Low under-canopy decomposition was ostensibly due to decreased soil movement associated with high under-canopy herbaceous biomass. Decomposition rates where canopies were removed three weeks prior to study initiation were comparable to those beneath intact canopies, suggesting that decomposition was driven by mesquite legacy effects on herbaceous cover-soil movement linkages. Decomposition rates where shrubs were removed 45 and 70 years prior to study initiation were comparable to intercanopy rates, suggesting that legacy effects persist less than 45 years. Accurate decomposition modeling has proved challenging in arid and semiarid systems but is critical to understanding biogeochemical responses to woody encroachment and brush management. Predicting brush-management effects on decomposition will require information on shrub-grass interactions and herbaceous biomass influences on soil movement at decadal timescales. Inclusion of microsite factors controlling soil accumulation on litter would improve the predictive capability of decomposition models.

Postfire stand structure in a semiarid savanna: cross-scale challenges estimating biomass.

Algorithms relating remotely sensed woody cover to biomass are often the basis for large-scale inventories of aboveground carbon stocks. However, these algorithms are commonly applied in a generic fashion without consideration of disturbances that might alter vegetation structure. We compared field and remote sensing estimates of woody biomass on savannas with contrasting disturbance (fire) histories and assessed potential errors in estimating woody biomass from cover without considering fire history. Field surveys quantified multilayer cover (MLC) of woody and succulent plants on sites experiencing wildfire in 1989 or 1994 and on nearby unburned (control) sites. Remote sensing estimates of the woody cover fraction (WCF) on burned and control sites were derived from contemporary (2005) dry-season Landsat Thematic Mapper imagery (during a period when herbaceous cover was senescent) using a probabilistic spectral mixture analysis model. Satellite WCF estimates were compared to field MLC assessments and related to aboveground biomass using allometry. Field-based MLC and remotely sensed WCFs both indicated that woody cover was comparable on control areas and areas burned 11-16 years ago. However, biomass was approximately twofold higher on control sites. Canopy cover was a strong predictor of woody biomass on burned and control areas, but fire history significantly altered the linear cover-biomass relationship on control plots to a curvilinear relationship on burned plots. Results suggest predictions of woody biomass from "generic" two-dimensional (2-D) cover algorithms may underestimate biomass in undisturbed stands and overestimate biomass in stands recovering from disturbance. Improving the accuracy of woody-biomass estimates from field and/or remotely sensed cover may therefore require disturbance-specific models or detection of vegetation height and transforming 2-D vegetation cover to 3-D vegetation volume.

Climatic/edaphic controls on soil carbon/nitrogen response to shrub encroachment in desert grassland.

The proliferation of woody plants in grasslands over the past 100+ years can alter carbon, nitrogen, and water cycles and influence land surface-atmosphere interactions. Although the majority of organic carbon in these ecosystems resides belowground, there is no consensus on how this change in land cover has affected soil organic carbon (SOC) and total nitrogen (TN) pools. The degree to which duration of woody plant occupation, climate, and edaphic conditions have mediated SOC and TN responses to changes in life-form composition are poorly understood. We addressed these issues at a desert grassland site in Arizona, USA, where the leguminous shrub velvet mesquite (Prosopis velutina) has proliferated along an elevation/precipitation/temperature gradient and on contrasting soil morphologic surfaces. On sandy loam complexes of mid-Holocene origin, mean SOC and TN of soils in the grassland matrix increased approximately 68% and approximately 45%, respectively, with increasing elevation. Soil organic carbon pools were comparable and TN pools were approximately 23% higher in Pleistocene-aged clay loam complexes co-occurring with Holocene-aged soils at the upper elevation/climatic zone. Across the site, belowground resources associated with large Prosopis plants were 21-154% (SOC) and 18-127% (TN) higher than those in the grassy matrix. The variance in SOC and TN pools accounted for by Prosopis stem size (a rough surrogate for time of site occupation) was highest at the low- and mid-elevation sites (69-74%) and lowest at the upper elevation site (32-38%). Soil delta15N values ranged from 5.5 per thousand to 6.7 per thousand across the soil/elevation zones but were comparable in herbaceous and shrub-impacted soils and exhibited a weak relationship with Prosopis basal stem diameter (r2 < 0.1) and TN (r2 < 0.08). The SOC delta13C values decreased linearly with increasing Prosopis basal diameter, suggesting that size and isotopic composition of the SOC pool is a function of time of Prosopis site occupation. Isotopic mixture models indicate that encroachment of C3 woody plants has also promoted SOC additions from C4 plant sources, indicative of long-term herbaceous facilitation. Grassy sites in contrasting soil/elevation combinations, initially highly distinctive in their SOC pool size and delta13C, appear to be converging on similar values following approximately 100 years of woody plant proliferation.

Seedling responses to water pulses in shrubs with contrasting histories of grassland encroachment.

Woody plant encroachment into grasslands has occurred worldwide, but it is unclear why some tree and shrub species have been markedly more successful than others. For example, Prosopis velutina has proliferated in many grasslands of the Sonoran Desert in North America over the past century, while other shrub species with similar growth form and life history, such as Acacia greggii, have not. We conducted a glasshouse experiment to assess whether differences in early seedling development could help explain why one species and not the other came to dominate many Sonoran Desert grasslands. We established eight watering treatments mimicking a range of natural precipitation patterns and harvested seedlings 16 or 17 days after germination. A. greggii had nearly 7 times more seed mass than P. velutina, but P. velutina emerged earlier (by 3.0±0.3 d) and grew faster (by 8.7±0.5 mg d⁻¹). Shoot mass at harvest was higher in A. greggii (99±6 mg seedling⁻¹) than in P. velutina (74±2 mg seedling⁻¹), but there was no significant difference in root mass (54±3 and 49±2 mg seedling⁻¹, respectively). Taproot elongation was differentially sensitive to water supply: under the highest initial watering pulse, taproots were 52±19 mm longer in P. velutina than in A. greggii. Enhanced taproot elongation under favorable rainfall conditions could give nascent P. velutina seedlings growth and survivorship advantages by helping reduce competition with grasses and maintain contact with soil water during drought. Conversely, A. greggii's greater investment in mass per seed appeared to provide little return in early seedling growth. We suggest that such differences in recruitment traits and their sensitivities to environmental conditions may help explain ecological differences between species that are highly similar as adults and help identify pivotal drivers of shrub encroachment into grasslands.

Woody plant encroachment into grasslands: spatial patterns of functional group distribution and community development.

Woody plant encroachment into grasslands has been globally widespread. The woody species invading grasslands represent a variety of contrasting plant functional groups and growth forms. Are some woody plant functional types (PFTs) better suited to invade grasslands than others? To what extent do local patterns of distribution and abundance of woody PFTs invading grasslands reflect intrinsic topoedaphic properties versus plant-induced changes in soil properties? We addressed these questions in the Southern Great Plains, United States at a subtropical grassland known to have been encroached upon by woody species over the past 50-100 years. A total of 20 woody species (9 tree-statured; 11 shrub-statured) were encountered along a transect extending from an upland into a playa basin. About half of the encroaching woody plants were potential N2-fixers (55% of species), but they contributed only 7% to 16 % of the total basal area. Most species and the PFTs they represent were ubiquitously distributed along the topoedaphic gradient, but with varying abundances. Overstory-understory comparisons suggest that while future species composition of these woody communities is likely to change, PFT composition is not. Canonical correspondence analysis (CCA) ordination and variance partitioning (Partial CCA) indicated that woody species and PFT composition in developing woody communities was primarily influenced by intrinsic landscape location variables (e.g., soil texture) and secondarily by plant-induced changes in soil organic carbon and total nitrogen content. The ubiquitous distribution of species and PFTs suggests that woody plants are generally well-suited to a broad range of grassland topoedaphic settings. However, here we only examined categorical and non-quantitative functional traits. Although intrinsic soil properties exerted more control over the floristics of grassland-to-woodland succession did plant modifications of soil carbon and nitrogen concentrations, the latter are likely to influence productivity and nutrient cycling and may, over longer time-frames, feed back to influence PFT distributions.

Variation in woody plant delta(13)C along a topoedaphic gradient in a subtropical savanna parkland.

delta(13)C values of C(3) plants are indicators of plant carbon-water relations that integrate plant responses to environmental conditions. However, few studies have quantified spatial variation in plant delta(13)C at the landscape scale. We determined variation in leaf delta(13)C, leaf nitrogen per leaf area (N(area)), and specific leaf area (SLA) in April and August 2005 for all individuals of three common woody species within a 308 x 12-m belt transect spanning an upland-lowland topoedaphic gradient in a subtropical savanna in southern Texas. Clay content, available soil moisture, and soil total N were all negatively correlated with elevation. The delta(13)C values of Prosopis glandulosa (deciduous N(2)-fixing tree legume), Condalia hookeri (evergreen shrub), and Zanthoxylum fagara (evergreen shrub) leaves increased 1-4 per thousand with decreasing elevation, with the delta(13)C value of P. glandulosa leaves being 1-3 per thousand higher than those of the two shrub species. Contrary to theory and results from previous studies, delta(13)C values were highest where soil water was most available, suggesting that some other variable was overriding or interacting with water availability. Leaf N(area) was positively correlated with leaf delta(13)C of all species (p < 0.01) and appeared to exert the strongest control over delta(13)C along this topoedaphic gradient. Since leaf N(area) is positively related to photosynthetic capacity, plants with high leaf N(area) are likely to have low p (I)/p (a) ratios and therefore higher delta(13)C values, assuming stomatal conductance is constant. Specific leaf area was not correlated significantly with leaf delta(13)C. Following a progressive growing season drought in July/August, leaf delta(13)C decreased. The lower delta(13)C in August may reflect the accumulation of (13)C-depleted epicuticular leaf wax. We suggest control of leaf delta(13)C along this topoedaphic gradient is mediated by leaf N(area) rather than by stomatal conductance limitations associated with water availability.