THE CARBON CYCLE IN DRYLANDS Penelope Serrano-Ortiza Oscar Pérez-Priegob, Enrique P. Sánchez-Cañetea, Ana Werea, Francisco Domingoa, Cecilio Oyonartec and Andrew S. Kowalskib. a Estación Experimental de Zonas Àridas (EEZA, CSIC), 04120 Almería, Spain Dept. Física Aplicada, Universidad de Granada, 18071 Granada, Spain c Dept. Edafología & Química Agrícola, Universidad de Almería, 04120 Almería, Spain b 1. Introduction Drylands are characterised by patches of vegetation and bare soil exposed to erratic rainfall events producing water-stressed vegetation during the drought period (Domingo et al. 1999). These water-limited ecosystems exist on every continent and comprise nearly a third of the total land surface corresponding to 60 million km 2 (Okin 2001; Schlesinger 1990). Dryland rangelands support about 50% of the world's livepool and provide forages for both domestic animals and wildlife (Puigdefábregas 1998). Although drylands withstand extreme climatic conditions, they are very sensitive to perturbations such as drought, fires or climate change, leading to desertification (Mouat and Lancaster 2006). Globally, the organic and inorganic carbon (C) storage in such water-limited systems is about 20-30% of the terrestrial global total (Eswaran et al. 2000; Rasmussen 2006; Safriel et al. 2005). In this context, the main characteristic of drylands is the capacity to store about 95% of the soil inorganic C (SIC) globally via “caliche formation” (Marion et al. 2008). However, the role of the SIC pool in relation to climate change is less well understood (Lal and Kimble 2000). In addition, due to arid conditions and the large percentage of bare soil and carbonate rocks, some other processes besides photosynthesis and respiration contribute to C sequestration or C gaseous emissions to the atmosphere. These include weathering processes, formation of secondary carbonates and subsoil ventilation that can even dominate the ecosystem C exchange during the dry season. Such contributions limit the use of biological models to provide estimates of C pool in drylands. This work reviews measurements of annual C sink capacities together with analysis of the main drivers controlling the potential C sequestration capacity and principal processes involved. 2. Materials and Methods This work analyzes the published literature concerning the carbon cycle over drylands using the eddy covariance technique. The eddy covariance technique is one of the most important methods used all over the world and has the potential to quantify how whole ecosystems respond to a spectrum of climate regimes (Baldocchi et al. 2001). The use of this powerful tool requires micrometeorological expertise and technological advances (Aubinet et al. 2000). Fluxes of CO2 and H2O are estimated using an infrared gas analyzer to measure densities of CO2 and H2O and a sonic anemometer to measure wind speed at a frequency higher than 10 Hz. 3. Results and Discussion 3.1 Main natural processes involved in carbon sequestration and loss 3.1.1 Biological processes The organic C stored in soils and biomass is mainly due to the balance between photosynthesis (net CO2 uptake) and respiration (net CO2 release via decomposition, degradation and diffusion processes). Such biological processes are mainly responsible for annual NEEs in most ecosystems (forest, wetlands, cropland, etc.). Thus, the FLUXNET community (Baldocchi et al. 2001) interprets CO2 fluxes measured using micrometeorological techniques (Dabberdt et al. 1993) as a biological flux neglecting non-biological processes (Falge et al. 2001; Reichstein et al. 2005; Stoy et al. 2006; Valentini et al. 2000). However, many CO2 flux measurements over drylands indicate contributions of abiotic processes to the NEE (Emmerich 2003; Ferlan et al. 2011; Hastings et al. 2005; Inglima et al. 2009; Mielnick and Dugas 2000; Were et al. 2010) (Wohlfahrt et al. 2008; Xie et al. 2008).These processes can dominate the flux during the dry season (Kowalski et al. 2008) with annual contributions >50% depending mainly on meteorological conditions (Serrano-Ortiz et al. 2009). 3.1.2 Weathering processes Some soils are derived from carboniferous (calcareous) rocks and/or include additional carbonates (secondary carbonates or caliche) as a result of weathering and precipitation processes (Eq. 1). H 2O( aq) CO2( g ) CaCO3( s ) 2HCO3 Ca(2aq ) (1) In terms of NEE, Eq. (1) specifies that for each molecule of CaCO 3 dissolved a molecule of atmospheric CO2 is consumed. During precipitation, one molecule of CO2 is released for every molecule of CaCO3 deposited to the surface. Thus, either water lost by evapotranspiration or additional sources of Ca2+ would enrich aqueous concentrations and enhance the deposition of CaCO3 from the aqueous solution, and release CO2 to the atmosphere (Eq. (1) to the left). Globally, and over long time scales, weathering processes are balanced with respect to CO2 (Berner 2003; Lasaga et al. 1994). However, at annual and seasonal scales the predominance of dissolution or precipitation processes may be relevant to local NEEs (Serrano-Ortiz et al. 2010) and contribute to the observed CO2 fluxes (Emmerich 2003; Mielnick and Dugas 2000). In addition, although precipitation processes imply CO2 release at short time scales, caliche formation can be considered a net atmospheric CO2 sink over long timescales: every two molecules of bicarbonate, previously formed during the growing season by dissolution of two molecules of CO 2, react with one molecule of additional Ca2+ to form one molecule of calcite and release one molecule of CO2. The formation of this secondary C form depends on land use and soil/crop management systems (Lal 2004). Addition of biomass, whose decomposition increases the partial pressure of CO2 in the soil, together with irrigation, increases SIC in agricultural soils (Entry et al. 2004). While its contribution to the total annual atmospheric CO2 sink may be less than 10% (Eswaran et al. 2000; Liu and Zhao 2000), it is unclear how SIC responds to rainfall and temperature changes predicted under the climate change scenarios (Rasmussen 2006). 3.1.3 Subsoil ventilation processes Drylands over carbonate rocks with cracks, pores and cavities, together with soils with deep vadose-zones, show a high capacity to store CO2 belowground. Since such CO2 storage may represent as much as 60% of the annual atmospheric sink (Serrano-Ortiz et al. 2011), the subsurface can be considered a temporal depot for CO2 coming from different processes (mainly weathering and respiration) (Serrano-Ortiz et al. 2010). In addition to diffusion processes, such soils have the potential to emit the stored CO2 via ventilation (Sanchez-Cañete et al. 2011) and contribute to ecosystem CO2 exchange observed. Ventilation is a transport process due to net movements of air in and out of an enclosed space. The behaviour of ventilation processes, in caves for example, is controlled by the degree of connection between the cavities and the aboveground system (Cuezva et al. 2011) and, thus, such processes have only been detected when the soil is dry (Cuezva et al. 2011; Sanchez-Cañete et al. 2011; Serrano-Ortiz et al. 2009; Were et al. 2010). The main meteorological drivers controlling soil CO2 ventilation due to pressure pumping are wind speed and turbulence (Jassal et al. 2005; Lewicki et al. 2010). Therefore, the non-negligible role of subsurface as a temporal depot of CO2, along with seasonal ventilation can contribute to the annual net ecosystem carbon balance (Serrano-Ortiz et al. 2010). 3.2 Carbon sink capacity at ecosystem level The processes mentioned above (weathering, ventilation and erosion processes) act together in drylands and contribute to the measured annual net C exchange (Table 1). For deserts located in southwest of the U.S.A. and Baja California, published studies have determined that the most important driver controlling CO2 fluxes is not the amount of rainfall but mostly its timing (Hastings et al. 2005; Mielnick et al. 2005; Wohlfahrt et al. 2008). However, published annual net C exchanges are not in agreement. While two desert shrublands located in the Mojave Desert and Baja California, with similar annual precipitation, act as annual net C sinks of 106 ± 70 and 52 g C m-2 yr-1, respectively, the Chihuahuan Desert site emitted 145 g C m-2 annually. Grasslands located in North America (New Mexico, Arizona and California) and Europe (Southeast Portugal and Southeast of Spain) are C sources ranging from 141 (source) to -190 (sink) g C m-2 yr-1 depending mostly on the total amount of rainfall (Aires et al. 2008; Anderson-Teixeira et al. 2011) (Emmerich 2003; Ma et al. 2007; Scott et al. 2006) and also wind speed for the particular site located in Southeast of Spain (Rey et al. 2011). While ecosystem C sink capacity in grasslands located in Northern Asia also depend on optimal temperature in summer (10-20ºC) (Kato et al. 2006; Wang et al. 2008). For savannas, Scott et al. 2009 measured an annual net C release ranging from 14 to 95 g C m-2 yr-1 in a semiarid savannah in southern Arizona, while Ma et al. (2007) measured an annual net C uptake ranging from 56 to 155 g C m -2 yr-1 in a savannah site located in California with higher annual precipitation and lower temperature. Finally, shrublands in drylands act mostly as small sinks for atmospheric CO2 (uptake from 2 to 75 g C m-2 yr-1) (Anderson-Teixeira et al. 2011; Luo et al. 2007; Serrano-Ortiz et al. 2009; Zhao et al. 2006), with some exceptions. In riparian areas where shrubs and woody plants have the capacity to exploit water resources by growing deep roots (Domingo et al. 1999), annual NEE can be higher than 200 g C m-2 yr-1 (Scott et al. 2006). On the contrary, the annual C loss estimation for the Lucky Hills site located in Arizona was 144 g C m -2 yr-1. The source of this C appears to be from the large inorganic carbon pool in these soils (Emmerich 2003). In summary, according to published studies, the carbon sink capacity at ecosystem level in drylands is very variable depending on the ecosystem type, inorganic C pool and mostly on rainfall timing and temperature during growing season. Table 1 Annual net ecosystem C exchange (g C m-2) measured mostly using the eddy covariance technique, together with annual temperature and rainfall. Negative sign (-) for net ecosystem exchange indicates a C sink Vegetation Desert and bare soil Location Experimental Site Mean annual temperature (ºC) 20 Mean annual precipitation (mm) 210 Net ecosystem C exchange (g C m-2 yr-1) -106±70 Mojave Desert (USA) Desert on the Nevada Test Site, 120 km northwest of Las Vegas Wohlfahrt et al. 2008 Chihuahuan Desert (USA) About 40 km northeast of Las Cruces, New Mexico Mielnick et al. 2005 – 272 145*1 Baja California (Mexico) 15 km west of the city of La Paz and 1.5 km from the Bay of La Paz (CIBNOR) “Cabo de Gata Natural Park” Almería (Andalucía) Hastings et al. 2005 24*2 147 197 -39 -52 Rey et al. 2011 17 210 251 294 66 144 92 Southeast Portugal Monte do Tojal, Évora in Southern Portugal Aires et al. 2008 14.7 364 751 49 -190 New Mexico (USA) Sevilleta LTER in Central New Mexico 13 244 30 Arizona (USA) The Kendall grassland Agricultural Research Service Walnut Gulch Experimental Watershed AndersonTeixeira et al. 2011 Scott et al. 2010 14.5 Emmerich 2003 17 313 312 274 246 162 356 -69 -98 -55 -47 21 126*1 Foodplain terraces along the San Pedro River Scott et al. 2006 17 234 -63 Foothills of the Sierra Nevada Ma et al. 2007 17±1 562±193 Southwest Spain Grassland California (USA) 3 (-88)–141 Qinghai-Tibetan Plateau (China) Alpine meadow Kato et al. 2006 -0.65 -0.91 -1.53 561* -79 -92 -173 Northern China Inner Mongolia Autonomous region Wang et al. 2008 Central Mongolia Arizona (USA) Hentiy province of Mongolia The Santa Rita mesquite savanna site located on the Santa Rita Experimental Range (SRER) Li et al. 2005 Scott et al. 2009 California (USA) Southwest Spain Foothills of the Sierra Nevada Mediterranean plateau, 25 km from the coast Ma et al. 2007 Serrano-Ortiz et al. 2009 2.5 1.3 1.7 1.2 19 20 20 19 17±1 12 297 174 215 196 285 335 289 330 562±193 475 10 30 -15 -41 60 14 95 30 (-155) – (-56) -2±23 New Mexico (USA) Sevilleta LTER in Central New Mexico 14 244 -30 San Diego (USA) Southern California, 75 km east of Pacific Ocean AndersonTeixeira et al. 2011 Luo et al. 2007 15 349 -52 Arizona(USA) Foodplain terraces along the San Pedro River Scott et al. 2006 17 234 -212 Lucky Hills, Agricultural Research Service Walnut Gulch Experimental Watershed Emmerich 2003 17 256 144*1 Alpine Potentilla fruticosa at the Haibei Research Station Zhao et al. 2006 2.3 2.2 542 493 -59 -75 Savanna Shrubland Reference Qinghai-Tibetan Plateau (China) *1Data not measured by the eddy covariance technique *2 Estimated data using information of Figure 5.8.1 of the cited reference *3 Annual average precipitation for 1981-2000 Running projects - Balance de carbono en el olivar: efecto de la presencia de la cubierta vegetal (CARBOLIVAR). Proyecto de Excelencia. Junta de Andalucía. From 01/2012 to 12//2014. Principal Investigador: Andrew S. Kowalski (Profesor titular UGR) - Efecto del cambio global en el papel de las turberas como sumideros de carbono. Ministerio de Ciencia e Innovación, Plan nacional de I+D+I (2008-2011). Programa: Internacionalización de la I+D. Subprograma: proyectos internacionales. Modalidad: Proyectos de Movilidad (Acciones Integradas). From: 01/2012 to 12//2012. Principal Investigador: Penélope Serrano Ortiz (EEZA, CSIC, Almería, España). - Equipamiento para la cuantificación de los flujos de gases de efecto invernadero en ecosistemas. UNGR10-1E-107. Fondos de Infraestructura FEDER 2010.From: 01/2012 to 12//2012. Principal Investigador: Andrew S. Kowalski (Profesor titular UGR). - Red de monitorización de los flujos de carbono en ecosistemas mediterráneos españoles – cuantificación y estudio de procesos (CGL2010-22193-C04-02) Ministerio de Ciencia e Innovación. From: 01/2011 to 12/2013. 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