BGD Biogeosciences Discussions BGD Biogeosciences Discuss. 1810-6285 Göttingen, Germany 10.5194/bgd-10-851-2013 Biological soil crusts on initial soils: organic carbon dynamics and chemistry under temperate climatic conditions Dümig A. 1 Veste M. 2 Hagedorn F. 3 Fischer T. https://orcid.org/0000-0001-6235-2261 4 Lange P. 4 Spröte R. 5 Kögel-Knabner I. https://orcid.org/0000-0002-7216-8326 1 6 Lehrstuhl für Bodenkunde, Department für Ökologie und Ökosystemmanagement, Wissenschaftszentrum Weihenstephan für Ernährung, Landnutzung und Umwelt, Technische Universität München, 85350 Freising-Weihenstephan, Germany Centrum für Energietechnologie Brandenburg e.V., Friedlieb-Runge-Strasse 3, 03046 Cottbus, Germany Department of Biogeochemistry, WSL, Swiss Federal Institute of Forest, Snow and Landscape Research, Zürcherstr. 111, 8903, Birmensdorf, Switzerland Brandenburg University of Technology at Cottbus, Faculty of Environmental Sciences and Process Engineering, Central Analytical Laboratory, Konrad-Wachsmann-Allee 6, 03046 Cottbus, Germany Chair of Soil Protection and Recultivation, Brandenburg University of Technology, Konrad-Wachsmann-Allee 6, 03046 Cottbus, Germany Institute for Advanced Study, Technische Universität München, Lichtenbergstrasse 2a, 85748 Garching, Germany 16 01 2013 10 1 851 894 Copyright: © 2013 A. Dümig et al. 2013 This work is licensed under the Creative Commons Attribution 3.0 Unported License. To view a copy of this licence, visit https://creativecommons.org/licenses/by/3.0/ This article is available from https://bg.copernicus.org/preprints/10/851/2013/bgd-10-851-2013.html The full text article is available as a PDF file from https://bg.copernicus.org/preprints/10/851/2013/bgd-10-851-2013.pdf

Numerous studies have been carried out on the community structure and diversity of biological soil crusts (BSCs) as well as their important functions on ecosystem processes. However, the amount of BSC-derived organic carbon (OC) input into soils and its chemical composition under natural conditions has rarely been investigated. In this study, different development stages of algae- and moss-dominated BSCs were investigated on a~natural (<17 yr old BSCs) and experimental sand dune (<4 yr old BSCs) in northeastern Germany. We determined the OC accumulation in BSC-layers and the BSC-derived OC input into the underlying substrates for bulk materials and fractions <63 &mu;m. The chemical composition of OC was characterized by applying solid-state <sup>13</sup>C NMR spectroscopy and analysis of the carbohydrate-C signature.<sup>14</sup>C contents were used to assess the origin and dynamic of OC in BSCs and underlying substrates. Our results indicated a rapid BSC establishment and development from algae- to moss-dominated BSCs within only 4 yr under this temperate climate. The distribution of BSC types was presumably controlled by the surface stability according to the position in the slope. We found no evidence that soil properties influenced the BSC distribution on both sand dunes. <sup>14</sup>C contents clearly indicated the existence of two OC pools in BSCs and substrates, recent BSC-derived OC and lignite-derived "old" OC (biologically refractory). The input of recent BSC-derived OC strongly decreased the mean residence time of total OC. The downward translocation of OC into the underlying substrates was only found for moss-dominated BSCs at the natural sand dune which may accelerate soil formation at these spots. BSC-derived OC mainly comprised O-alkyl C (carbohydrate-C) and to a lesser extent also alkyl C and N-alkyl C in varying compositions. Accumulation of alkyl C was only detected in BSCs at the experimental dune which may induce a~lower water solubility of BSC-derived extracellular polymeric substances when compared to BSCs at the natural sand dune indicating that hydrological effects of BSCs on soils depend on the chemical composition of the extracellular polymeric substances.

 References Amelung,&nbsp;W., Cheshire,&nbsp;M.&nbsp;V., and Guggenberger,&nbsp;G.: Determination of neutral and acidic sugars in soil by capillary gas-liquid chromatography after trifluoroacetic acid hydrolysis, Soil Biol. Biochem., 28, 1631–1639, 1996. Amit,&nbsp;R. and Harrison,&nbsp;J.&nbsp;B.&nbsp;J.: Biogenic calcic horizon development under extremely arid conditions, Nizzana sand dunes, Israel, Adv. GeoEcol., 28, 65–88, 1995. Anderson,&nbsp;D.&nbsp;C., Harper,&nbsp;K.&nbsp;T., and Holmgren,&nbsp;R.&nbsp;C.: Recovery of cryptogamic soil crusts from grazing on Utah winter ranges, J. Range Man., 35, 355–359, 1982. Baldock,&nbsp;J.&nbsp;A., Oades,&nbsp;J.&nbsp;M., Vassallo,&nbsp;A.&nbsp;M., and Wilson,&nbsp;M.&nbsp;A.: Solid-state CP/MAS <sup>13</sup>C NMR analysis of bacterial and fungal cultures isolated from a&nbsp;soil incubated with glucose, Aust.&nbsp;J. Soil Res., 28, 213–225, 1990. Becker-Heidmann,&nbsp;P., Liang-Wu,&nbsp;L., and Scharpenseel,&nbsp;H.&nbsp;W.: Radiocarbon dating of organic matter fractions of a&nbsp;Chinese mollisol,&nbsp;Z. Pflanzenernähr. Bodenk., 151, 37–39, 1988. Belnap,&nbsp;J.: Nitrogen fixation in biological soil crusts from southeast Utah, USA, Biol. Fert. Soils, 35, 128–135, 2002. Belnap,&nbsp;J.: The potential roles of biological soil crusts in dryland hydrological cycles, Hydrol. Process., 20, 3159–3178, 2006. Belnap,&nbsp;J. and Eldridge,&nbsp;D.: Disturbance and recovery of biological soil crusts, in: Biological Soil Crusts: Structure, Function, and Management, edited by: Belnap,&nbsp;J. and Lange,&nbsp;O.&nbsp;L., Springer-Verlag Berlin, Heidelberg, 363–383, 2003. Belnap,&nbsp;J. and Gardner,&nbsp;J.&nbsp;S.: Soil microstructure in soils of the Colorado Plateau: the role of the cyanobacterium Microcoleus vaginatus, Great Basin Nat., 53, 40–47, 1993. Belnap,&nbsp;J. and Gillette,&nbsp;D.&nbsp;A.: Disturbance of biological soil crusts: impacts on potential wind erodibility of sandy desert soils in Southeastern Utah, Land Degrad. Develop., 8, 355–362, 1997. Belnap,&nbsp;J. and Lange,&nbsp;O.&nbsp;L.: Biological Soil Crusts: Structure, Function and Management, Ecological Studies 150, Springer, Berlin, 2001. Belnap,&nbsp;J., Phillips,&nbsp;S.&nbsp;L., and Miller,&nbsp;M.&nbsp;E.: Response of desert biological soil crusts to alterations in precipitation frequency, Oecolgia, 141, 306–316, 2004. Bertocchi,&nbsp;C., Navarini,&nbsp;L., Cesáro,&nbsp;A., and Anastasio,&nbsp;M.: Polysaccharides from cyanobacteria, Carbohydr. Polym., 12, 127–153, 1990. Beymer,&nbsp;R.&nbsp;J. and Klopatek,&nbsp;J.&nbsp;M.: Potential contribution of carbon by microphytic crusts in Pinyon-Juniper Woodlands, Arid Soil Res. Rehab., 5, 187–198, 1991. Beyschlag,&nbsp;W., Wittmann,&nbsp;M., Jentsch,&nbsp;A., and Steinlein,&nbsp;T.: Soil crusts and disturbance benefit plant germination, establishment and growth on nutrient deficient sand, Basic Appl. Ecol., 9, 243–252, 2008. Brankatschk,&nbsp;R., Fischer,&nbsp;T., Veste,&nbsp;M., and Zeyer,&nbsp;J.: Succession of N cycling processes in biological soil crusts on a&nbsp;Central European inland dune, FEMS Microbiol Ecol., 83, 149–160, https://doi.org/<a href="http://dx.doi.org/10.1111/j.1574-6941.2012.01459.x">10.1111/j.1574-6941.2012.01459.x</a>, 2012. Brock,&nbsp;T.&nbsp;D.: Effect of water potential on a&nbsp;<i>Microcoleus</i> (<i>Cyanophydeae</i>) from a&nbsp;desert crust, J. Phycol., 11, 316–320, 1975. Bronk Ramsey,&nbsp;C.: Development of the radiocarbon program OxCal, Radiocarbon, 43, 355–363, 2001. Castillo-Monroy,&nbsp;A.&nbsp;P., Bowker,&nbsp;M.&nbsp;A., Maestre,&nbsp;F.&nbsp;T., Rodríguez-Echeverría,&nbsp;S., Martínez,&nbsp;I., Barraza-Zepeda,&nbsp;C.&nbsp;E., and Escolar,&nbsp;C.: Relationships between biological soil crusts, bacterial diversity and abundance, and ecosystem functioning: insights from a&nbsp;semi-arid Mediterranean environment, J. Vegetat. Sci., 22, 165–174, 2001. Chamizo,&nbsp;S., Cantón,&nbsp;Y., Miralles,&nbsp;I., and Domingo,&nbsp;F.: Biological soil crust development affects physicochemical characteristics of soil surface in semiarid ecosystems, Soil Biol. Biochem., 49, 96–105, 2012. Chen,&nbsp;R., Zhang,&nbsp;Y., Li,&nbsp;Y., Wei,&nbsp;W.,&nbsp;Zhang,&nbsp;J., and Wu,&nbsp;N.: The variation of morphological features and mineralogical components of biological soil crusts in the Gurbantunggut Desert of Northwestern China, Environ. Geol., 57, 1135–1143, 2009. Dojani,&nbsp;S., Büdel,&nbsp;B., Deutschewitz,&nbsp;K., and Weber,&nbsp;B.: Rapid succession of biological soil crusts after experimental disturbance in the Succulent Karoo, South Africa, Appl. Soil Ecol., 48, 263–269, 2011. Drever,&nbsp;J.&nbsp;I. and Stillings,&nbsp;L.&nbsp;L.: The role of organic acids in mineral weathering, Colloid. Surf. A: Physicochem. Eng. Asp., 120, 167–181, 1997. Duncan,&nbsp;T.&nbsp;M.: <sup>13</sup>C chemical shielding in solids, J. Phys. Chem. Ref. Data, 16, 125–151, 1987. Eder,&nbsp;E., Spielvogel,&nbsp;S., Kölbl,&nbsp;A., Albert,&nbsp;G., and Kögel-Knabner,&nbsp;I.: Analysis of hydrolysable neutral sugars in mineral soils: improvement of alditol acetylation for gas chromatographic separation and measurement, Org. Geochem., 41, 580–585, 2010. Elbert,&nbsp;W., Weber,&nbsp;B., Büdel,&nbsp;B., Andreae,&nbsp;M.&nbsp;O., and Pöschl,&nbsp;U.: Microbiotic crusts on soil, rock and plants: neglected major players in the global cycles of carbon and nitrogen?, Biogeosciences Discuss., 6, 6983–7015, https://doi.org/<a href="http://dx.doi.org/10.5194/bgd-6-6983-2009">10.5194/bgd-6-6983-2009</a>, 2009. Elbert,&nbsp;W., Weber,&nbsp;B., Burrow,&nbsp;S., Steinkamp,&nbsp;J., Büdel,&nbsp;B., Andreae,&nbsp;M.&nbsp;O., and Pöschl,&nbsp;U.: Contribution of cryptogamic covers to the global cycles of carbon and nitrogen, Nat. Geosci., 5, 459–462, 2012. Eldridge,&nbsp;D.&nbsp;J.: Biological soil crusts and water relations in Australia deserts, in: Biological Soil Crusts: Structure, Function and Management, edited by: Belnap,&nbsp;J. and Lange,&nbsp;O.&nbsp;L., Springer, Berlin, 315–325, 2003. Eldridge,&nbsp;D.&nbsp;J., and Greene,&nbsp;R.&nbsp;S.&nbsp;B.: Microbiotic soil crusts: a&nbsp;review of their roles in soil and ecological processes in the rangelands of Australia, Aust.&nbsp;J. Soil Res., 32, 389–415, 1994. Evans,&nbsp;R.&nbsp;D. and Johansen,&nbsp;J.&nbsp;R.: Microbiotic crusts and ecosystem processes, Crit. Rev. Plant Sci., 18, 183–225, 1999. Evans,&nbsp;R.&nbsp;D. and Lange,&nbsp;O.&nbsp;L.: Biological soil crusts and ecosystem nitrogen and carbon dynamics, in: Biological Soil Crusts: Structure, Function and Management, edited by: Belnap,&nbsp;J. and Lange,&nbsp;O.&nbsp;L., Springer-Verlag, Berlin, 2003. Fischer,&nbsp;T., Veste,&nbsp;M., Schaaf,&nbsp;W., Dümig,&nbsp;A., Kögel-Knabner,&nbsp;I., Wiehe,&nbsp;W., Bens,&nbsp;O., and Hüttl,&nbsp;R.&nbsp;F.: Initial pedogenesis in a&nbsp;topsoil crust 3 <abbr>years</abbr> after construction of an artificial catchment in Brandenburg, NE Germany, Biogeochemistry, 101, 165–176, 2010a. Fischer,&nbsp;T., Veste,&nbsp;M., Wiehe,&nbsp;W., and Lange,&nbsp;P.: Water repellency and pore clogging at early successional stages of biological soil crusts on inland dunes, Brandenburg, NE Germany,&nbsp;Catena, 80, 47–52, 2010b. Fischer,&nbsp;T.,&nbsp;Veste,&nbsp;M., Eisele,&nbsp;A., Bens,&nbsp;O., Spyra,&nbsp;W., and Hüttl,&nbsp;R.&nbsp;F.: Small scale spatial heterogeneity of normalized difference vegetation indices (NDVI) and hot spots of photosynthesis in biological soil crusts, Flora, 207, 159–167, 2012. Flemming,&nbsp;H.-C. and Wingender,&nbsp;J.: The biofilm matrix, Nat. Rev. Microbiol., 8, 623–633, 2010. Fröberg,&nbsp;M., Tipping,&nbsp;E., Stendahl,&nbsp;J., Clarke,&nbsp;N., and Bryant,&nbsp;Ch.: Mean residence time of O horizon carbon along a&nbsp;climatic gradient in Scandinavia estimated by <sup>14</sup>C measurements of archived soils, Biogeochemistry, 104, 227–236, 2011. Harper,&nbsp;K.&nbsp;T. and Marble,&nbsp;J.&nbsp;R.: A&nbsp;role for nonvascular plants in management of arid and semiarid rangelands, in: Applications of Plant Sciences to Rangeland Management and Inventory, edited by: Tuller,&nbsp;P.&nbsp;T., Kluwer, Amsterdam, 135–169, 1988. Hatcher,&nbsp;P.&nbsp;G., Wilson,&nbsp;M.&nbsp;A., Vasallo,&nbsp;A.&nbsp;M., and Lerch,&nbsp;H.&nbsp;E.: Studies of angiospermous woods in Australian brown coals by nuclear magnetic resonance and analytical pyrolysis: new insights into early coalification, Int.&nbsp;J. Coal Geol., 13, 99–126, 1989. Ingerpuu,&nbsp;N., Liira,&nbsp;J., and Pärtel,&nbsp;M.: Vascular plants facilitated bryophytes in a&nbsp;grassland experiment, Plant Ecol., 180, 69–75, 2005. Jeffries,&nbsp;D.&nbsp;L., Link,&nbsp;S.&nbsp;O., and Klopatek,&nbsp;J.&nbsp;M.: CO<sub>2</sub> fluxes of cryptogamic crusts. I. response to resaturation, New Phytol., 125, 163–173, 1993. Jentsch,&nbsp;A. and Beyschlag,&nbsp;W.: Vegetation ecology of dry acidic grasslands in the lowland area of central Europe, Flora, 189, 3–26, 2003. Kidron,&nbsp;G.&nbsp;J., Vonshak,&nbsp;A., and Abeliovich,&nbsp;A.: Microbiotic crusts as biomarkers for surface stability and wetness duration in the Negev Desert, Earth Surf. Process. Landf., 34, 1594–1604, 2009. Kidron,&nbsp;G.&nbsp;J., Vonshak,&nbsp;A., Dor,&nbsp;I., Barinova,&nbsp;S., and Abeliovich,&nbsp;A.: Properties and spatial distribution of microbiotic crusts in the Negev Desert, Israel, Catena, 82, 92–101, 2010. Klouda,&nbsp;B., Lewis,&nbsp;C.&nbsp;W., Rasmussen,&nbsp;R.&nbsp;A., Rhoderick,&nbsp;G.&nbsp;C., Sams,&nbsp;R.&nbsp;L., Stevens,&nbsp;R.&nbsp;K., Currie,&nbsp;L.&nbsp;A., Donahue,&nbsp;D.&nbsp;J., Jull,&nbsp;T.&nbsp;A.&nbsp;J., and Seila,&nbsp;R.&nbsp;L.: Radiocarbon measurements of atmospheric volatile organic compounds: quantifying the biogenic contribution, Environ. Sci. Technol., 30, 1098–1105, 1996. Knicker,&nbsp;H. and Lüdemann, H-D.: N-15 and C-13 CPMAS and solution NMR studies of N-15 enriched plant material during 600 days of microbial degradation, Org. Geochem., 23, 329–341, 1995. Kögel-Knabner,&nbsp;I.: <sup>13</sup>C and <sup>15</sup>N NMR spectroscopy as a&nbsp;tool in soil organic matter studies, Geoderma, 80, 243–270, 1997. Kögel-Knabner,&nbsp;I., Hatcher,&nbsp;P.&nbsp;G., Tegelaar,&nbsp;E.&nbsp;W., and De Leeuw,&nbsp;J.&nbsp;W.: Aliphatic components of forest soil organic matter as determined by solid-state <sup>13</sup>C NMR and analytical pyrolysis, Sci. Total Environ., 113, 89–106, 1992. Kroy,&nbsp;K., Sauermann,&nbsp;G., and Herrmann,&nbsp;H.&nbsp;J.: Minimal model for sand dunes, Phys. Rev. Lett., 88, 1– 4, 2002. Lange,&nbsp;O.&nbsp;L., Kidron,&nbsp;G., Büdel,&nbsp;B., Meyer,&nbsp;A., Kilian,&nbsp;E., and Abeliovich,&nbsp;A.: Taxonomic composition and photosynthetic characteristics of the "biological soil crusts" covering sand dunes in the western Negev Desert, Funct. Ecol., 6, 519–527, 1992. Lange,&nbsp;O.&nbsp;L., Belnap,&nbsp;J., Reichenberger,&nbsp;H., and Meyer,&nbsp;A.: Photosynthesis of green algal soil crust lichens from arid lands in southern Utah, USA: role of water content on light and temperature response of CO<sub>2</sub> exchange, Flora, 192, 1–15, 1997. Langhans,&nbsp;T.&nbsp;M., Storm,&nbsp;C., and Schwabe,&nbsp;A.: Community assembly of biological soil crusts of different successional stages in a&nbsp;temperate sand ecosystem, as assessed by direct determination and enrichment techniques, Microbial Ecol., 58, 394–407, 2009. Lázaro,&nbsp;R., Cantón,&nbsp;Y., Solé-Benet,&nbsp;A., Bevan,&nbsp;J., Alexander,&nbsp;R., Sancho,&nbsp;L.&nbsp;G., and Puigdefábregas,&nbsp;J.: The influence of competition between lichen colonization and erosion on the evolution of soil surfaces in the Tabernas badlands (SE Spain) and its landscape effect, Geomorphology, 102, 252–266, 2008. Levin,&nbsp;I. and Kromer,&nbsp;B.: The tropospheric <sup>14</sup>CO<sub>2</sub> level in mid-latitudes of the Northern Hemisphere (1959–2003), Radiocarbon, 46, 1261–1272, 2004. Löbel,&nbsp;S., Dengler,&nbsp;J., and Hobohm,&nbsp;C.: Species richness of vascular plants, bryophytes and lichens in dry grasslands: the effects of environment, landscape structure and competition, Folia Geobot., 41, 377–393, 2006. Maestre,&nbsp;F.&nbsp;T., Huesca,&nbsp;M.&nbsp;T., Zaady,&nbsp;E., Bautista,&nbsp;S., and Cortina,&nbsp;J.: Infiltration, penetration resistance and microphytic crust composition in contrasted microsites within a&nbsp;Mediterranean semi-arid steppe, Soil Biol. Biochem., 34, 895–898, 2002. Maestre,&nbsp;F.&nbsp;T., Bowker,&nbsp;M.&nbsp;A., Cantón,&nbsp;Y., Castillo-Monroy,&nbsp;A.&nbsp;P., Cortina,&nbsp;J., Escolar,&nbsp;C., Escudero,&nbsp;A., Lázaro,&nbsp;R., and Martínez,&nbsp;I.: Ecology and functional roles of biological soil crusts in semi-arid ecosystems of Spain, J. Arid Environ., 75, 1282–1291, 2011. Mager,&nbsp;D.: Carbohydrates in cyanobacterial soil crusts as a&nbsp;source of carbon in the southwest Kalahari, Botswana, Soil Biol Biochem. 42, 313–318, 2010. Mazor,&nbsp;G., Kidron,&nbsp;G.&nbsp;J., Vanshak,&nbsp;A., and Abeliovich,&nbsp;A.: The role of cyanobacterial exopolysaccharides in structuring desert microbial crusts, FEMS Microbiol. Ecol., 21, 121–130, 1996. Mehra,&nbsp;O.&nbsp;P. and Jackson,&nbsp;M.&nbsp;L.: Iron oxide removal from soils and clays by a&nbsp;dithionite–citrate system buffered with sodium bicarbonate, in: Proceedins of the 7th National Conference on Clays and Clay Minerals, edited by Ingerson,&nbsp;E., Pergamon Press, London, 317–327, 1960. Nierop,&nbsp;K.&nbsp;G.&nbsp;J., van Lagen,&nbsp;B., and Buurman,&nbsp;P.: Composition of plant tissues and soil organic matter in the first stages of a&nbsp;vegetation succession, Geoderma, 100, 1–24, 2001. Oades&nbsp;J.&nbsp;M.: Soil organic matter and structural stability: mechanisms and implications for management, Plant Soil, 76, 319–337, 1984. Paul,&nbsp;E.&nbsp;A., Follett,&nbsp;R.&nbsp;F., Leavitt,&nbsp;S.&nbsp;W., Halvorson,&nbsp;A., Peterson,&nbsp;G.&nbsp;A., and Lyon,&nbsp;D.&nbsp;J.: Radiocarbon dating for determination of soil organic matter pool sizes and dynamics, Soil Sci. Soc. Am.&nbsp;J., 61, 1058–1067, 1997. Peersen,&nbsp;O.&nbsp;B., Wu,&nbsp;X., Kustanovich,&nbsp;I., and Smith,&nbsp;S.&nbsp;O.: Variable-amplitude cross-polarization MAS NMR, J. Magn. Reson., 104, 334–339, 1993. Popper,&nbsp;Z.&nbsp;A. and Fry,&nbsp;S.&nbsp;C.: Primary cell wall composition of bryophytes and charophytes, Ann. Bot-London, 91, 1–12, 2003. Prasse,&nbsp;R. and Bornkamm,&nbsp;R.: Effect of microbiotic soil surface crusts on emergence of vascular plants, Plant Ecol., 150, 65–75, 2000. Prietzel,&nbsp;J., Dechamps,&nbsp;N., and Spielvogel,&nbsp;S.: Analysis of non-cellulosic polysaccharides helps to reveal the history of thick organic surface layers on calcareous Alpine soils, Plant Soil, https://doi.org/<a href="http://dx.doi.org/10.1007/s11104-012-1340-2">10.1007/s11104-012-1340-2</a>, 2012. Reynolds,&nbsp;R., Belnap,&nbsp;J., Reheis,&nbsp;M., Lamothe,&nbsp;P., and Luiszer,&nbsp;F.: Aeolian dust in Colorado Plateau soils: nutrient inputs and recent change in source, Proc. Natl. Acad. Sci. USA, 98, 7123–7127, 2001. Rossi,&nbsp;F., Potrafka,&nbsp;R.&nbsp;M., Pichel,&nbsp;F.&nbsp;G., and De Philippis,&nbsp;R.: The role of the exopolysaccharides in enhancing hydraulic conductivity of biological soil crusts, Soil Biol. Biochem., 46, 33–40, 2012. Rumpel,&nbsp;C. and Dignac,&nbsp;M.-F.: Gas chromatographic analysis of monosaccharides in a&nbsp;forest soil profile: analysis by gas chromatography after trifluoroacetic acid hydrolysis and reduction-acetylation, Soil Biol. Biochem., 38, 1478–1481, 2006. Rumpel,&nbsp;C., Knicker,&nbsp;H., Kögel-Knabner,&nbsp;I., and Hüttl,&nbsp;R.&nbsp;F.: Airborne contamination of immature soil (Lusatian mining district) by lignite-derived materials: its detection and contribution to the soil organic matter budget, Water Air Soil Pollut., 105, 481–492, 1998. Rumpel,&nbsp;C., Balesdent,&nbsp;J., Grootes,&nbsp;P., Weber,&nbsp;E., and Kögel-Knabner,&nbsp;I.: Quantification of lignite- and vegetation-derived soil carbon using <sup>14</sup>C activity measurements in a&nbsp;forested chronosequence, Geoderma, 112, 155–166, 2003. Schaaf,&nbsp;W., Bens,&nbsp;B., Fischer,&nbsp;A., Gerke,&nbsp;H.&nbsp;H., Gerwin,&nbsp;W., Grünewald,&nbsp;U., Holländer,&nbsp;H.&nbsp;M., Kögel-Knabner,&nbsp;I., Mutz,&nbsp;M., Schloter,&nbsp;M., Schulin,&nbsp;R., Veste,&nbsp;M., Winter,&nbsp;S., and Hüttl,&nbsp;R.&nbsp;F.: Patterns and processes of initial terrestrial-ecosystem development, J. Plant Nutr. Soil Sc., 174, 229–239, 2011. Schaefer,&nbsp;J. and Stejskal,&nbsp;E.&nbsp;O.: Carbon-13 nuclear magnetic resonance of polymers spinning at magic angle, J. Am. Chem. Soc., 98, 1031–1032, 1976. Scharpenseel,&nbsp;H.&nbsp;W. and Becker-Heidmann,&nbsp;P.: Twenty-five years of radiocarbon dating soils: paradigm of erring and learning, Radiocarbon, 34, 541–549, 1992. Scharpenseel,&nbsp;H.-W. and Becker-Heidmann,&nbsp;P.: Shifts in <sup>14</sup>C patterns of soil properties due to bomb carbon, including effects of morphogenetic and turbation processes, Radiocarbon, 31, 627–636, 1997. Schwertmann,&nbsp;U.: Differenzierung der Eisenoxide des Bodens durch Extraktion mit Ammoniumoxalat-Lösung,&nbsp;Z. Pflanzenernähr. Düng. Bodenkd., 105, 194–202, 1964. Spröte,&nbsp;R., Fischer,&nbsp;T., Veste,&nbsp;M., Raab,&nbsp;T., Wiehe,&nbsp;W., Lange, Ph., Bens,&nbsp;O., and Hüttl,&nbsp;R.&nbsp;F.: Biological topsoil crusts at early successional stages on Quaternary substrates dumped by mining in Brandenburg, NE Germany, Geomorphologie, 4, 359–370, 2010. Stach,&nbsp;E., Mackowsky,&nbsp;M.-T., Teichmüller,&nbsp;M., Taylor,&nbsp;G.&nbsp;H., Chandra,&nbsp;D., and Teichmüller,&nbsp;R.: Stach's Textbook of Coal Petrology, Gebrüder Borntraeger, Berlin, 1982. Stuiver,&nbsp;M. and Polach,&nbsp;H.&nbsp;A.: Discussions of reporting <sup>14</sup>C data, Radiocarbon, 19, 355–363, 1977. Terauchi,&nbsp;K. and Kondo,&nbsp;T.: The cyanobacterial Circadian Clock and the KaiC Phosphorylation cycle, in: The Cyanobacteria: Molecular Biology, Genomics and Evolution, edited by: Herrero,&nbsp;A. and Flores,&nbsp;E., Caister Academic Press, Norfolk, 199–215, 2008. Trumbore,&nbsp;S.&nbsp;E.: Applications of accelerator mass spectrometry to soil science, in: Mass Spectrometry of Soils, edited by: Boutton,&nbsp;T.&nbsp;W. and Yamasaki,&nbsp;S., Marcel Dekker, New York, 311–340, 1996. Trumbore,&nbsp;S.: Age of soil organic matter and soil respiration: radiocarbon constraints on belowground C dynamics, Ecol. Appl., 10, 399–411, 2000. Trumbore,&nbsp;S.: Radiocarbon and soil carbon dynamics, Annu. Rev. Earth Planet. Sci., 37, 12.1–12.20, 2009. Trumbore,&nbsp;S.&nbsp;E., Vogel,&nbsp;J.&nbsp;S., and Southon,&nbsp;J.&nbsp;R.: AMS <sup>14</sup>C measurements of fractionated soil organic matter: an approach to deciphering the soil carbon cycle, Radiocarbon, 31, 644–654, 1989. Trumbore,&nbsp;S.&nbsp;E., Davidson,&nbsp;E.&nbsp;A., de Camargo,&nbsp;P.&nbsp;B., Nepstad,&nbsp;D.&nbsp;C., and Martinelli,&nbsp;L.&nbsp;A.: Belowground cycling of carbon in forests and pastures of eastern Amazonia, Global Biogeochem. Cy., 9, 515–528, 1995. Veste,&nbsp;M. and Littmann,&nbsp;T.: Dewfall and its geo-ecological implication for biological surface crusts in desert sand dunes (north-western Negev, Israel), J. Arid Land Stud., 16, 139–147, 2006. Veste,&nbsp;M., Littmann,&nbsp;T., Breckle,&nbsp;S.-W., and Yair,&nbsp;A.: The role of biological soil crusts on desert sand dunes of the north-western Negev (Israel), in: Sustainable Land-Use in Deserts, edited by: Breckle,&nbsp;S.-W., Veste,&nbsp;M., and Wucherer,&nbsp;W., Springer, Heidelberg, New York, Tokyo, 357–367, 2001a. Veste,&nbsp;M., Littmann,&nbsp;T., Friedrich,&nbsp;H., and Breckle,&nbsp;S.-W.: Microclimatic boundary conditions for activity of soil lichen crusts in sand dunes of the north-western Negev desert, Israel, Flora 196, 465–476, 2001b. Veste,&nbsp;M., Breckle,&nbsp;S.-W., Eggert,&nbsp;K., and Littmann,&nbsp;T.: Vegetation pattern in arid sand dunes controlled by biological soil crusts along a&nbsp;climatic gradient in the Northern Negev desert, Basic Appl. Dryland Res., 5, 1–16, 2011. von Lützow, M., Kögel-Knabner, I., Ekschmitt, K., Flessa, H., Guggenberger, G., Matzner, E., and Marschner, B.: SOM fractionation methods: relevance to functional pools and to stabilization mechanisms, Soil Biol. Biochem., 39, 2183–2207, 2007. Wilske,&nbsp;B., Burgheimer,&nbsp;J., Karnieli,&nbsp;A., Zaady,&nbsp;E., Andreae,&nbsp;M.&nbsp;O., Yakir,&nbsp;D., and Kesselmeier,&nbsp;J.: The CO<sub>2</sub> exchange of biological soil crusts in a&nbsp;semiarid grass-shrubland at the northern transition zone of the Negev desert, Israel, Biogeosciences, 5, 1411–1423, https://doi.org/<a href="http://dx.doi.org/10.5194/bg-5-1411-2008">10.5194/bg-5-1411-2008</a>, 2008. Wilson,&nbsp;M.&nbsp;A., Pugmire,&nbsp;R.&nbsp;J., and Grant,&nbsp;D.&nbsp;M.: Nuclear magnetic resonance spectroscopy of soils and related materials. Relaxation of <sup>13</sup>C nuclei in cross polarization nuclear magnetic resonance experiments, Org. Geochem., 5, 121–129, 1983. Yair,&nbsp;A., Almog,&nbsp;R., and Veste,&nbsp;M.: Differential hydrological response of biological topsoil crusts along a&nbsp;rainfall gradient in a&nbsp;sandy arid area: Northern Negev desert, Israel, Catena, 87, 326–333, 2011. Zaady,&nbsp;E., Ben-David,&nbsp;E.&nbsp;A., Sher,&nbsp;Y., Tzirkin,&nbsp;R., and Nejidat,&nbsp;A.: Inferring biological soil crust successional stage using combined PLFA, DGGE, physical and biophysiological analyses, Soil Biol. Biochem., 42, 842–849, 2010. Zhao,&nbsp;H.-L., Guo,&nbsp;Y.-R., Zhou,&nbsp;R.-L., and Drake,&nbsp;S.: Biological soil crust and surface soil properties in different vegetation types of Horqin Sand Land, China, Catena, 82, 70–76, 2010.