Climate and small scale factors determine functional diversity shifts of biological soil crusts in Iberian drylands

Austin AT, Yahdjian L, Stark JM, Belnap J, Porporato A, Norton U, Ravetta D, Schaeffer SM (2003) Water pulses and biogeochemical cycles in arid and semiarid ecosystems. Oecologia 141:221–235 Bates D, Maechler M, Bolker B (2012) lme4: linear mixed-effects models using S4classes. http://CRAN.R-project.org/package=lme4 Belnap J (2006) The potential roles of biological soil crusts in dryland hydrologic cycles. Hydrol Process 20:3159–3178 Belnap J, Lange OL (2003) Biological soil crusts: structure, function, and management. Springer, Berlin Botta-Dukát Z (2005) Rao’s quadratic entropy as a measure of functional diversity based on multiple traits. J Veg Sci 16:533–540 Bowker M, Belnap J, Chaudhary VB, Johnson NC (2008) Revisiting classic water erosion models in drylands: the strong impact of biological soil crusts. Soil Biol Biochem 40:2309–2316 Bowker MA, Maestre FT, Escolar C (2010a) Biological crusts as model system for examining the biodiversity-ecosystem function relationship in soils. Soil Biol Biochem 42:405–417 Bowker MA, Soliveres S, Maestre FT (2010b) Competition increases with abiotic stress and regulates the diversity of biological soil crusts. J Ecol 98:551–560 Bowker MA, Maestre FT, Mau RL (2013) Diversity and patch-size distributions of biological soil crusts regulate dryland ecosystem multifunctionality. Ecosystems 16:1–11 Chaudhary VB, Bowker MA, O’Dell TE, Grace JB, Redman AE, Rillig MC, Johnson NC (2009) Untangling the biological contributions to soil stability in semiarid shrublands. Ecol Appl 19:110–122 Chave J, Coomes D, Jansen S, Lewis SL, Swenson NG, Zanne AE (2009) Towards a worldwide wood economics spectrum. Ecol Lett 12:351–366 Concostrina-Zubiri L, Martínez I, Rabasa SG, Escudero A (2013a) The influence of environmental factors on biological soil crust: from a community perspective to a species level approach. J Veg Sci. doi:10.1111/jvs.12084 Concostrina-Zubiri L, Huber-Sannwald E, Martínez I, Flores Flores JL, Escudero A (2013b) Biological soil crusts greatly contribute to small-scale soil heterogeneity along a grazing gradient. Soil Biol Biochem 64:28–36 Cornelissen J, Lavorel S, Garnier E et al (2003) A handbook of protocols for standardised and easy measurement of plant functional traits worldwide. Aust J Bot 51:335–380 Cornelissen JHC, Lang SI, Soudzilovskaia NA, During HJ (2007) Comparative cryptogam ecology: a review of bryophyte and lichen traits that drive biogeochemistry. Ann Bot 99:987–1001 Cornwell WK, Ackerly DD (2009) Community assembly and shifts in plant trait distributions across an environmental gradient in coastal California. Ecol Monogr 79:109–126 Cornwell WK, Schwilk DW, Ackerly DD (2006) A trait-based test for habitat filtering: convex hull volume. Ecology 87:1465–1471 Darby BJ, Neher DA, Belnap J (2010) Impact of biological soil crusts and desert plants on soil microfaunal community composition. Plant Soil 328:421–431 de Bello F, Lavorel S, Díaz S, Harrington R, Cornelissen JHC et al (2010a) Towards an assessment of multiple ecosystem processes and services via functional traits. Biodivers Conserv 19:2873–2893 de Bello F, Lavergne S, Meynard CN, Lepš J, Thuiller W (2010b) The partitioning of diversity: showing Theseus a way out of the labyrinth. J Veg Sci 21:992–1000 de Bello F, Lavorel S, Lavergne S, Albert CH, Boulangeat I et al (2013) Hierarchical effects of environmental filters on the functional structure of plant communities: a case study in the French Alps. Ecography 36:393–402 Delgado-Baquerizo M, Maestre FT, Gallardo A (2013) Biological soil crusts increase the resistance of soil nitrogen dynamics to changes in temperatures in a semi-arid ecosystem. Plant Soil 366:35–47 Dias AT, Berg MP, de Bello F, Oosten AR, Bíla K, Moretti M (2012) An experimental framework to identify community functional components driving ecosystem processes and services delivery. J Ecol 101:29–37 Díaz S, Lavorel S, de Bello F, Quétier F, Grigulis K, Robson TM (2007) Incorporating plant functional diversity effects in ecosystem service assessments. PNAS 104:20684–20689 Elbert W, Weber B, Burrows S, Steinkamp J, Büdel B et al (2012) Contribution of cryptogamic covers to the global cycles of carbon and nitrogen. Nat Geosci 5:459–462 Eldridge DJ, Greene RSB (1994) Microbiotic soil crusts: a review of their roles in soil and ecological processes in the rangelands of Australia. Aust J Soil Res 32:389–415 Eldridge DJ, Rosentreter R (1999) Morphological groups: a framework for monitoring microphytic crusts in arid landscapes. J Arid Environ 41:11–25 Eldridge DJ, Tozer ME (1997) Environmental factors relating to the distribution of terricolous bryophytes and lichens in semi-arid eastern Australia. Bryologist 100:28–39 Ellis CJ, Coppins BJ (2006) Contrasting functional traits maintain lichen epiphyte diversity in response to climate and autogenic succession. J Biogeogr 33:1643–1656 Escolar C, Martinez I, Bowker MA, Maestre FT (2012) Warming reduces the growth and diversity of biological soil crusts in a semi-arid environment: implications for ecosystem structure and functioning. Philos Trans R Soc B 367:3087–3099 Escudero A, Martínez I, de la Cruz A, Otálora M, Maestre FT (2007) Soil lichens have species-specific effects on the seedling emergence of three gypsophile plant species. J Arid Environ 70:18–28 Garnier E, Cortez J, Billès G, Navas ML, Roumet C et al (2004) Plant functional markers capture ecosystem properties during secondary succession. Ecology 85:2630–2637 Garnier E, Lavorel S, Ansquer P, Castro H, Cruz P et al (2007) Assessing the effects of land-use change on plant traits, communities and ecosystem functioning in grasslands: a standardized methodology and lessons from an application to 11 European sites. Ann Bot 99:967–985 Giordani P, Brunialti G, Bacaro G, Nascimben J (2012) Functional traits of epiphytic lichens as potential indicators of environmental conditions in forest ecosystems. Eco Indic 18:413–420 Giordani P, Incerti G, Rizzi G, Rellini I, Nimis PL, Modenesi P (2013) Functional traits of cryptogams in Mediterranean ecosystems are driven by water, light and substrate interactions. J Veg Sci. doi:10.1111/jvs.12119 Harper KT, Belnap J (2001) The influence of biological soil crusts on mineral uptake by associated vascular plants. J Arid Environ 47:347–357 Hauck M, Jürgens SR, Willenbruch K, Huneck S, Leuschner C (2009) Dissociation and metal-binding characteristics of yellow lichen substances suggest a relationship with site preferences of lichens. Ann Bot 103:13–22 Jiménez Aguilar A, Huber–Sannwald E, Belnap J, Smart DR, Moreno JTA (2009) Biological soil crusts exhibit a dynamic response to seasonal rain and release from grazing with implications for soil stability. J Arid Environ 73:1158–1169 Karnieli A, Kokaly RF, West NE, Clark RN (2003) Remote sensing of biological soil crusts. In: Belnap J, Lange OL (eds) Biological soil crusts, structure, function, and management, 2nd edn. Springer, Berlin, pp 431–455 Lakatos M, Rascher U, Büdel B (2006) Functional characteristics of corticolous lichens in the understory of a tropical lowland rain forest. New Phytol 172:679–695 Laliberté E, Legendre P (2010) A distance-based framework for measuring functional diversity from multiple traits. Ecology 91:299–305 Lalley JS, Viles HA (2005) Terricolous lichens in the northern Namib Desert of Namibia: distribution and community composition. Lichenol 37:77–92 Lavorel S, Grigulis K, McIntyre S et al (2008) Assessing functional diversity in the field-methodology matters! Funct Ecol 22:134–147 Loreau M, Mouquet N, Gonzalez A (2003) Biodiversity as spatial insurance in heterogeneous landscapes. PNAS 100:12765–12770 MA (Millennium Ecosystem Assessment) (2005) Ecosystems and human well-being: desertification synthesis. Island Press, Washington, DC Maestre FT, Escolar C, Martínez I, Escudero A (2008) Are soil lichen communities structured by biotic interactions? A null model analysis. J Veg Sci 19:261–266 Maestre FT, Bowker MA, Puche MD, Hinojosa MB, Martínez I et al (2009) Shrub encroachment can reverse desertification in semi-arid Mediterranean grasslands. Ecol Lett 12:930–941 Malam Issa O, Défarge C, Trichet J, Valentin C, Rajot JL (2009) Microbiotic soil crusts in the Sahel of Western Niger and their influence on soil porosity and water dynamics. Catena 77:48–55 Martínez I, Escudero A, Maestre FT, de la Cruz A, Guerrero C, Rubio A (2006) Small-scale patterns of abundance of mosses and lichens forming biological soil crusts in two semi-arid gypsum environments. Aust J Bot 54:339–348 Mason NW, Bello F, Mouillot D, Pavoine S, Dray S (2012) A guide for using functional diversity indices to reveal changes in assembly processes along ecological gradients. J Veg Sci 24:794–806 Mason NW, Mouillot D, Lee WG, Wilson JB (2005) Functional richness, functional evenness and functional divergence: the primary components of functional diversity. Oikos 111:112–118 McCulloch CE, Searle SR (2001) Generalized linear mixed models (GLMMs). In: McCulloch CE, Searle SR, Neuhaus JM (eds) Generalized, linear, and mixed models, 2nd edn. Wiley, New York, pp 220–246 Miller ME, Belote RT, Bowker MA, Garman S (2011) Alternative states of a semiarid grassland ecosystem: Implications for ecosystem services. Ecosphere 5: art55 Mokany K, Ash J, Roxburgh S (2008) Functional identity is more important than diversity in influencing ecosystem processes in a temperate native grassland. J Ecol 96:884–893 Molnár K, Farkas E (2010) Current results on biological activities of lichen secondary metabolites: a review. Z Naturforsch C 65:157–173 Mouchet MA, Villéger S, Mason NWH, Mouillot D (2010) Functional diversity measures: an overview of their redundancy and their ability to discriminate community assembly rules. Funct Ecol 24:867–876 Nash TH (2008) Lichen Biology. Cambridge University Press, Cambridge Nimis PL, Martellos S (2004) Keys to the lichens of Italy I. Terricolous species. Edizioni Goliardiche. Trieste, Italy Nimis PL, Martellos S (2008) ITALICA. The information system on Italian lichens. Version 4.0. http://dbiodbs.univ.trieste.it/italic/italic03. Accessed 24 June 2013 Ochoa–Hueso R, Hernandez RR, Pueyo JJ, Manrique E (2011) Spatial distribution and physiology of biological soil crusts from semi–arid central Spain are related to soil chemistry and shrub cover. Soil Biol Biochem 43:1894–1901 Palmqvist K, Sundberg B (2000) Light use efficiency of dry matter gain in five macro-lichens: relative impact of microclimate conditions and species-specific traits. Plant Cell Environ 23:1–14 Paoli L, Pisani T, Munzi S, Gaggi C, Loppi S (2010) Influence of sun irradiance and water availability on lichen photosynthetic pigments during a Mediterranean summer. Biologia 65:776–783 Pérez-Harguindeguy N, Díaz S, Garnier E, Lavorel S, Poorter H et al (2013) New handbook for standardised measurement of plant functional traits worldwide. Aust J Bot 61:167–234 Petchey OL, Gaston KJ (2002) Functional diversity (FD), species richness and community composition. Ecol Lett 5:402–411 Petchey OL, Gaston KJ (2006) Functional diversity: back to basics and looking forward. Ecol Lett 9:741–758 Pinho P, Bergamini A, Carvalho P, Branquinho C, Stofer S et al (2011a) Lichen functional groups as ecological indicators of the effects of low-intensity land-use in Mediterranean ecosystems. Ecol Indic 15:36–42 Pinho P, Dias T, Cruz CC, Tang YS, Sutton M et al (2011b) Using lichen functional-diversity to assess the effects of atmospheric ammonia in Mediterranean woodlands. J Appl Ecol 48:1107–1116 Prieto M, Aragón G, Martínez I (2010a) The genus Catapyrenium s. lat. (Verrucariaceae) in the Iberian Peninsula and the Balearic Islands. Lichenol 42:637–684 Prieto M, Martínez I, Aragón G (2010b) The genus Placidiopsis in the Iberian Peninsula and the Balearic Islands. Mycotaxon 114:463–472 R Development Core Team (2010) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna Rao CR (1982) Diversity and dissimilarity coefficients: a unified approach. Theor Popul Biol 21:24–43 Ricotta C, Moretti M (2011) CWM and Rao’s quadratic diversity: a unified framework for functional ecology. Oecologia 167:181–188 Spasojevic MJ, Suding KN (2012) Inferring community assembly mechanisms from functional diversity patterns: the importance of multiple assembly processes. J Ecol 100:652–661 Stofer S, Bergamini A, Aragón G, Carvalho P, Coppins BJ et al (2006) Species richness of lichen functional groups in relation to land use intensity. Lichenol 38:331–353 Stubbs WJ, Wilson JB (2004) Evidence for limiting similarity in a sand dune community. J Ecol 92:557–567 Villéger S, Mason NW, Mouillot D (2008) New multidimensional functional diversity indices for a multifaceted framework in functional ecology. Ecology 89:2290–2301 Violle C, Navas ML, Vile D, Kazakou E, Fortunel C et al (2007) Let the concept of trait be functional! Oikos 116:882–892 Warren SD (2003) Synopsis: influence of biological soil crusts on arid land hydrology and soil stability. In: Belnap J, Lange OL (eds) Biological soil crusts: structure, function, and management. Springer, Berlin, pp 349–360 Weiher E, Keddy P (2001) Ecological assembly rules: perspectives, advances, retreats. Cambridge University Press, Cambridge West NE (1990) Structure and function of microphytic soil crusts in wildland ecosystems of arid to semi-arid regions. In: Begon M (ed) Advances in ecological research. Academic Press, New York, pp 179–223 Zedda L, Grongroft A, Schultz M, Petersen A, Mills A, Rambold G (2011) Distribution patterns of soil lichens across the principal biomes of southern Africa. J Arid Environ 75:215–220