Biological soil crusts are highly functional soil surface communities of mosses, lichens and cyanobacteria that are vulnerable to disturbances such as grazing. We developed an eight-tier habitat classifcation based upon soil properties including texture, carbonate and gypsum content, and presence of shrinking-swelling clays.
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A Simple Classification of Soil Types as Habitats of Biological Soil Crusts on the Colorado Plateau, USA 2008 Journal of Vegetation Science
Biological soil crusts are highly functional soil surface communities of mosses, lichens and cyanobacteria that are vulnerable to disturbances such as grazing. We developed an eight-tier habitat classifcation based upon soil properties including texture, carbonate and gypsum content, and presence of shrinking-swelling clays.
Biological soil crusts are highly functional soil surface communities of mosses, lichens and cyanobacteria that are vulnerable to disturbances such as grazing. We developed an eight-tier habitat classifcation based upon soil properties including texture, carbonate and gypsum content, and presence of shrinking-swelling clays.
- A simple classifcation of biological soil crust habitat on the Colorado Plateau - 831
Journal of Vegetation Science 19: 831-840, 2008
doi: 10.3170/2008-8-18454, published online 15 April 2008 IAVS; Opulus Press Uppsala. A simple classifcation of soil types as habitats of biological soil crusts on the Colorado Plateau, USA Bowker, Matthew A. 1,2* & Belnap, Jayne 3 1 Department of Biological Sciences, Northern Arizona University, Box 5640, Flagstaff, AZ 86011,USA; Current Address: rea de Biodiversidad y Conservacin, Universidad Rey Juan Carlos, c/ Tulipn s/n., E-28933 Mstoles (Madrid), Spain; 3 Southwest Biological Science Center, U.S. Geological Survey, 2290 S.W. Resource Boulevard, Moab, UT 84532, USA; E-mail: Jayne_Belnap@usgs.gov; * Corresponding author; E-mail Matthew.Bowker@urjc.es Abstract Question: Can a simple soil classifcation method, accessible to non-experts, be used to infer properties of the biological soil crust (BSC) communities such as species richness, evenness, and structure? Location: Grand Staircase-Escalante National Monument, an arid region of the Colorado Plateau, USA. Methods: Biological soil crusts are highly functional soil surface communities of mosses, lichens and cyanobacteria that are vulnerable to soil surface disturbances such as grazing. We sampled BSC communities at 114 relatively undisturbed sites. We developed an eight-tier BSC habitat classifcation based upon soil properties including texture, carbonate and gypsum content, and presence of shrinking-swelling clays. We used simple structural equation models to determine how well this classifcation system predicted the evenness, richness, and community structure of BSC relative to elevation and annual precipitation. Results: We found that our habitat classifcation system explained at least 3.5 more variance in BSC richness (R 2
= 0.57), evenness (R 2 = 0.59), and community structure (R 2
= 0.34) than annual precipitation and elevation combined. Gypsiferous soils, non-calcareous sandy soils, and limestone- derived soils were all very high in both species richness and evenness. Additionally, we found that gypsiferous soils were the most biologically unique group, harboring eight strong to excellent indicator species. Conclusions: Community properties of BSCs are overwhelm- ingly infuenced by edaphic factors. These factors can be summarized effciently by land managers and laypeople using a simple soil habitat classifcation, which will facilitate incor- poration of BSCs into assessment and monitoring protocols and help prioritize conservation or restoration efforts. Keywords: Biodiversity conservation; Cryptogam; Ecological indicator; Ecosystem engineer; Land management; Microbiotic crust; Soil chemistry. Nomenclature: Zander 2008 (Bryophytes); Esslinger 2008 (Lichen). Abbreviation: BSC = Biological soil crust. Introduction Biological soil crusts (BSCs) are a type of thin (< 1 cm), desiccation tolerant microbial mat of cyanobacteria, subsequently colonized by mosses and lichens, living at the soil surface in drylands. These BSCs act as ecosystem engineers (Jones et al. 1997) and contribute strongly to dryland ecosystem function (Belnap 2002; Beymer & Klopatek 1991). For example, the flamentous cyanobac- teria of BSCs chemically and physically aggregate the soil surface into a thin horizontal layer, increasing erosion resistance and infuencing water redistribution (Mazor et al. 1996). In arid lands, vascular plants are known to create fertility islands, but BSCs are a dominant infuence in the creation and maintenance of soil fertil- ity in interspaces between vascular plants. Mechanisms through which BSCs increase fertility include carbon (C)-fxation (Beymer & Klopatek 1991), nitrogen (N)- fxation (Belnap 2002), and dust trapping (Reynolds et al. 2000), among others. Biological soil crusts also have potential to be highly valuable as ecological indicators of ecosystem health (Tongway & Hindley 1996). For these reasons and others, scientists should enable the inclusion of these information-rich ecological indicators in the management of public lands and other commons, in addition to private lands. Many previous studies have related environmental factors to the community composition and structure of BSCs at both small (< 1 m; Bowker et al. 2006a; Rosentreter 1986; Martinez et al. 2006) and large scales (100s of km 2 ; Ponzetti & McCune 2001; Bowker et al. 2005; Thompson et al. 2006). Often, a confusing array of inter-correlated factors also correlate with specifc components of the BSC community (Eldridge & Tozer 1997). Not surprisingly, BSC abundance, composition, and distribution are infuenced by climate variables such as annual precipitation, maximal temperature, and proximity to maritime air (Rogers 1972a; Nash & Mo- ser 1982). Another strong infuence is the physical and chemical environment of the soil (Downing & Selkirk 1993; Ponzetti & McCune 2001; Bowker et al. 2005). 832 Bowker, M.A. & Belnap, J Soil chemical and physical properties are especially diffcult to consider independently, for example pH, CaCO 3 , electrical conductivity, and available fractions of immobile nutrients (P, Mn, Zn, Cu, Fe) are usually strongly autocorrelated. Surface disturbances, such as livestock grazing, are also highly infuential in reducing BSC abundance (Brotherson et al. 1983). Potentially, a much simpler means of describing the soil habitat would be a desirable step in both descriptive and predictive modeling of BSC distribution and foristic patterns. Biological soil crusts are easily damaged or destroyed by compressional forces associated with several distur- bance types, including off road vehicles, foot traffc of humans and other animals, and mining and associated exploration activities (reviewed in Belnap & Eldridge 2003). Of these, livestock grazing is not the most severe (Belnap & Eldridge 2003), but is the most widespread of disturbance factors affecting the large majority of the western USA, and most arid rangelands of the world (Dregne & Chou 1992). Because BSCs are also notoriously slow to recover from disturbance, usually requiring decades (Belnap & Eldridge 2003), BSCs are rarely observed at their potential abundance, diversity, and functional state. Land managers increasingly require methods for inferring information about potential BSC communities to incorporate into assessment and moni- toring protocols, and also to prioritize conservation or restoration efforts after overgrazing and other soil surface disturbances. An easy to use scheme for inferring com- munity properties of BSCs is also potentially useful in traditional conservation applications (Goldstein 1999), such as identifying regions of high biological uniqueness or diversity (Myers et al. 2000). Thus, we developed a simple scheme of classifying soils with similar chemistry, texture, and sometimes appearance that was informative in predicting overall abundance of broad functional groups of BSCs (Table 1, Bowker et al. 2006b). The ability of this soil classifcation scheme to characterize BSC community composition and structure has not been studied previously. Our classifca- tion system does not require special expertise in either BSC taxonomy or ecology, or in soil science, but it does allow the user to infer consider information about the form and function of the BSC community. This classifcation is based solely on soil surface characteristics and can be derived from widely available information such as geo- logic maps or soil surveys, and characteristics that can be measured in the feld (Schoeneberger et al. 1998). Our classifcation system included eight soil types that include the majority of the soil types found on the Colorado Plateau (Table 1). This soil classifcation explained 17-73% of the variance in 19 specifc physico-chemical soil descriptors. We hypothesized that our soil classifcation system would also perform well in predicting BSC community charac- teristics, and outperform elevation and annual precipita- tion as predictors. The former was expected because: (1) soil chemistry and texture (calcium carbonate, gypsum, and clay and silt content) infuence inherent soil stability and nutrient availability, (2) shrinking and swelling clays create unstable substrates for slow growing soil surface organisms, and (3) increased pore space (determined by texture) provides an easier substrate for flamentous growth forms (e.g. cyanobacterial flaments). In contrast, precipitation ultimately determines activity time of these desiccation tolerant organisms, and elevation is a surrogate for temperature which strongly infuences photosynthetic rates of BSC organisms (Lange et al. 1998). Material and Methods Sampling design and survey methods We conducted sampling over three years in the ca. 800 000-ha Grand Staircase-Escalante National Monu- ment (Utah, USA). Our sampling design and survey meth- ods were designed to capture a broad array of ecological gradients in our sampling strategy and are described in detail in Bowker et al. (2006b). We divided the study area into three precipitation brackets ( 20 cm.a 1 , 20 - 30 cm.a 1 , 30 cm.a 1 ). All sites were classifed as one of eight mutually exclusive soil types (Table 1): bentonitic clay soils, calcareous sandy soils, non-calcareous sandy soils, gypsiferous soils, siliceous sandy soils (these are also non-calcareous, but are distinguished by large grain size, and siliceous cementing in parent materials), non- bentonitic fne soils, Kaiparowits-derived soils (a sandy textured parent material that is unique because it forms highly erodible badlands), and limestone-derived soils. Soils were assigned to these soil types using information in the GSENM Soil Survey (US Department of Agriculture- Natural Resource Conservation Service [USDA-NRCS] 2005). All possible combinations (some did not exist in GSENM) of soil type precipitation bracket were sampled and replicated (average n = 6, range 2 - 18) for a total of 114 sites. Only sites with relatively minor soil surface disturbance impacts were chosen for sampling, thus we did not have the luxury of a balanced random design. Rarity on the landscape accounted for low replication in a few soil type precipitation combinations, but common combina- tions were well replicated. At the completion of sampling, we had 6 - 37 replicate sites of each soil type, and 19 - 49 replicate samples for each precipitation bracket. To measure crust cover and composition, we used a step point-intercept transect (modifed from Evans & Love 1957) consisting of 300 points (spaced ca. 2 m apart) at each of the 114 sampling sites. At each point, ground cover measures were recorded, including mosses - A simple classifcation of biological soil crust habitat on the Colorado Plateau - 833 Table 1. A dichotomous key and descriptions of soil types. 1a. Soils primarily consist of clay- and silt-sized particles. Clay content is almost always greater than 30%. Surfaces generally exhibit a roughened popcorn- like texture due moderate to high shrink-swell capacity ( 4.5%).Bentonitic fne soils 1b. Soils are either more sand-dominated, or in some cases roughly equal proportions of sand, and clay size fractions. Clay content is almost always less than 30%. Soil surfaces may be roughened due to frost-heaving of biological soil crusts, but shrink-swell capacity is generally low2 2a. Soils contain at least 10% gypsum. Soil texture generally not sand-dominated, more often loamy. Soils often have visible exposures of white-gray gypsum beds.Gypsiferous soils. 2b. Soils contain less than 10% gypsum. Soil texture may be loamy, but is more often sand-dominated. Exposures of gypsum are lack- ing..3 3a. Soils are non-calcareous. There is little or no effervescence of HCL, [0, 0-3, or occasionally 1-5% calcium carbonate (CaCO 3 ) equivalent in USDA soil survey]. Textures are clearly sand-dominated4 4a. Soils are highly sand-dominated (> 75% sand sized particles). May be weathered in place from siliceously-cemented bedrock (e.g. Navajo Sandstone) or eolian deposits of similar material Siliceous sandy soils 4b. Soils are sand-dominated but contain < 75% sand-sized particles. May be weathered in place (not from Navajo Sandstone), or on well-weathered alluvial or fuvial terraces, or derived from parent material poor in CaCO 3 Non-calcareous soils 3b. Soils are calcareous. There is at least moderate effervescence of HCL at soil surface, usually > 5 maximum CaCO 3 equivalent. Textures are variable5 5a. Soils effervesce strongly, generally containing maximal CaCO 3 equivalent > 15%. Derived from either Kaiparowits Formation (Fm.), Timpoweap Member (Mbr.) of Moenkopi Fm., Kaibab Fm., or non-gypsiferous, non-limestone members of the Carmel and Moenkopi Fms. ..6 6a. Soils are derived from the Kaiparowits Fm. and commonly occur in highly eroded, gray-colored badlands, soil has sandy salt-and-pepper appearance Kaiparowits-derived soils 6b. Soils are often light colored (e.g. gray, yellowish-gray, pale orange), and are derived from limestone (Timpoweap Mbr. of Moenkopi Fm. or Kaibab Fm. in GSENM)..Limestone-derived soils 6c. Soils are deep orange or red grading into vermillion, and are derived from non- gypsiferous, non-limestone members of the Moenkopi Fm. and Carmel Fm., often channery at surface Non-bentonitic fne soils 5b. Soils are usually less effervescent, and less calcareous, but defnitely effervescent. Soils are not derived from above listed formations, and do not generally occur in badlands. Most of these soils are sand-dominated, but include some loamy textured soils as well.Calcareous sandy soils Bentonitic fne soils (B): Weathered in place from Tropic Fm., Blue Gate & Tununk Mbrs. of Mancos Fm., Brushy Basin Mbr. of Morrison Fm., and various shale members of the Chinle Fm.; Colors variable but often impressive, may include grays, reds, mauve, and green; Forming badland landscapes supporting very sparse vegetation, including the dwarf Atriplex corrugata; highly effervescent due to CaCO 3 and other carbonates; may be somewhat gypsif- erous, but not containing whitish gypsum beds; clay sized particles predominate and smectitic clay minerals generate high shrink-swell capacity; Generally lacking in BSC cover. Gypsiferous soils (G): Weathered in place from Shnabkaib Mbr. of Moenkopi Fm., Crystal Creek and Paria River Mbrs. of Carmel Fm., and Paradox Fm.; Gypsum-rich patches generally whitish or grayish, surrounding soil surface is reddish in many cases; Vegetation variable (Artemisia, Atriplex, Ephedra and Pinus-Juniperus); Containing CaCO 3 , but gypsum always more abundant; Contain beds or pockets of gypsum, including alabaster cliffs in some cases; Gypsum-rich microsites dominated by silt and clay sized particles, surrounding matrix may be more sandy; Potentially supporting high cover of moss and lichen rich BSCs. Siliceous sandy soils (S): Weathered in place or wind transported from non-calcareous siliceously-cemented sandstones (e.g. Navajo Sandstone and Coconino Sandstone); Color may be nearly white to pink to deep orange; Vegetation variable, depth-dependent (includes desert grasslands and Artemisia communities); CaCO 3 and gypsum lacking; Texture strongly skewed toward coarse sand; Potentially supporting high cover of cyanobacterial soil crusts. Non-calcareous sandy soils (NC ): Derived from various parent materials (usually weakly calcareous, including Judd Hollow Mbr. of Straight Cliffs Fm., or Shinarump Mbr. of Chinle Fm. but exclusive of siliceous sandstones), but surface weathering has been strong enough to leach out CaCO 3 ; Colors vary, dependent on parent material; Vegetation various, commonly Artemisia; Most abundant in relatively high precipitation areas due to greater leaching; Contain- ing little or no CaCO 3 or gypsum; Dominated by sand sized particles (often poorly sorted), but generally less coarse than Siliceous sandy soils; Potentially supporting moderate to high cover of cyanobacterially dominated BSCs, with a rich moss and lichen component. Kaiparowits-derived soils (K): Weathered in place from the Kaiparowits Fm. Occur in gray colored badlands, with variable climate-dependent vegetation; Violently effervescent due to carbonates, lacking gypsum; Texture is sand-dominated, with distinctive salt-and-pepper appearance. Generally lacking BSC cover. Limestone-derived soils (L): Weathered in place from any limestone (e.g. Kaibab Fm. or the Timpoweap Mbr. of Moenkopi Fm.; Colors range from grays to pale yellow or orange coloration (dependent on parent material); Vegetation climate-dependent, usually Artemisia or Pinus-Juniperus dominated; Moderately effervescent due to presence of CaCO 3 , gypsum not abundant; Texture generally loamy, but dependent on parent material; Potentially supporting high cover of moss and lichen rich BSCs. Non-bentonitic fne soils (NB): Weathered in place from non-limestone, non-gypsiferous, and non-bentonitic fuvial shale red beds (e.g. Mbrs. of Moenkopi Fm., Carmel Fm., Halgaito Fm., Organ Rock Fm., Hermit Fm.); Notably deep orange, red or purple colored; Vegetation often sparse, including Coleogyne, Artemisia, and Pinus-Juniperus; Moderately effervescent due to presence of CaCO 3 , gypsum not abundant; Texture usually loamy, but sand fraction is fne; conspicuous fat channers often on surface, exposed soil forms a vesicular physical crust; Potentially supporting very low to moderate cover of moss and lichen BSCs, dependent on moisture. Calcareous sandy soils (C): Derived at least partially from any calcareous sandstone, exclusive of the Kaiparowits Fm. (e.g. Kayenta Fm., Entrada Sandstone, Dakota Fm. among several others), weathering has not been suffcient to remove all CaCO 3 from soil surface; Colors variable, dependent on specifc parent material; Vegetation highly variable, climate dependent; Usually moderately effervescent, gypsum content minor; Texture generally skewed toward more sandy size classes, although some soils are loamy; Potentially supporting low to moderate cyanobacterial BSC cover, with some mosses and lichens. 834 Bowker, M.A. & Belnap, J and lichens to species, light cyanobacterial BSCs (early successional BSCs, almost exclusively Microcoleus spp.), and dark cyanobacterial BSCs (later succesional BSCs dominated by Microcoleus spp., Nostoc spp. and Scytonema spp.). Statistical analysis To determine the relative contribution of three environmental predictors linked to soil stability and fertility, and physiological activity of BSC organisms (soil type, annual mean precipitation [modeled within a 5.08 cm resolution], elevation [values obtained us- ing GPS]) to BSC richness, evenness (Pielous J), and community structure, we used three simple structural equation models (SEM) in Amos 5.0 (SPSS Inc., Anon. 2003). These models were essentially equivalent to multiple regression models except that they allowed the inclusion of the categorical predictor soil type as a composite variable (Grace 2006), allowed the inclusion of correlated predictors by specifying their relationship to one another (e.g. our models specifcally state that annual precipitation is infuenced by elevation), and used a maximum likelihood technique to estimate regression weights simultaneously. Often, researchers using SEM advance a causal hypothesis and test the goodness of ft of their model structure. In this case, the causal order- ing of the variables in the models is known with high confdence; thus, a test of ft was not of interest. Instead, because we were interested only in maximum likelihood estimates of regression weights and R 2 , we allowed all of the predictors to freely covary, creating a saturated model. Saturation precludes an overall goodness of ft test, which is not of consequence here. Community structure was a synthetic response variable derived from a non-metric multidimensional scaling (NMDS) ordina- tion. We used the slow and thorough autopilot setting in PC-ORD 4.0 (MJM Software Design 1999), which selects optimal dimensionality of the ordination using a Monte Carlo test. In this and subsequent NMDS ordina- tions, we used the Bray-Curtis distance measure because it is compatible with the distribution problems common to community data. Data were relativized to species maximum (i.e. data for each species was rescaled from 0-1). This transformation suppresses the effect of a few dominant taxa upon the analysis, and gives equal weight to all community members; this is valuable in broadening inference from the species to the community level. The ordination was one-dimensional, thus the axis scores were used as a univariate variable in our SEM. To determine how specifc soil types differed in species richness and evenness, we used two ANOVA models in JMP IN 5.0 (SAS Inst. 2005). To determine specifcally how community structure changed among soil types, we used indicator species analysis (Dufrene & Legendre 1997) in PC-ORD (MJM Software Design 1999). Indicator species analysis uses abundance and frequency data to yield a percent of perfect indication value for each variable-group combination and uses a Monte Carlo test to determine the probability of obtain- ing a given indicator value (IV) due to chance alone. We defned a strong indicator of a particular soil type as one with an IV of 0.25-0.50, a very strong indicator as one with an IV of 0.51-0.75, and an excellent indicator as one with an IV 0.76. Fig. 1. Three structural equation models of identical structure using three predictors (soil type, elevation, and annual pre- cipitation) as determinants of three responses: BSC species richness, evenness, and community structure. Rectangles represent measured variables. The composite variable (Grace 2006) soil type is represented by a diamond. Unidirectional arrows represent a directed causal infuence of one variable upon another. Bidirectional arrows represent undefned covariance between a pair of variables. Path coeffcients, appearing above unidirectional arrows, are equivalent to regression weights or partial correlation coeffcients (width of arrows is proportional to path coeffcients). R 2 appears above every endogenous vari- able and is interpreted as for any linear model. In all three cases, soil type is the best predictor of BSC parameters. a correlations of elevation with soil types: B = 0.10, G = 0.26, L = 0.35, NB = 0.06, NC = 0.17, K = 0.04, S = 0.02. b correlations of pre- cipitation with soil types: B = 0.24, G = 0.11, L = 0.23, NB = 0.12, NC = 0.22, K = 0.04, S = 0.03. **** P < 0.0001 - A simple classifcation of biological soil crust habitat on the Colorado Plateau - 835 Results BSC community properties as a function of environ- mental predictors Using simple structural equation models with soil type, elevation, and precipitation as predictors, we were able to explain more than half of the variance in BSC species richness (R 2 = 0.57), about a third of the variance in community structure (R 2 = 0.34), and over half in BSC evenness (R 2 = 0.59) (Fig. 1). In the case of richness and community structure, soil type was much more informative than the other variables, although they did make contributions. We found that in the case of evenness, soil type (r = 0.77) accounted for virtually all of the variance explained (Fig. 1). BSC community properties as a function of soil type Both species richness (F = 17.7429, P < 0.0001) and evenness (F = 68.1449, P < 0.0001) were strongly infuenced by soil type (Fig. 2). Richness was about six times higher in gypsiferous soils compared to bentonitic fne soils and Kaiparowits-derived soils; the other soil types were intermediate. Evenness exhibited a similar pattern except that limestone-derived soils were the most even and were similar to gypsiferous soils; again bentonitic fne soils and Kaiparowits-derived soils were much lower (about fve-fold) and other soil types were intermediate. Because BSC community structure was also largely a function of soil type, we conducted an indicator spe- cies analysis. A total of 19 out of 59 species and other taxonomic groups were strong to excellent indicators (defned as IV > 25) of individual soil types (Table 1). Bentonitic fne soils, calcareous sandy soils, Kaiparowits- derived soils and non-bentonitic fne soils had no strong indicator species. Gypsiferous soils had eight strong to excellent indicators; seven of these were very strong, (Lecanora gypsicola, Psora decipiens, Didymodon nevadensis, Diploschistes diacapsis, Squamarina len- tigera, Acarospora nodulosa ssp. nodulosa), and two were excellent indicators (Catapyrenium spec., Ful- gensia bracteata) primarily due to their fdelity to this soil type. Limestone-derived soils were characterized by fve strong indicators, of which Aspicilia aspera and Psora cerebriformis were very strong. Siliceous sandy soils and non-calcareous sandy soils each had two strong indicators. However they were not as strong as those for the gypsiferous soils or limestone-derived soils due to lack of fdelity in the case of siliceous sandy soils, and a lack of consistency in the case of non-calcareous sandy soils. Discussion These data suggest that soil characters are by far the most infuential natural abiotic predictor of the richness, evenness, and community structure of BSCs of the Colorado Plateau. A simple system of eight soil types provides a relatively non-technical means of sum- marizing BSC community properties. This information is potentially applicable in inferring reference states in range management (Bowker et al. 2006b) and ecological restoration (Bowker 2007), conservation of BSC function (Bowker et al. in press), and reserve design for conserv- ing biodiversity of BSCs. A useful framework for classifcation of BSC habitat on the Colorado Plateau The majority of soils of the Colorado Plateau are characterized primarily by residuum weathered in place from sedimentary rocks. Of the fve factors of soil for- mation (Jenny 1941), parent material (i.e. geology) has Fig. 2. Species richness and evenness as a function of soil type. Error bars represent one standard error. B = bentonitic fne soils, K = Kaiparowits-derived soils, C = calcareous sandy soils, S = siliceous sandy soils, NB = non-bentontitic fne soils, NC = non calcareous sandy soils, G = gypsiferous soils, L = limestone-derived soils. Shared letters indicate Tukey-Kramer test, P > 0.05. 836 Bowker, M.A. & Belnap, J Table 2. Results of indicator species analysis, by community member or community type. Soil type refers to the soil type for which a community member has the highest indicator value IV. B = bentonitic fne soils, C = calcareous sandy soils, G = gypsiferous soils, K = Kaiparowits-derived soils, NB = non-bentontitic fne soils, NC = non calcareous sandy soils, S = siliceous sandy soils. Bold I.V. and associated P-values indicate species that are strong to excellent indicators. Species Taxonomic Group Soil type I.V. P light cyanobacterial crust Cyanobacterial community C 15.9 0.11 dark cyanobacterial crust Cyanobacterial community S 33.8 0.0002 Aloina bifrons Moss G 20.3 0.03 Aspicilia hispida Lichen L 73.7 0.0002 Aspicilia aspera Lichen L 14.5 0.09 Aspicilia desertorum Lichen NC 34.7 0.0028 Aspicilia or Lecanora spec. Lichen S 12.7 0.19 Acarospora nodulosa Lichen G 58 0.0002 Acarospora schleicheri Lichen NC 25 0.010 Bryum argenteum and lanatum Moss S 32.3 0.003 Bryum kunzei Moss G 7.4 0.57 Bryum caespiticium Moss L 16.2 0.09 Crossideum spec. Moss G 18 0.03 Collema coccophorum Lichen L 41.1 0.0004 Catapyrenium spec. Lichen G 90 0.0002 Placidium spp. Lichen G 22.6 0.04 Ceratodon purpureus Moss NC 24.6 0.03 Cladonia pyxidata Lichen S 19.2 0.05 Collema tenax Lichen G 20.9 0.14 Candellariella terrigena Lichen NC 26.9 0.05 Calopolaca tominii Lichen G 23.6 0.03 Caloplaca jungermanii Lichen L 18.6 0.02 Caloplaca lactea Lichen L 20 0.01 Diploschistes muscorum Lichen NC 19 0.03 Diploschistes diacapsis Lichen G 64.3 0.0002 Didymodon nevadensis Moss G 66.7 0.0002 Desmatodon spec. Moss C 2.7 - Didymodon vinealis Moss C 2.7 - Endocarpon pusillum Lichen NB 20.1 0.06 Encalypta spp. Moss S 10.8 0.13 Fulgensia desertorum Lichen G 6.5 0.91 Fulgensia bracteata Lichen G 87.8 0.0002 Gypsoplaca macrophylla Lichen G 23.4 0.01 Grimmia orbicularis Moss G 33.3 0.0006 Grimmia anodon Moss L 10 0.23 Brachythecium collinum Moss C 2.7 - Heppia spec. Lichen NC 6.2 0.68 Lecanora gypsicola Lichen G 74.6 0.0002 Leptogium lichenoides Lichen B 3.4 0.81 Lecanora muralis Lichen NC 21.3 0.02 Lecanora cf. zosterae Lichen B 3.5 0.93 Nostoc cf. fagelliforme Cyanobacterium L 23.5 0.02 Rinodina spec. Lichen NC 6.2 0.67 Psora cerebriformis Lichen L 57.7 0.0002 Psora globifera Lichen L 6.6 0.48 Psora pruinosa Lichen NC 19.8 0.04 Psora decipiens Lichen G 67.8 0.0002 Pterygoneurum ovatum Moss L 24.4 0.01 Peltigera rufescens Lichen NC 12.5 0.11 Pterygoneurum subsessile Moss L 8.2 0.24 Psora tuckermanii Lichen L 26.9 0.0128 Peltula patellata Lichen G 14.3 0.24 Unknown pyrenolichen Lichen C 2.7 - Squamarina lentigera Lichen G 58.3 0.0002 Syntrichia caninervis Moss L 29.4 0.004 Syntrichia ruralis Moss S 24.1 0.06 Tortula brevipes Moss G 10.1 0.19 Tortula mucronifolia Moss K 16.7 0.05 Toninia sedifolia Lichen L 45.9 0.002 - A simple classifcation of biological soil crust habitat on the Colorado Plateau - 837 the dominant infuence because of the very different textures and chemistries of the various rock formations and due to the relative young age of many of the soils. This differs considerably from the Great Basin, Mojave, and Sonoran Deserts where older soils are composed primarily of transported material of various origins, and where climate, age of alluvial parent materials, and slope may play more important roles in soil development (Scull et al. 2004). In contrast, the complex and diverse mosaic of geological formations in the Colorado Plateau results in more stark differences and heterogeneity among soils, but also more predictable soil properties. Thus, considerable information can be inferred about BSCs and other biota from geological surveys. The system of eight soil functional types used here is based upon observations and formal surveys of hundreds of sites across the Colorado Plateau. Classifcation rules (Table 1) are based upon available information in soil surveys (e.g. soil physico-chemical properties and often parent material; USDA-NRCS, Anon. 2005), geological surveys (e.g. descriptions of parent materials; Doelling & Davis 1989), and feld-observable characteristics (Schoeneberger et al.1998). Bowker et al. (2006b) found that this system explained variance in 19 different soil properties, reducing the coeffcient of variation by 35% on average. It is broad enough that it can be applied to most soils of the Colorado Plateau and potentially to other regions. To truly broaden our system to cover the entire Colorado Plateau, additional soil types may need to be invoked. Basalt-derived soils, and granite-derived soils, are two potential additional groups that could be added to this classifcation system in the future. As with many groups of biota, BSCs appear to contain species that are generalists and specialists. At one extreme, some of the most common taxa occur in virtually all deserts of the world, e.g. Microcoleus vaginatus, Collema tenax or coccophorum, and Anomo- bryum argenteum (Belnap & Lange 2003). Other taxa have a high fdelity for particular soil chemical or textural characters, e.g. Aspicilia hispida or A. aspera and CaCO 3 , and Lecanora gypsicola and gypsum. Both calcium carbonate and gypsum, can reduce the avail- ability of immobile nutrients such as P, and free Ca+ ions can also displace exchangeable cations such as K+ and Mg+ in cells (Lajtha & Schlesinger 1988; Bates & Farmer 1990). Habitat specialization based upon calcium compounds in the soil may refect a disparity in mechanisms for nutrient uptake. Furthermore, in the case of gypsum specialization, sulfur may play an important role as three of our very strong gypsum indica- tors (Table 1; Diploschistes diacapsis, Psora decipiens, and Squamarina lentigera) are also known to inhabit the edges of sulfur-rich thermal springs (Rosentreter & Belnap 2003). Soil texture can potentially have many effects. High silt and clay content is likely to increase water holding capacity of the soil which could favor less drought-resistant BSC taxa (Anderson et al. 1982). Sandy soils also tend to be less aggregated and therefore less stable, so perhaps the fastest-growing BSC organisms (the cyanobacteria) are favored on the sandier substrates (Dougill & Thomas 2004). In addition, the greater pore spaces may present fewer physical barriers to flamentous cyanobacteria. Poten- tially, some combination of the latter two mechanisms may have led to establishment of well-developed dark cyanobacterial crusts on siliceous sandy soils (Table 1), the sandiest soils in our study region. No species favored bentonitic clay soils, in fact these substrates were nearly devoid of BSCs (also observed in Bowker et al. 2005). Although we cannot rule out a chemical mechanism to explain this, the simplest explanation is that shrinking and swelling clays and high susceptibil- ity to water erosion make these substrates very diffcult for BSCs to colonize. In total, over half of the species encountered exhibited at least some degree of habitat specialization, and over one third of these were quite strong (Table 1). Although the mechanisms behind these physico-chemical soil preferences of specialist species are not well understood, the phenomenon is highly predictable using the soil types outlined here. The specifc classifcation rules listed in Table 1 are designed for use across the 337 000-km 2 Colorado Plateau region of the USA, but could be modifed for use in other arid regions of the world. The key in accom- plishing this is identifcation of the important abiotic gradients which predict BSC community properties, and classifcation of soils based upon possible combina- tions of these properties. Examples of the same abiotic gradients having predictive power for BSC community properties in other regions include: soil texture Aus- tralia (Eldridge & Tozer 1997; Thompson et al. 2006), shrink-swell clays Italy (Loppi et al. 2004), gypsum content Tunisia and Australia (Ullman & Bdel 2003), Spain (Guerra et al. 1995), Namibia (Lalley & Viles 2005), and calcium carbonate content Tunisia and Australia (Ullman & Bdel 2003); Columbia Basin, USA (Ponzetti & McCune 2001). Fortunately many of these characteristics are measured in or can be inferred from geological surveys and soil surveys, and in many cases maps already exist which can be used in GIS applications. 838 Bowker, M.A. & Belnap, J An application: reserve design for conserving biodi- versity of BSCs We previously developed an approach to conserva- tion and restoration prioritization based upon BSC functional signifcance which uses the same predic- tors to determine where on the landscape BSCs may play disproportionate roles in ecosystem function, and overlays the information upon additional information describing probability of surface disturbance (Bowker et al. in press). Thus, we can identify the areas where BSCs have the greatest signifcance and are simultaneously most likely to be disturbed in the present or future; the former might be considered restoration priority areas, and the latter may be considered conservation priority areas. Additionally, our soil classifcation system has been used to model potential cover of key BSC types for use as a reference condition parameter in either range management or restoration-rehabilitation applications (Bowker et al. 2006). In contrast to these applications, when BSC biodiversity is the object of conservation, a reserve design incorporating key gradients and regional hotspots could be more useful. Of course, by acting to conserve both organisms and function we could most comprehensively protect these systems. Traditional approaches to conservation of various groups of taxa have often sought to determine which spatial locations or habitat types support the greatest biodiversity or biological uniqueness (Myers et al. 2000, Stohlgren et al. 2005). Based upon the present work, we can clearly say that conservation of BSC taxa would be most effective via reserves on three soil types: non- calcareous sandy soils, gypsiferous soils, and limestone- derived soils. These three soil types were the richest and also most even communities. Among the three groups, there were a total of 15 indicator species, indicating that these soil types contain many habitat specialists. Non-specialist species are also well represented in this soil type, indeed only a handful were never represented in either gypsiferous soils, non-calcareous sandy soils, and limestone derived soils. It appears that to a large degree, the BSC fora of other soil types are a subsets of the fora found on these three soil types. For at least three decades, gypsiferous soils have been known to harbor both a diverse and unique community of soil crusts in both North America and Europe (Anderson & Rushforth 1976; Guerra et al. 1995; Martinez et al. 2006). While fairly common in the Mediterranean regions of Europe (Martinez et al. 2006), gypsiferous soils are rare on the Colorado Plateau. Examples of this unique and poorly studied fora of Colorado Plateau gypsiferous soils include three globally rare species frst described in the 1990s (Didymodon nevadensis, Lecanora gypsicola), one more broadly distributed species that was undetected in North America until the 1980s (Acarospora nodulosa ssp. nodulosa), and one globally rare genus and species (Gypsoplaca macrophylla; St. Clair & Warrick 1987; Zander et al.1995; Rajvanshi et al. 1998). In contrast, most of the species of non-calcareous and limestone-derived soils are widespread, however the species turnover between these two habitat types is large. In addition to representing these regional-scale hotspots that are rich in habitat specialists and are biologically unique, it is important to capture important ecological gradients in reserve design in heterogenous areas (Diamond 1975). Climatic gradients are impor- tant, but as we have shown here, edaphic gradients are more important to the structure of BSC communities. Thus, reserve design could primarily be based upon edaphic gradients. Aside from gypsum content, perhaps the most important gradient to capture is CaCO 3 content. Calcareousness of substrates, either rock or soil, have long been known to infuence the potential lichen and moss communities (Downing & Selkirk 1993; Ponzetti & McCune 2000). Additionally, soil texture has long been recognized as an important determinant of BSC development (Anderson et al. 1982). Thompson et al. (2006) suggest that for BSCs, small reserves would be adequate because species-area curves saturate quickly. Although we did not study species-area relationships specifcally, our observations agree with this statement as long as the system of small reserves (each of perhaps several hectares) capture variance in the key gradients mentioned above. However, it is important to explicitly state that this assertion applies only to biodiversity conservation of BSC, and that small reserves cannot appreciably conserve BSC function in the larger ecosystem. Reserves are merely islands in a matrix of landscape being used for livestock production, a major negative infuence upon BSC development. Incorporation of BSCs into land management ap- plications such as reserve design and resource manage- ment plans (Bowker et al. 2006b; Bowker et al. 2007) requires the delivery of accessible tools to managers, who cannot be expected to also be BSC experts. With the application of the BSC habitat classifcation system described here, non-specialists can make informed deci- sions about where BSC biodiversity or function might be conserved, and where BSCs might be most useful as ecological indicators. - A simple classifcation of biological soil crust habitat on the Colorado Plateau - 839 Acknowledgements. 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