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Long-Term Changes in Biological Soil Crust Cover and Composition Eva Dettweiler-Robinson1*, Jeanne M Ponzetti2 and Jonathan D Bakker3

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Long-Term Changes in Biological Soil Crust Cover and Composition Eva Dettweiler-Robinson1*, Jeanne M Ponzetti2 and Jonathan D Bakker3 Dettweiler-Robinson et al. Ecological Processes 2013, 2:5 http://www.ecologicalprocesses.com/content/2/1/5 RESEARCH Open Access Long-term changes in biological soil crust cover and composition Eva Dettweiler-Robinson1*, Jeanne M Ponzetti2 and Jonathan D Bakker3 Abstract Introduction: Communities change over time due to disturbances, variations in climate, and species invasions. Biological soil crust communities are important because they contribute to erosion control and nutrient cycling. Crust types may respond differently to changes in environmental conditions: single-celled organisms and bryophytes quickly recover after a disturbance, while lichens are slow growing and dominate favorable sites. Community change in crusts has seldom been assessed using repeated measures. For this study, we hypothesized that changes in crust composition were related to disturbance, topographic position, and invasive vegetation. Methods: We monitored permanent plots in the Columbia Basin in 1999 and 2010 and compared changes in crust composition, cover, richness, and turnover with predictor variables of herbivore exclosure, elevation, heat load index, time since fire, presence of an invasive grass, and change in cover of the invasive grass. Results: Bryophytes were cosmopolitan with high cover. Dominant lichens did not change dramatically. Indicator taxa differed by monitoring year. Bryophyte and total crust cover declined, and there was lower turnover outside of herbivore exclosures. Lichen cover did not change significantly. Plots that burned recently had high turnover. Increase in taxon richness was correlated with presence of an invasive grass in 1999. Change in cover of the invasive grass was positively related to proportional loss and negatively related to gain. Conclusions: Composition and turnover metrics differed significantly over 11 years, though cover was more stable between years. This study can be a baseline for assessing change in crust composition due to anthropogenic influences. Keywords: Biological soil crust, Columbia basin, Composition change, Taxa turnover rate, Fire, Bromus tectorum, Livestock exclosure Introduction perform many roles in the ecosystem; the presence of ma- Biological communities change over time due to factors ture crust is associated with reduced wind and water ero- such as disturbance (Pickett et al. 2008; Platt and Connell sion and increased carbon and nitrogen fixation (Belnap 2003), changes in climate (Tylianakis et al. 2008), and spe- and Lange 2001). The structure and composition of crust cies invasion (Mack et al. 2000). Understanding back- communities are related to precipitation levels, topo- ground rates of change is necessary when evaluating graphic position, soil type, and disturbance history (Belnap human impacts (Magurran et al. 2010). Arid lands are im- and Lange 2001; Ponzetti et al. 2007). portant ecosystems because they cover ~ 40% of Earth’s Changes in crust composition over time depend on site land surface and are rapidly changing due to biophysical and vegetation characteristics, disturbance history, and and social factors such as climate, cropland expansion, and characteristics of the crust taxa (Rosentreter et al. 2001). overgrazing (Geist and Lambin 2004). Sites with higher resource availability, such as moist Biological soil crusts are communities of photosynthetic microsites, can support more species (Garcia-Pichel and organisms that live on the soil surface of arid lands. Crusts Belnap 2001) and can have higher productivity and more stability (Hooper et al. 2005). Invasive species such as the * Correspondence: [email protected] annual grass Bromus tectorum can decrease crust cover 1Department of Biology, University of New Mexico, MSC03 2020, 1 University of New Mexico, Albuquerque, NM 98131, U.S.A and species richness (Belnap et al. 2006). This grass may Full list of author information is available at the end of the article © 2013 Dettweiler-Robinson et al.; licensee Springer. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/2.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Dettweiler-Robinson et al. Ecological Processes 2013, 2:5 Page 2 of 10 http://www.ecologicalprocesses.com/content/2/1/5 prevent crust from recovering (Dettweiler-Robinson et al. et al. 2004; Kaltenecker et al. 1999). In addition, these 2013) by competing for resources and/or reducing the fire studies used a space-for-time approach, tended to focus return interval (Brooks et al. 2004). Disturbances such as on total crust cover rather than individual taxa, and oc- fire, intense grazing, or drought can directly remove or curred in sites dominated by cyanobacterial crust. In this damage crust. Disturbances can also indirectly affect crust study, we monitored permanent plots (Ponzetti et al. by triggering abiotic changes, such as loss of soil stability 2007) to document changes in a bryophyte- and lichen- (Bowker et al. 2004) or invasion by non-native plants dominated crust community 11 years apart. We assessed (Chambers et al. 2007). The intensity, timing, recurrence, changes in crust taxa by comparing cover, frequency of and type of disturbance can affect the rate of crust recov- occurrence, and indicator species between years. We ery even up to 30 years post disturbance (Bowker et al. quantified changes in community composition, total 2004; Ford and Johnson 2006; Lalley and Viles 2008; cover, covers of bryophytes and of lichens, and taxon Marble and Harper 1989; Zaady et al. 2007). For example, richness, as well as proportional taxon loss, gain, and the first fire that burns through intact shrub cover follow- turnover. We examined the relationship between these ing a long fire-free period would be of high intensity and variables and potential abiotic and biotic explanatory therefore should have larger effects on the crust than sub- factors including exclosure from large herbivores, eleva- sequent fires that are less intense because of the reduced tion, heat load index (HLI), time since fire (TSF), initial fuel loads. Crust taxa differ in susceptibility to damage and presence of B. tectorum, and change in B. tectorum removal by different disturbances (Ponzetti et al. 2007). cover. We expected (1) more change in cover and com- For example, organisms that live under plant canopies position in recently burned quadrats because communi- would be less likely to survive a high-severity fire than ties would be reassembling, (2) less change at higher organisms that live in plant interspaces (Hilty et al. 2004). elevation and cooler HLI as these areas have more mois- Succession may proceed differently based on timing and ture available to support high cover and diverse crust severity of disturbance, what portion of the community communities, and (3) more change in quadrats that ini- survived the disturbance, and the spatial arrangement tially contained B. tectorum or that had a large increase of disturbed and undisturbed patches (White and in B. tectorum cover. Jentsch 2001). Because some crust taxa are small, cryptic, and have Methods slow growth rates, community change can be difficult to Site description detect within short time scales. These changes reflect The Horse Heaven Hills (HHH) complex (46.2°N, differences among taxa in growth rate and in response 119.5°W) in south-central Washington is managed by to environmental factors. Cyanobacteria and other the Bureau of Land Management. The climate is semi- single-celled components can colonize a site and expand arid, with an average of 188 mm of precipitation per rapidly in cover (Belnap 1993). In a scraping disturbance year, mostly falling between October and April. Heavy experiment, cyanobacterial cover quickly recovered to fog is common in the winter; we have noted that crusts the level of the control treatment within 8 months and can be active (green, indicating photosynthetic activity) did not differ up to 32 months after treatment (Dojani on these foggy days. Mean daily temperatures are 0°C in et al. 2011). Bryophyte cover can increase rapidly in the January and 22°C in July (Western Region Climate Cen- absence of disturbance (Antos et al. 1983; Bates et al. ter 2012). Soil types include Ritzville silt loams and 2009), and young mosses can grow more rapidly than Kiona very stony silt loams (Soil Survey Staff and Nat- older mosses (Barker et al. 2005). Lichens generally have ural Resources Conservation Service 2011). Plots are lo- slow growth rates that may decline with age (Armstrong cated on elevations from 250–550 m with generally and Bradwell 2010). Crustose lichens can invade soon northerly aspects. after disturbance (Johansen et al. 1984) but may take a The dominant disturbances in the area are fire and graz- long time to increase in cover. Foliose and fruticose li- ing. Fires occurred in the area in 1986, 1998, 2002, and chens may colonize later in succession because they re- 2007; fire history before 1980 is uncertain due to lack of quire stable substrates (Belnap and Eldridge 2001). Dark spatially explicit data. Much of the area was grazed from cyanobacterial, moss, and lichen crusts responded more the 1960s to the 1980s. There have been no active grazing slowly and variably than light cyanobacteria in the scrap- leases since 1986, though there are scattered reports of ing experiment (Dojani et al. 2011), suggesting that longer trespass by cattle from nearby pastures. time scales
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