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Blackbrush (Coleogyne Ramosissima Torr.) 145 This file was created by scanning the printed publication. Errors identified by the software have been corrected; however, some errors may remain. BLACKBRUSH (COLEOGYNE RAMOS/SSJMATORR.) State of Our Knowledge and Future Challenges Rosemw)i L. Pendleton, Burton K. Pendleton, Susan E. Meye1; B1yce Richardson, Todd Esque) and Stanley G. Kitchen ABSTRACT its current distribution would be unlikely to Blackbrush (Co!eogyne ramosissimaTorr.) is succeed. Common garden experiments under a regionally dominant shrub species found in multiple climate conditions will give us a the transition zone between North American better understanding of climatic tolerances. warm and cold deserts where it occupies Further research is needed to develop new millions of hectares on National Park Service, and better restoration techniques. Bureau of Land Management, and National Forest lands. Blackbrush habitat is under INTRODUCTION severe threat of loss from the combined effects of exotic annual grasses, increased fire Blackbrush (Coleogyne ramosissima -Torr.) frequency, and rapid climate change. Attempts occurs in an ecotonal band between warm to restore blackbrush habitat have met with and cold desert regions extending from limited success. This research synthesizes California through Utah and Arizona (Meyer existing information on blackbrush biology and Pendleton 2005). It is an iconic landscape and ecology. Our goal is to guide successful dominant in many of the national parks restoration efforts. Recent genetic work has and monuments. In the Colorado Plateau, revealed the existence of two metapopulations blackbrush is found throughout much of corresponding to the Mojave and Colorado southeastern Utah primarily in upland areas Plateau regions. This work, along with ger­ along the Colorado and San Juan River mination and reciprocal transplant studies, drainages and into the desert canyon areas of demonstrates the need to use appropriate northern Arizona. Blackbrush communities seed sources. Blackbrush plants produce of the Colorado Plateau are bounded by salt large seed crops infrequently; however, seed deset1 shrub (Atriplex sp.), sand sagebrush can be stored for 10--12 _years with minimal (Artemisiafi!ifolia), and grass communities at loss of viability. Establishment success may lower elevations and by pinyon-juniper (Pinus be enhanced by planting seeds in the fall edulis-Juniperus sp.) and sagebrush-grass at to meet chilling requirements, and in small higher elevations. Blackbrush communities groups at 2.5--4.0 em in depth to mimic rodent also occur in the Mojave, where they are caches. Cunent observations and projected bounded by the creosote-bursage (Larrea changes in climate suggest that attempts to tridenlata-Ambrosia dumosa) community restore blackbrush at lower elevations than at lower elevations and by pinyon-juniper- Blackbrush (Co/eogyne ramosissima Torr.) 143 sagebrush at upper elevations (Lei and dissertation was subsequently published Walker 1995). Blackbrush is found in some as a research report (Bowns and West of the most scenic areas of the Southwest, 1976), which incJuded a distribution map and it provides cover and food for a number for blackbrush compiled from a number of wildlife species, most notably heteromyid of sources. More recently, Richardson and rodents (Auger 2005), nongame birds, deer, Meyer (20 12) developed a map of predicted and desert bighom (Bradley 1965; Brown and distribution based on bioclimatic profiles Smith 2000). (figure 10.1). Unlike the map ofBowns and The blackbrush community is currently West (1976), the newer map shows a distinct under threat from the combined effects of break between Colorado Plateau and Mojave exotic species, large-scale fires, and climate Desert populations. change. Predicted effects of climate change This break in distribution is reinforced on recruitment ofblackbrush make it unlikely by genetic analysis (Richardson and Meyer that blackbrush will persist in place. Black­ 20 12), which revealed the existence of two brush has little resistance to invasion by exotic metapopulations corresponding to Colorado species and very little resilience to subsequent Plateau and Mojave Desert regions. This same large-scale fires. Blackbrush itself is not fire study also refuted the long-held assumption adapted and does not resprout when burned that olackbrush was a paleoendemic species, (Brooks 2009). Large areas of the blackbrush generally lacking in genetic variation (Steb­ community have burned in the last decade, bins 1972; Stebbins and Major 1965). The although primarily in the Mojave. Natural assumption was based primarily on the fact recove1y from these and other large-scale that the genus is monotypic with few close disturbance events is apparently not possible, relatives (see Potter et al. 2007). Current dis­ and the resulting vegetation then becomes tribution was thought to represent a restriction dominated primarily by exotic weed species from a much wider distribution during the (Brooks 2009). Cretaceous period to xeric "islands" during In this chapter, we summarize the current the onset of more mesic Tertiary· conditions state of knowledge of blackbrush biology, during which the gene pool became depauper­ including reproductive biology, establish­ ate (Stebbins 1972). In contrast, studies have ment ecology, interaction with herbivores, now revealed a relatively high level of genetic response to disturbance, and genetic stmc­ diversity within blackbrush populations, as ture. We report on current studies of inter­ well as significant population differentiation population variation and potential response (Richardson and Meyer 2012; Schuster to climate change. Finally, we assess current et al. 1994). The existence of two metapop­ efforts at restoration and identify needs for ulations also provides context for prior work additional research. demonstrating that the two population groups differ in other important ways related to their germination characte1istics and establishment DISTRIBUTION AND GENETIC strategies (Meyer and Pendleton 2005; Pend­ STRUCTURE leton and Meyer 2004 ). Prior to the early 1970s, very little was known about blackbrush, despite its occurrence as SOILS AND SOIL MICROORGANISMS a landscape dominant on over three million hectares (Pendleton and Meyer 2004). The Blackbrush stands typically occur on upland first intensive study of blackbrush biology terraces, ridges, open plains, and alluvial was done by Jim Bowns as part of his PhD slopes (Bowns 1973; Bowns and West 1976; research (Bowns 1973). Most of Bowns' Turner 1994). Edaphic conditions largely 144 Pendleton et al. Figure 10. 1. Predicted distribution of blackbrush based on a bioclimatic profile from Richardson and Meyer (2012). control the amount of blackbrush present Romney 1972) and does not occur on soils within a vegetative community (Thatcher with high salt content (Bowns and West 1975). Soils are generally described as 1976; Shreve 1942). A more comprehensive coarse, shallow, poorly developed, and description of blackbrush soil relationships calcareous (Anderson 200 l; Kearney and across the geographic range is sorely needed, Peebles 1960; Shreve 1942). However, more as available soils data are somewhat limited. specific descriptions are needed. Parent In the Colorado Plateau, blackbmsh sites materials in the Colorado Plateau frequently characteristically exhibit well-developed consist of sandstone or siltstone, whereas biotic crusts, which help to stabilize sandy mixed alluvium and basalts are more com­ soils in the relatively large perennial plant mon on Mojave sites, and often they have a canopy interspaces (Belnap and Lange pronounced petrocalcic layer (Bowns 1973). 2003; Munson, Belnap, and Okin 2011). Blackbrush is saline sensitive (Wallace and Soil crusts in the Mojave are less developed Blackbrush (Coleogyne ramosissima Torr.) 145 Figure 10.2. Photograph of a pedestalled blackbrush plant showing the high degree of soil erosion that often follows destruction of biological soil crusts on the Colorado Plateau. and are dominated by cyanobacteria with mycorrhizal and dark septate fungal associa­ occasional patches of lichen and moss. Soil tions (Green, Porras-Alfaro and Sinsabaugh organic matter, nitrogen, and phosphorus are 2008; Hawkes 2003). This mechanism may reportedly higher under blackbrush plants well be operating in the well-defined soil than in interspaces, which is consistent with crusts of the Colorado Plateau and, to a more the fertile island effect common to shrub lands limited extent, in the Mojave. In greenhouse (Bowns and West 1976; Thompson et al. studies, blackbrush seedlings exhibited a 2005). Wallace and Romney (1972) attributed positive growth response to the presence of the increase in nitrogen to symbiotic nitro­ arbuscular mycorrhizae and biological soil gen fixation within the shrub rhizosphere. crusts (Pendleton, Pendleton, and Howard However, biological soil crusts are the 1999). Large-scale hot fires can disrupt these predominant source of nitrogen in many arid soil relationships, and loss of soil crust from ecosystems (Evans and Ehleringer 1993), and any kind of disturbance can lead to significant nitrogen fixed by soil crust cyanobacteria erosion (figure 10.2; Belnap and Lange 2003; can be translocated to plant roots through Munson et al. 2011 ). 146 Pendleton et al. REPRODUCTIVE BIOLOGY production (Pendleton 2008; Pendleton, Meyer, and Pendleton 1995). Flowering in blackbrush typically occurs Blackbrush fruits ripen between late May during April and early May and
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