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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 is induced and the third week of July depending on by moderate to heavy winter precipitation. elevation and weather conditions (Pendleton The timing and degree of flowering varies 2008). Natural seed-fall is correlated with rain significantly from year to year (Beatley 1974). showers, which dislodge the achenes from the Population flowering occurs over a period of floral cup (Auger 2005). Achenes have no 2-3 weeks (Pendleton and Meyer 2004). Indi­ special morphological traits that would aid vidual flowers are open 4-5 days, and most of in dispersal, but they are gathered and cached the anthers dehisce within the first 24 hours by heteromyid rodents (Auger 2005; Bowns (Pendleton and Pendleton 1998). Ftowers are and West 1976). In the Colorado Plateau perfect and regular, with a perianth usually (Auger 2005), these consist primarily of Ord 's consisting of four persistent yellow petaloid kangaroo rat (Dipodomys ordii) and the plains sepals (Welsh et al. 1993). The fntit is a gla­ pocket mouse (Perognathus.flavescens). In the brous chestnut-brown achene. Blackbmsh is Mojave, the chisel-toothed kangaroo rat (D. wind-pollinated and largely self-incompatible, microps) is the major disperser (Beatley 1976). but it exhibits characteristics indicative of evolution from an insect-pollinated ancestor. In addition to the showy petaloid sepals, rare GERMINATION AND ESTABLISHMENT individuals with l--4 yellow petals have been Freshly collected seed ofblackbrush is largely observed in most populations, and plants dormant. Dry after-ripening under summer occasionally produce more than one fruit per field conditions may reduce initial donnancy, flower (Pendleton and Pendleton 1998). and winter chilling removes any remaining Blackbrush is mast-seeding, which is dormancy (Meyer and Pendleton 2005; defined as the "synchronous intermittent Pendleton and Meyer 2004 ). Th~ amount of production of large seed crops in perennial chill required to break donnancy is correlated plants" that "results from weather conditions with collection site's elevation. Seeds from or is an evolved plant reproductive strategy" ]ow-elevation sites have shorter chilling for satiating predators (Kelly and Sork requirements than seeds from higher elevation 2002, 427). Mast-seeding in blackbrush sites. Also, seeds from higher, colder sites, is- a composite of both forces. The size of where winter snow regularly occurs, have the seed crop is related to precipitation and a lower optimum temperature for chilling plant resource reserves, which accumulate and can genninate faster when incubated at slowly in woody desert species (Pendleton, near-freezing temperatures (Pendleton and Meyer, and Pendleton 1995). Mast-seeding Meyer 2004). In the field, germination takes events deplete resource reserves such that, place in late winter, under winter rain condi­ even when adequate winter precipitation tions at warm desert Mojave sites, and under occurs in successive years, black brush does snow at colder Colorado Plateau sites (Meyer not produce successive large fruit crops. and Pendleton 2005). This ecotypic variation A mast-year's crop comprises almost all functions to time germination optimally in of the seed production at low elevations, habitats with contrasting chilling regimes whereas some seeds are produced in more (Pendleton and Meyer 2004). mesic higher-elevation sites in all but the Blackbmsh does not form a persistent seed driest intermast years (Pendleton, Meyer, and bank, and recruitment depends on seed pro­ Pendleton 1995). Periods between mast-seed duced during the previous summer. Seeds are crops often exceed five years. Late frosts collected by heteromyid rodents (kangaroo can reduce or eliminate flowering and fruit Blackbrush (Coleogyne ramosissima Torr.) 147

rats and pocket mice) and cached in scatter invasive grasses, and increased fire frequency. hoards (Auger 2005). Scatter hoarding is the Global climate change models predict higher collection of seed by heteromyid rodents and temperatures and an increase in extreme subsequent burial in small surface caches drought events for the Southwest (Archer (Langland et al. 2001 ). The rodents later and Predick 2008; Cay an et al. 201 0; Seager retum to recollect the seed and store it in et al. 2007). Drought has multiple effects on their burrows (larder hoards). Larder-hoarded seed set and seedling recruitment. Increased seeds are removed from the seed bank, but drought during seed fill can negatively affect scatter hoards that were not recollected form seed production of long-lived mast-seeding a short-term seed bank. Burial by rodents woody species (Perez-Ramos 201 0), and the is generally required for blackbrosh seed stress period for seedlings is the time between to genninate and for successful seedling the loss of winter soil moisture and the onset establishment. of monsoonal moisture in July-September. Following germination, several weeks of While mature plants undergo dormancy during root growth occurs prior to shoot emergence this time, the majority of seedling mortality in early spring (Pendleton 2008). Timing of occurs during this period. Greater than normal emergence varies with region: February for precipitation during this usually dry period wann winter Mojave Desert populations and appears to be the key to seedling survival. March for Colorado Plateau populations. Predicted increases in summer temperatures, Seeds and emerging seedlings are very pal­ coupled with a higher incidence of drought, atable to heteromyid rodents, and therefore, will exacerbate seedling stress and make successful recruitment is primarily limited successful recruitment an even rarer event. to periods following a mast~seed crop when Predicted effects of climate change on some of the cached seed can escape predation recruitment of blackbrush make it unlikely (Auger 2005). Successful establishment is a that blackbrush will persist in place. Current relatively rare event, requiring two years of observations suggest an upward ,.elevational above-average precipitation--one to produce shift in distribution for this species in response a seed crop of sufficient size and another for to a long-term warming trend (Meyer and seedlings to get through the summer. Emerged Pendleton 2005). Data from packrat middens seedlings have very low survivorship over reveal past migration of the blackbrush time due to herbivory and summer moisture ecotone. Blackbrush has been reported from limitation (Bowns and West 1976). Meyer and Late Pleistocene midden sites along the lower Pendleton (2005) found establishment success Colorado River Valley, far to the south of its to be greatest for seeds from local popula­ current range, as well as at lower elevations of tions, indicating ecotypic differentiation in the Grand Canyon (Cole 1990a, 1990b; Hunter emergence and establishment strategies for and McAuliffe 1994; King and VanDevender Mojave and Colorado Plateau populations. 1977; VanDevender 1990). During the late Overall establishment success, however, was Pleistocene and early Holocene, blackbrush much greater on the Colorado Plateau where migrated northward and upward in elevation cooler temperatures and more dependable into the Mojave Desert and Colorado Plateau, summer moisture prevail (figure 10.3). likely reaching its historic distribution in the late Holocene (Coates, Cole, and Mead 2008; Cole 1990a, 1990b; Hunter and McAuliffe RESPONSE TO DISTURBANCE 1994). The paleo records, therefore, indicate AND CLIMATE CHANGE that blackbrush has the ability to migrate in Blackbrush communities are under threat response to changes in climate; however the from the combined effects of climate change, speed at which future changes occur may 35

30

1: 25 0 ~ .~ 20 Q. "<::) -Mojave Desert ~ 15 Q. Colorado E E 10 Plateau

5

0 1 2 3 4 5 6 7 8 9 10 11 12 Month 40 B 35

30 Colorado 25 Plateau - Mean u Max =~ 20 7 Colorado Plateau -Mean ~ 15 , Q.l Min Q. ' E 10 \ -Mojave Desert­ ~ Mean Max 5 +-----~;~------~---- ; 0 -!--_,_....------~ Mojave Desert­ Mean Min -5

-10 1 2 3 4 5 6 7 8 9 10 11 12 Month

Figure 10.3 . Long-term average precipitation (A) and temperature (B) profiles for Colorado Plateau and Mojave Desert blackbrush habitats. For temperature, heavy lines depict mean monthly highs, while broken lines depict mean monthly lows. Graphs represent composite averages for six locations in each region . (Modified from Pendleton et al. 2012.) Blackbrush (Cofeogyne ramosissima Torr.) 149

preclude natural migration for long-lived stand-replacing. Seedlings are not fire-tolerant, species with episodic recruitment. but if a fire occurs in the summer after seeds By far the greatest threat to the blackbrush are cached (June), seeds may survive the fire community comes from the combination of and emerge in the winter and spring (Zitzer exotic grasses and accompanying changes in 2009). Rodent behavior is at least partially the fire regime. Climate change models pre­ responsible for the lack of recovery following dict an increase in the incidence of large-scale large fires, as seed movement by granivores wildfire (Marlon et al. 2009). As stated earlier, into a burned area is restricted (Beatley 1976). the blackbrush community has little resistance In the Mojave, the kangaroo rat (D. microps), to invasion by exotic annual grasses and is which occupies the unburned blackbrush com­ extremely vulnerable to fire (Brooks and munity, does not cache seeds in open areas, Chambers 20 ll ; Brooks and Matchett 2006). such as bums, due to predation threat. On the Historically, low amounts of fine fuels present other hand, small-scale disturbances, such as in shrub interspaces limited the spread offires pipelines, road cuts, and rights-of-way, can be except under extreme conditions (Brooks, revegetated with seed caches and will recover Esque, and Duck 2007), and recovery often in time (Beatley 1976). occurred within a few decades (Brooks and While concern over loss of blackbrush Matchett 2006). Natural fire-return intervals habitat has centered around Mojave popu­ were likely on the order of centuries (Webb lations, the Colorado Plateau is not immune et al. 1987). Extensive burning of blackbrush from similar losses. There are many reports stands in the mid-1900s to improve livestock of dramatic increases in exotic annual grass production had unpredictable outcomes and cover in grasslands of the Colorado Plateau mostly resulted in less desirable vegetation (Belnap, Phillips, and Troxler 2006; Enserink (Bowns and West 1976; Brooks, Esque, and 1999; Hanson 1999; Rimer and Evans 2006; Duck 2007). Burned blackbrush stands took SchwiTll1ing et al. 2008). Although cheatgrass many years to recover, if at all. (Bromus tectorum) does not cun:ently occur Exotic grasses now present in most extensively in blackbrush habitat,- new blackbrush stands in the Mojave create large strains of cheatgrass may be able to adapt amounts affine fuels foJlowing years of high and persist. Furthermore, the impact that rainfall (Brooks and Matchett 2006). Ensuing climate change will have on invasive species fires can bum large areas and encourage dom­ in general and annual grasses specifically inance by early-seral exotic annuals that, in is not well understood. Predictions range turn, perpetuate shorter fire-return intervals from an increase in the number of invasive (Brooks 2009; Brooks and Pyke 2002; Engel species and the expansion of exotic annual and Abella 2011 ). For example, Esque (2009) grasses (e.g., Abatzoglou and Kolden 2011; modeled blackbrush habitat and fire perim­ D'Antonio and Vitousek 1992) to a decrease eters in the Mojave immediately following in potential habitat for cheatgrass, at least, years of high rainfall. Their study revealed in the Southwest (Bradley 2009). Conse­ that nearly 7,700 km2 of black brush habitat quently, while blackbrush communities in the burned in 2005-2006 alone. The Ely Bureau Colorado Plateau have not yet experienced of Land Management District had the highest the same kind of fire-cycle increase that has impact, with about 40% (112,502 ha) of the occurred in the Mojave, it is possible that new district's blackbrush communities burned introductions of invasive species, combined during the 2005 and 2006 fire seasons. with predicted alterations in temperature and Recruitment following fire is seed limited. precipitation patterns that will accompany Partially burned adult plants may recover to climate change, could alter fire patterns on some extent, but most large-scale fires are the Colorado Plateau. 150 Pendleton et al.

RESTORATION POTENTIAL growing in clusters, and natural recruitment AND GUIDELINES often occurs under established nurse plants (Jones 20 12; S. Kitchen unpublished data). Blackbrush is considered to be one of the most It may therefore be useful to establish nurse challenging ecosystems to restore (Brooks plants on large bumed areas by including and Matchett 2006; Hansen and Ostler 2008). other species that have high seedling success There have been attempts to seed blackbrush rates in a seed mix (Abella and Newton (Abella and Newton 2009), but success has 2009), or through the use of transplants to been limited. Transplants have also met with establish shrub islands. It is important, too, mixed success (Monsen, Stevens, and Shaw to use an appropriate seed source, as Mojave 2004). ]tis our hope that an understanding of and Colorado Plateau populations differ in the biology of blackbrush will increase the their germination and establishment strate­ success of future restoration efforts. gies (Meyer and Pendleton 2005; Pendleton Blackbrush mast-seed crops are produced and Meyer 2004 ). There are also elevational infrequently and, therefore, seed availability differences among populations in chilling is likely to be a limiting factor in restoration. requirements. However, blackbrush seed does maintain Restoration may also be accomplished viability in storage. Gennination and emer­ through the use of transplants. Production gence tests on 32 seed collections that had of blackbrush ptants for transplanting is not been kept at room temperature for 12-27 difficult (Pendleton 2008), but use of a well­ years revealed that seed lots with good initial drained soil medium is necessary as seedlings viability maintained a viability of greater than are susceptible to overwatering (Pendleton 80% for 10-12 years (Pendleton et al. 20 12). 2008; Wallace and Romney 1972). Even Large quantities of good quality seed can older, stored seed can be successfully used easily be collected in mast-years and stored to produce seedlings. We have successfully for future use. For example, in 2008, volun­ produced seedlings from seed s~ored for 27 teers collected a large mast crop of Mojave years by first germinating seeds in a growth blackbntsh seed for use in future plant pro­ chamber (Pendleton et al. 2012). In the duction and seeding projects. This volunteer­ Mojave, transplants would need additional collecting effort is a good example of a water to become established, likely for cost-effective method of providing a supply multiple years (Winkel eta!. 1995; Pendleton of seed for use after fire or other disturbances unpublished data). In contrast, Colorado Pla­ (Brean 2008). teau plantings have been established without Germination and establishment success second year supplemental water due to the may be enhanced by planting seeds in the more predictable summer moisture (Hughes fall to meet chilling requirements and at a and Weglinski 1991). depth of 2.5-4.0 em to mimic rodent cache Whether by seedings or transplants, resto­ depth. It may also be necessary to protect ration efforts will meet greater success at the emerging seedlings from rodent predation upper blackbmsh ecotone or above. Predicted (Jones In emergence experiments in , 2012). climate change variables of i.ncreasing C02 the Mojave and Colorado Plateau, protective higher nighttime winter temperatures, and hardware cloth cages were effective in lack of spring and early summer moisture all reducing seed and seedling predation, which suggest that attempts to restore blackbrush resulted in higher survivorship than for communities at lower elevations would be uncaged seeds (Meyer and Pendleton 2005). unlikely to succeed. A recent restoration Seeding in groups (caches) also seems to study using black.brush seed reported larger improve survival. Blackbrush is adapted to and more numerous seedlings at mid and high Blackbrush (Co/eogyne ramosissima Torr.) 151

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