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SPECIES: Sarcobatus vermiculatus | Black Greasewood
GENERAL BOTANICAL CHARACTERISTICS : Black greasewood is a native, long-lived, winter deciduous, perennial shrub which grows 3.3 to 8.2 feet (1.0-2.5 m) in height [11,46,55]. It is most typically a freely-branched and spreading shrub, although rounded and erect growth forms also occur [20,55]. Branches, some of which end in spines, are rigid and numerous [20,55]. Bark is smooth and whitish, and graying at maturity [21,55]. Black greasewood typically has a long taproot. Roots can extend 20 to 57 feet (6.1-17.4 m) below the soil surface [13]. Leaves of black greasewood are simple, linear, alternate, and fleshy, with entire margins [20,55]. The leaves are shed in winter. Numerous staminate flowers are borne on fleshy, catkinlike terminal spikes, whereas pistillate flowers form singly or in pairs in the axils of leaflike bracts [7,55]. Fruit is a small, coriaceous achene which is winged at the middle [55,59,61]. The fruit contains small brown seeds [34,61]. RAUNKIAER LIFE FORM : Phanerophyte REGENERATION PROCESSES : Seed: Black greasewood is generally monoecious. Staminate and pistillate flowers mature at different times, thus maximizing opportunities for cross-pollination [49]. According to McArthur and Sanderson [30], the investment in male and female functions varies from dry to mesic sites, allowing for efficient occupation of patchy, stressful, and heterogeneous environments. Seeds mature in the fall and are dehisced over the winter [49]. Some seed is produced annually [39,55], but abundant seed production is limited to occasional years [4]. Seed production also apparently varies according to levels of disturbance. Roundy and others [49] report that black greasewood is a poor seed producer on undisturbed sites. Only 20 percent of the plants on undisturbed sites produced seed, with an average total of less than 20 seeds per plant [49]. However, where 30 percent of the black greasewood had been killed, resprouted plants produced an average of 250 seeds [49]. Seed fill varies from 16 to 94 percent [11]. Germination: An afterripening period of 30 to 60 days is generally required for the embryo to mature [11]. Laboratory studies have indicated that optimum germination occurs within an average of 5.5 days at a constant temperature of 52 degrees F (11 degrees C) and -16 moisture requirement bars [50]. Under natural conditions, most seeds germinate during relatively long periods of high soil moisture [46]. The presence or absence of light does not appear to affect germination rates [50]. The presence of the membranous pericarp and enclosing papery bracts, however, does influence germination [64]. When the pericarp was broken, the embryo uncoiled within an hour, and root hairs developed within 24 hours [11]. When the pericarp remained undamaged, the seeds quickly imbibed water, but germination was slow and many seeds had not germinated after 30 days [11]. Both inter- and intra-population differences in germination have been observed at varying osmotic potentials and salt concentrations, and ecotypic variation in germination characteristics is suspected [46]. A certain proportion of seed commonly fails to germinate at low osmotic potential or where salt concentrations are high. These remaining seeds may germinate later, thus representing an adaptation to a range of environmental conditions [46]. Vegetative regeneration: Black greasewood typically sprouts after fire, application of herbicides, and other types of disturbance [62,65]. SITE CHARACTERISTICS : Black greasewood grows on dry, sunny, flat valley bottoms, on lowland floodplains, in ephemeral stream channels, and at playa margins [49,57]. It is a dominant plant throughout much of the Great Basin and Mojave Desert. Black greasewood occurs in salt shrublands, inland saltgrass, northern and southern desert shrub, and pinyon-juniper communities [34,37]. Black greasewood communities generally occur below the moister sagebrush or shadscale zones [49]. In high saline areas, black greasewood often grows in nearly pure stands, although on less saline sites it commonly grows with a number of other shrub species and typically has a grass understory [29]. Soil: The growth and distribution of black greasewood is affected by soil salinity, ionic concentrations, soil depth, and total water potential [48]. Black greasewood is often considered an indicator of saline-sodic or relatively moist soils [44,50]. Sites typically have clay-loam, silt-loam, or deep, fine, sandy loam soils with high salinity or alkalinity [11,21]. Although black greasewood most commonly develops on finely textured saline or alkaline soils, it occasionally grows on coarsely textured nonsaline soils [46]. Black greasewood is often abundant on outcrops of alkaline or salt-bearing shales with little soil development [38,53]. In some areas, the salt content of the soil 3 feet (1 m) below the surface reaches 1.08 percent [13]. When salinity increases above this level, black greasewood becomes yellow and dwarfed [13]. Representative pH levels at a black greasewood site are as follows [6]: pH Na (me/100g) Ca (me/100g) upper soil 7.9 9.1 9.1 lower soil 8.0 12.0 15.4 Exchangeable sodium can range up to 12.0 me/100 g, with a pH as high as 9.82 [11]. Black greasewood is highly tolerant of boron, as well as sodium, in the soil beneath the canopy [43,48]. Climate: Black greasewood is tolerant of a wide range of climatic conditions but most commonly grows in areas with hot, dry summers [44]. The distribution of black greasewood is thought to be highly dependent on a high soil moisture content below the depth of seasonal precipitation percolation [45,57]. It commonly occurs in areas with a seasonally high water table and is often the only green shrub in pluvial desert sites with available groundwater [63]. Average annual precipitation ranges from 5 to 10 inches (12-25 cm) [48]. The water table is generally within 14.8 feet (4.5 m) of the soil surface, and may rise seasonally to 5 to 3.3 feet (1.5-1 m) below the surface [48]. Elevation: Elevational ranges by geographic area have been documented as follows [10,34,61]: from 3,000 to 7,000 feet (914-2,134 m) in CA 4,500 to 8,500 feet (1,373-2,593 m) in CO 4,000 to 7,118 feet (1,220-2,170 m) in UT 3,600 to 7,300 feet (1,098-2,227 m) in WY SUCCESSIONAL STATUS : Black greasewood, a long-lived species, occurs as a nonclimatic climax indicator in many saltbush, sagebrush, or sagebrush-grassland communities. It is described as a "stable dominant" under moist-sodic edaphic climax conditons [11]. Black greasewood can compete after disturbance and is also well represented in a number of early seral communities. Many Great Basin sites now occupied by black greasewood were formerly dominated by Great Basin wildrye (Leymus cinereus) or big sagebrush (Artemisia tridentata) [7,43]. Soil changes in the form of increased salinity brought about by litter deposition of greasewood may have led to the replacement of sagebrush [43]. With increasing salinity (above 1.08 percent), black greasewood is replaced by species such as inland saltgrass (Distichilis stricta var. stricta), pickleweed (Allenrolfea occidentalis), or samphire (Salicornia utahensis) [13]. SEASONAL DEVELOPMENT : Vegetative development: Leaves of black greasewood appear in mid to late spring [55]. Vegetative growth is initially slow, but a period of accelerated growth, lasting from 3 to 6 weeks, occurs annually from late May to early July [49]. Increased soil temperatures may promote accelerated growth of greasewood, while low soil moisture ends this rapid period of growth [49]. Accelerated leader growth at a Nevada site was documented as follows [49]: Onset Date Cessation Date Average Range Average Range May 15 May 10-25 June 30 June 25-July 10 May 15 May 15 June 15 June 5-25 May 15 May 10-25 June 20 June 5-July 10 Leaves of black greasewood are shed in early fall or over the winter months [49]. Phenological development of black greasewood at a southwestern Colorado site was documented as follows [5]: Stage Date dormant October-December early leaf March full leaf April flower buds developing April early bloom May fruit developing June seed ripening July seed shattering July-September Flowering: The appearance of spikes coincides with the onset of rapid growth, with spikes opening after maximum vegetative growth is completed [49]. Reddish-green, winged utricles are formed by the pistillate flowers from late July to late August [11,49]. Floral development at a Nevada site was as follows [49]: Staminate Staminate Staminate Winged calyxes Utricles flowers appear flowers open flowers dried formed matured May 25 June 8 July 1 July 20 October May 16 June 7 June 20 July 25 October Generalized flowering by geographic area is as follows [10,34]: Location Beginning of flowering End of flowering CA May August CO May August MT June June ND June July UT July October WY May September Seed matures from late July through November. Seeds are shed from late fall through the following spring [11,49].

Related categories for Species: Sarcobatus vermiculatus | Black Greasewood

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