SPECIES: Selaginella densa


Introductory

SPECIES: Selaginella densa
AUTHORSHIP AND CITATION : Crane, M. F. 1990. Selaginella densa. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ [].

ABBREVIATION : SELDEN SYNONYMS : Selaginella rubestris var. densa Selaginella scopulorum Selaginella engelmannii Selaginella engelmannii var scopulorum Selaginella engelmannii var standleyi Selaginella standleyi SCS PLANT CODE : SEDE2 SEDES SEDES2 COMMON NAMES : little spikemoss spikemoss Rocky Mountain selaginella prairie spikemoss little clubmoss TAXONOMY : The currently accepted scientific name of little spikemoss is Selaginella densa Rydb. [31]. There are three varieties of little spikemoss [31]: S. d. var. densa S. d. var. scopulorum (Maxon) Tryon S. d. var. standleyi (Maxon) Tryon LIFE FORM : Fern or Fern Ally FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY

Distribution and occurrence

DISTRIBUTION AND OCCURRENCE

SPECIES: Selaginella densa
GENERAL DISTRIBUTION : The range of little spikemoss extends south from the Alaska panhandle to northwest California and east into Manitoba and the Dakotas. It is common on the Great Plains and in the Rocky Mountains it grows as far south as eastern Utah and Arizona, New Mexico and Trans-Pecos Texas [13,27]. The distributions of the varieties of little spikemoss are as follows [13,26,27,42]: S. d. var. densa is the characteristic variety found on the northern Great Plains. It may occasionally extend into western Montana, southwest Alberta and southeast British Columbia. Further south it is found in the mountains of Colorado and the LaSal Mountains of Utah. S. d. var. scopulorum is the common variety in the mountains of the Northern Rocky Mountain Region through the LaSal and Uinta mountains of eastern Utah to west Texas. It is also found from British Columbia to northern California. It is less common in the mountains of Colorado. S. d. var. standleyi is uncommon and found only above timberline in the northern Rocky Mountains. ECOSYSTEMS : FRES10 White - red - jack pine FRES20 Douglas-fir FRES21 Ponderosa pine FRES23 Fir - spruce FRES26 Lodgepole pine FRES29 Sagebrush FRES30 Desert shrub FRES34 Chaparral - mountain shrub FRES35 Pinyon - juniper FRES36 Mountain grasslands FRES38 Plains grasslands FRES39 Prairie FRES40 Desert grasslands FRES44 Alpine STATES : AZ CA CO KS MT NE NM ND OK OR SD UT WA WY AB BC MB SK BLM PHYSIOGRAPHIC REGIONS : 1 Northern Pacific Border 2 Cascade Mountains 4 Sierra Mountains 5 Columbia Plateau 8 Northern Rocky Mountains 9 Middle Rocky Mountains 10 Wyoming Basin 11 Southern Rocky Mountains 12 Colorado Plateau 13 Rocky Mountain Piedmont 14 Great Plains 15 Black Hills Uplift 16 Upper Missouri Basin and Broken Lands KUCHLER PLANT ASSOCIATIONS : K007 Red fir forest K008 Lodgepole pine - subalpine forest K011 Western ponderosa forest K012 Douglas-fir forest K015 Western spruce - fir forest K016 Eastern ponderosa forest K017 Black Hills pine forest K018 Pine - Douglas-fir forest K019 Arizona pine forest K020 Spruce - fir - Douglas-fir forest K021 Southwestern spruce - fir forest K023 Juniper - pinyon woodland K037 Mountain mahogany - oak scrub K038 Great Basin sagebrush K039 Blackbrush K040 Saltbush - greasewood K051 Wheatgrass - bluegrass K052 Alpine meadows and barren K055 Sagebrush steppe K056 Wheatgrass - needlegrass shrubsteppe K057 Galleta - three-awn shrubsteppe K063 Foothills prairie K064 Grama - needlegrass - wheatgrass K065 Grama - buffalograss K066 Wheatgrass - needlegrass K067 Wheatgrass - bluestem - needlegrass K070 Sandsage - bluestem prairie K095 Great Lakes pine forest SAF COVER TYPES : 1 Jack pine 205 Mountain hemlock 206 Engelmann spruce - subalpine fir 207 Red fir 208 Whitebark pine 209 Bristlecone pine 210 Interior Douglas-fir 218 Lodgepole pine 219 Limber pine 220 Rocky Mountain juniper 237 Interior ponderosa pine 238 Western juniper 239 Pinyon - juniper SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : In the Great Plains grasslands of southern Canada and the northern United States, little spikemoss can be a dominant in three of the five major grass associations: the short-grass prairie, mixed prairie and fescue (Festuca scabrella) prairie [7,9]. At higher elevations in Glacier Park, it codominates with Idaho fescue (F. idahoensis) in a climax grassland association [48]. In the Custer National Forest of Montana, North and South Dakota and the Theodore Roosevelt National Park of North Dakota, little spikemoss is rare outside of the needle and thread grass (Stipa comata)/threadleaf sedge (Carex filifolia) habitat type where it is an important species [24,25]. Published classification schemes listing little spikemoss as an indicator species or a dominant part of vegetation in plant associations (pas) are presented below: Area Classification Authority Region 2: CO,NE,KS, general veg. pas Johnston 1987 SD,WY
Management Considerations

MANAGEMENT CONSIDERATIONS

SPECIES: Selaginella densa
IMPORTANCE TO LIVESTOCK AND WILDLIFE : Little spikemoss is a minor part of elk diets but is not a valuable elk browse [36]. During a Montana study domestic animals were never observed grazing on little spikemoss [57]. Bighorn sheep may eat some spikemoss (Selaginella spp.) in New Mexico [40]. PALATABILITY : The relish and degree of use shown by livestock and wildlife species for little spikemoss (var. densa) in several western states is rated as follows [16]: UT WY MT ND Cattle poor poor poor poor Sheep poor poor poor poor Horses poor poor poor poor Elk fair poor poor ---- Mule deer fair poor poor fair White-tailed deer ---- poor ---- fair Antelope fair poor poor fair Upland game birds fair fair ---- poor Waterfowl fair ---- ---- ---- Small nongame birds fair ---- ---- ---- Small Mammals fair fair ---- ---- NUTRITIONAL VALUE : The energy and protein value of little spikemoss forage is poor [16]. Aboveground parts of little spikemoss have a high ash content (19.7%). Most of this is silica which comprises about 16.5 percent of plant dry weight. Other results of this analysis are given below [59]: Percent of dry weight of aboveground portions of plant Calcium 0.20 Potassium 0.30 Iron 0.038 Zinc 0.023 Sodium 0.29 Phosphorus 0.11 Protein 8.50 COVER VALUE : Little spikemoss provides poor cover for small animals [16]. VALUE FOR REHABILITATION OF DISTURBED SITES : Little spikemoss has poor short-term or long-term revegetation potential [16]. It does grow well on gentle and moderate slopes and it has low to moderate erosion control potential. However, its potential biomass production is low and establishment requirements may be exacting [16]. OTHER USES AND VALUES : While little spikemoss does not provide forage, it does protect the soil from erosion by wind or water and reduces the effects of tramping [7]. With care, little spikemoss can be transplanted into gardens where it is an attractive ground cover that grows on rocks and in other dry situations [34]. OTHER MANAGEMENT CONSIDERATIONS : The role of little spikemoss in mixed prairie is not well understood, although its influence on other vegetation and the habitat has been thought to be slight because its requirements for water and nutrients are low [9,10,60]. During light showers, its dense mats and extensive, shallow roots may intercept and soak up all the available water [55,57]. During high intensity storms, the mats of little spikemoss act to increase infiltration and decrease runoff and surface erosion [57]. Its shallow root system also limits spikemoss growth to periods of the year when moisture is available at shallow depths [59]. Grazing effects: The effect of grazing on little spikemoss seems to be confounded by drought. In Canadian and Montana mixed prairie, little spikemoss is able to increase during periods of drought [10,18,58]. Also, little spikemoss increases when protected from grazing and decreases with grazing due to trampling of grazing animals [10]. Little spikemoss is relatively weakly rooted and easily damaged by trampling [37]. Decreases in response to grazing may also be explained by the exposure and drying of the surface soil when cover is removed by grazing [18]. In the short-grass prairie of southeastern Alberta and southwestern Saskatchewan, grasses can easily establish in areas with high cover of little spikemoss during years with above normal precipitation, but they are rarely able to establish during dry years [7]. A 4-year Montana study done during a period of normal rainfall compared ungrazed mixed prairie with mixed prairie moderately grazed by sheep. Cover of little spikemoss decreased from 21 to 17.6 percent on the ungrazed site but it decreased from 15.1 to 7.1 percent on the grazed area [58]. The strong decrease on grazed prairie was attributed to trampling, competition, and manure [57]. A 7-year study of three levels of grazing intensity on the short-grass prairie in southeastern Alberta and southwestern Saskatchewan was done during dry and drought years [9]. Little spikemoss increased under all levels of grazing, with the greatest increase in the most lightly grazed area. On ungrazed land little spikemoss increased in basal area from 11 percent to 16 percent. Results of a second 10-year study showed the basal area of little spikemoss increased by 37 percent on ungrazed and 27 percent on grazed land, while grass cover generally decreased. Looking at total vegetation cover the authors concluded that "climate rather than moderate grazing use was the principal factor affecting plant cover" between 1928 and 1939. During a third 5-year study, little spikemoss increased more under a medium level of grazing than under severe grazing intensity [9]. Little spikemoss increases more under rotational grazing than under continuous grazing [9,65]. A 3-year North Dakota mixed prairie study compared exclosures established 38 to 41 years previously with cattle-grazed range. Little spikemoss cover did not vary significantly despite major differences in species composition of grasses and silver sagebrush (Artemisia cana) [5,6]. Site differences were more important than grazing pressure [6]. Ross and Hunter [50] list spikemoss as increasing under grazing pressure in Montana. However, little spikemoss increases very slowly when taller, competing vegetation is partly or wholly removed [52]. Treatment with fertilizer: The primary effect of manure or fertilizer applications is to increase the competitiveness of taller grasses, thus decreasing little spikemoss [18]. However, in the northern plains response depends upon precipitation [54]. There is no evidence that nitrogen application alone kills little spikemoss [52]. When fertilizers containing nitrogen and phosphorus were added to grassland plots in each of 2 successive years, the cover of live little spikemoss decreased significantly. It was not determined whether this was due to the nutrients themselves or because of increased competition from other vegetation [57]. Manuring alone increased all plant cover the first season of a Montana study, but little spikemoss was subordinated to the more vigorous native grasses and decreased the second season [18]. After more than 30 years with normal weather, little spikemoss was still reduced in plots treated by annual applications of manure for 1 to 11 years [19]. Mulching, with or without added fertilizer, can nearly eliminate little spikemoss [66]. Mechanical treatment: Mechanical treatments reduced little spikemoss cover between 25 and 70 percent in northern Montana mixed prairie [52]. The effects of various combinations of manure treatment, disking, and harrowing over a 10-year period along with seeding of crested wheatgrass (Agropyron cristatum) and yellow sweetclover (Melilotus officinalis) have been studied [18,19]. Most treatments resulted in long-term decreases in little spikemoss, and combination treatments seemed most successful. When the treated area was resurveyed over 30 years later, little spikemoss cover was still so decreased that researchers concluded that little spikemoss does not easily reestablish on northern mixed prairie following mechanical treatment [19]. Little spikemoss cover and the cover of either other vegetation or litter were significantly and negatively correlated [19]. Chemical control: The most effective chemicals for control of little spikemoss while increasing the yield of desirable grasses are AMS (ammate) and atrazine [59]. Spring treatment is most effective. Monuron, paraquat, and bromacil control little spikemoss but also reduce the yield of or kill other vegetation [59]. Further range tests indicate that ammate, atrazine, and monuron are the best treatments [52]. Their effectiveness is increased with the use of a surfactant [52].
Botanical and ecological characteristics

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Selaginella densa
GENERAL BOTANICAL CHARACTERISTICS : Description: Little spikemoss is an evergreen, nonflowering herb that forms dense, cushionlike mats which are seldom more than 1 inch (2.5 cm) in height [10,57]. The short, leafy, compactly branched stems lie along the soil surface and may be 4 inches (1 dm) across [27]. The simple, awn-tipped leaves are very small, up to about 0.1 inch (2.5 mm) long and 0.02 inch (0.5 mm) wide, and arranged in a relatively dense spiral along the stem. There is usually a single, unbranched vein. The leaves underneath the stem are longer than leaves on top along the same part of the stem [42]. Little spikemoss has true roots which are very fine (0.008 inch or 0.2 mm in diameter) and minutely branched. They form a tangled mass and may comprise 86 percent of the plant dry matter [59]. Most roots occur within 0.78 to 2.0 inches (2-5 cm) of the soil surface [12]. Drought Resistance: Little spikemoss is very resistant to drought if the roots are not disturbed. In the laboratory, clumps allowed to desiccate for 6 and 33 months were then revived simply by watering and later planting [55,61]. The clump that was revived after 33 months developed reproductive cones within 3 months [61]. When all soil was removed from roots, no plants survived after 5 weeks of air drying [59]. Shoots that are dormant during drought have their leaves closely appressed to the stem and are a dull grayish-green instead of the ash-gray of dead shoots [61]. Young plants are equally drought resistant. On a site near Saskatoon, Saskatchewan, Webster and Steeves [61] found over 50 locations where sporelings had survived periods of drought. RAUNKIAER LIFE FORM : Undisturbed State: Chamaephyte Burned or Clipped State: Chamaephyte REGENERATION PROCESSES : The spikemoss life cycle differs from that of flowering plants. At branch ends, four-angled "cones" or strobili are formed. Each has two types of spore-bearing structures, microsporangia and megasporangia, in the leaf axils, with megasporangia located below the microsporangia [61]. Microspores are released passively to fall through the strobili to the ground [32]. At maturity they will release microscopic, flagellated sperm cells. The sperm require water to swim to the egg cell which develops from cells of the megasporangia. Where fertilization occurs is not known [52]. Tissue surrounding the fertilized egg may carry on photosynthesis as well as provide stored food for the developing embryo. This entire structure may be shed or held in the leaf axils of the strobili while the embryo develops. The necessity of water for sexual reproduction restricts the habitat of spikemosses [15], and young plants are not often reported [18,52]. As little spikemoss clumps grow the centers die, leaving a ring of live material surrounding a dead core [59]. Under range conditions lateral growth was observed to be less than 0.4 inch (1 cm) a year, making vegetative spreading a slow process [18,59]. During another field study growth of less than 0.2 inch (5 mm) in 5 seasons was observed [52]. Fragmentation might be another means of vegetative reproduction and dispersal [18], although stems of little spikemoss do not have a natural tendency to fragment when dry [55]. SITE CHARACTERISTICS : Little spikemoss is most often found on dry shallow soils that are gravelly or rocky or have gravelly or stony subsoils [57]. It is most common in grasslands, alpine tundra, and high subalpine communities which occupy dry, snow-free ridges. Little spikemoss is an extremely stress-tolerant species and can survive in cold alpine regions with a short growing season, drought, and summer frosts [62]. Little spikemoss is not common in forests but can be found in Douglas-fir (Pseudotsuga menziesii) with ponderosa pine (Pinus ponderosa) or lodgepole pine (P. contorta), Engelmann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa), or aspen (Populus tremuloides) communities [33,44]. Little spikemoss also grows in pinyon (Pinus edulis)-juniper (Juniperus spp.), sagebrush (Artemisia spp.), and krummholz communities [63]. In Jackson Hole, Wyoming the big sagebrush (A. tridentata) community has abundant little spikemoss [47]. In northern Montana aspen grove and grassland communities, it is found primarily with Hood's phlox (Phlox hoodii) on stony moraine summits, outwash fans and coarse gravels [38]. The abundance of little spikemoss in the aspen grove and grassland community decreases to the north in Saskatchewan [11]. Grassland Sites: Little spikemoss is most abundant in short-grass prairie. Its basal area may be equal to that of all other herbs and shrubs [7,9]. In mixed prairie, little spikemoss is slightly less abundant, occurring with an overall frequency of 74 to 98 percent and cover up to 25 percent [10]. It has much less cover and is not evenly distributed in the fescue prairie [7,11]. In the northern Red River Valley, it dominates with needle-and-thread grass [45]. On sites with standing surface water, little spikemoss will be unimportant [64]. However, in Montana, areas where it grows are closer to permanent water than areas where it does not grow [57]. In North Dakota, little spikemoss is common on a variety of grassland habitat types except those with extra moisture [64]. Alpine and subalpine sites: In the Washington and British Columbia Cascades, little spikemoss is common in herb field, the Bellard alpinesedge (Kobresia myosuroides), and purple reedgrass (Calamagrostis purpurascens) communities on sites that are essentially snow-free during the winter. On such sites it is exposed to high wind, low temperatures, and frequent frosts all winter [20]. On alpine sites in Montana, little spikemoss does well where Hooker mountainavens (Dryas octopetala) is dominant [2]. In alpine areas in Utah, it grows in cushion plant communities with alpine avens (Geum rossii) and on dry meadows and alpine turf [37]. On the Colorado Front Range, little spikemoss is part of several forest communities, but its greatest abundance is at higher elevations in Bellard alpinesedge meadows or under limber pine (Pinus flexilis) on exposed rocky sites [33,44]. It is well adapted to these and other alpine cushion-plant communities [62]. In contrast, near Schoolroom Glacier in the Teton Range of Wyoming, it is found in a meadow community with kentrophyta milkvetch (Astragalus kentrophyta) [53]. In Alberta it grows under limber pine on the rocky, dry slopes and summits of exposed ridges [41]. In northern California little spikemoss (var. scopulorum) grows on open rocky sites in red fir (Abies magnifica) and lodgepole pine forests of Siskiyou County [42]. Parent material: Little spikemoss grows in the drier interior of British Columbia except for a disjunct population on dry, well-aerated and warm limestone soils on the Queen Charlotte Islands [49]. In Montana, little spikemoss does well on sites with thin, rocky, and unstable soils derived from calcareous parent material [2]. In the Bighorn Mountains of Wyoming, it is present on granitic soils but almost absent on soils derived from sedimentary rock [57]. Soil texture: Little spikemoss is common on medium texture soils and sandy loams. It is less common on sandy soils and rare on clay [10,52,57]. In grasslands dominated by bluebunch wheatgrass (Pseudoroegneria spicata) little spikemoss is found in areas with shallower, rockier, and less sandy soils than areas without little spikemoss [57]. Optimum soil depth for little spikemoss (var. densa) is between 10 and 20 inches (25.4-50.8 cm) [16]. Growth of little spikemoss (var. densa) on various soils in several western states is ranked as follows [16]: UT WY MT ND Soil texture Gravel good fair fair fair Sand good poor fair fair Sandy loam good good fair fair Loam good good good good Clay loam good good good good clay fair fair fair good dense clay poor poor poor poor Organic soils fair poor poor ---- Acidic soils fair poor poor ---- Saline soils fair poor poor poor Sodic soils ---- poor poor fair Sodic-saline soils poor poor poor poor Elevation: Elevational ranges in some western regions are [16,26,30,42,63]: Minimum Maximum feet meters feet meters Arizona 11,470 3,496 11,470 3,496 California var. scopulorum 5,000 1,524 7,000 2,134 Colorado var. densa 4,000 1,219 12,000 3,658 var. scopulorum 8,000 2,438 12,500 3,810 var. standleyi 9,500 2,896 13,000 3,962 Montana var. scopulorum 6,300 1,920 6,300 1,920 Utah 8,858 2,700 14,107 4,300 Wyoming var. densa 6,000 1,829 10,800 3,292 var. scopulorum 5,600 1,707 10,800 3,292 var. standleyi 5,600 1,707 10,800 3,292 SUCCESSIONAL STATUS : In general, spikemosses (Selaginella spp.) are unable to compete with other plants in relatively moist habitats [55]. They also are not important pioneers. Instead spikemosses are found on open, xeric sites supporting particular plant communities but shift locally within those communities. On such open sites their ability to withstand desiccation allows them to survive and grow well [55]. On alpine sites of Montana's Beartooth Plateau, little spikemoss pioneers on the gravel mulch left by rodent disturbance in grassy meadows [29]. Little spikemoss (var. scopulorum) is found in alpine areas on unstable soils disturbed by frost and abraded by wind [29]. SEASONAL DEVELOPMENT : In North Dakota spore production begins in June and finishes in July [16]. In Canadian mixed prairie, vegetative growth begins in early April, strobili first appear late in April, and spores mature late in May [10]. In the Colorado alpine, leaves of little spikemoss remain green all winter when protected from wind but turn brown and erode where exposed [3]. The cones (strobili) change from green to brown in late May or early June when they are released from snow [3].
Fire ecology

FIRE ECOLOGY

SPECIES: Selaginella densa
FIRE ECOLOGY OR ADAPTATIONS : Little spikemoss would appear to have little defense against fire. However, it has been observed to survive on dry rocky hillsides where other vegetation and litter were sparse. On nearby moist areas where prefire vegetation was more dense, it was completely removed [51]. POSTFIRE REGENERATION STRATEGY : secondary colonizer; off-site spores carried to site after year two
Fire effects

FIRE EFFECTS

SPECIES: Selaginella densa
IMMEDIATE FIRE EFFECT ON PLANT : Little spikemoss is killed by fire except where it is protected by site conditions [51]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : Generally, spikemosses are lacking following recent fire or other major disturbance [55]. In a study of paired stands in mixed grass prairie in Manitoba, little spikemoss cover was 14.9 percent in a control plot, 6.2 percent following a single fire and 2.3 percent following two fires [67]. In paired stands in western North Dakota, the frequency of little spikemoss was much less following fires that burned from 3 months to 4 years before sampling [17]. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : NO-ENTRY FIRE MANAGEMENT CONSIDERATIONS : Prescribed fire can be used to reduce the cover of little spikemoss. Any method of reducing little spikemoss would be most successful if done during a cycle of wet years on the mixed prairie, as drought would severely set back the recovery of desirable vegetation [1].

Selaginella densa: References


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