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Tortula ruralis

Tortula ruralis gametophyes (left) and gametophytes with sporophytes (right). Photos courtesy of Michael Lüth, hosted by the USDA-NRCS PLANTS Database. In: Lüth, M. 2004. Pictures of bryophytes from Europe [CD-ROM]. Published by the author.


SPECIES: Tortula ruralis
AUTHORSHIP AND CITATION : Matthews, Robin F. 1993. Tortula ruralis. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: []. Revisions : On 25 January 2016, the above photos and the life cycle figure in the Regeneration Processes section were added to this Species Review. ABBREVIATION : TORRUR SYNONYMS : Tortula intermedia (Bridel) De Notaris Tortula ruraliformis (Besch.) Ingham Barbula ruralis Hedw. Syntrichia intermedia Brid. NRCS PLANT CODE [42]: TORU70 COMMON NAMES : twisted moss tortula moss star moss TAXONOMY : The scientific name of this moss is Tortula ruralis (Hedw.) Gaertn., Meyer, & Scherb. (Pottiaceae) [7,9,10,24,29]. The following varieties are recognized in North America [11]: Tortula ruralis var. ruralis Tortula ruralis var. crinita De Not Tortula ruralis var. hirsuta (Vent.) Kramer LIFE FORM : Bryophyte FEDERAL LEGAL STATUS : No special status OTHER STATUS : Information on state- and province-level protection status of plants in the United States and Canada is available at NatureServe.


SPECIES: Tortula ruralis
GENERAL DISTRIBUTION : Tortula ruralis is a cosmopolitan species found in arctic, boreal, temperate, and desert regions. It is distributed throughout Canada, much of the United States, Mexico, and the Pacific islands [16,24,29,40]. It apparently is more common in western North America than in the eastern provinces and states [18,24]. Tortula ruralis occurs in New York [42]. Tortula ruralis is widespread in Europe, Asia, the Middle East, North and South Africa, South America, and Australia [10,16,24]. ECOSYSTEMS : FRES10 White - red - jack pine FRES11 Spruce - fir FRES20 Douglas-fir FRES21 Ponderosa pine FRES22 Western white pine FRES23 Fir - spruce FRES24 Hemlock - Sitka spruce FRES25 Larch FRES26 Lodgepole pine FRES27 Redwood FRES29 Sagebrush FRES30 Desert shrub FRES31 Shinnery FRES32 Texas savanna FRES33 Southwestern shrubsteppe FRES34 Chaparral - mountain shrub FRES35 Pinyon - juniper FRES36 Mountain grasslands FRES37 Mountain meadows FRES38 Plains grasslands FRES39 Prairie FRES40 Desert grasslands FRES41 Wet grasslands FRES42 Annual grasslands FRES44 Alpine STATES : AK AZ CA CO CT DE HI ID IL IN IA KS KY ME MD MA MI MN MO MT NE NV NH NJ NM NY ND OH OK OR PA RI SD TX UT VT VA WA WV WI WY AB BC MB NB NF NT NS ON PE PQ SK YT MEXICO BLM PHYSIOGRAPHIC REGIONS : 1 Northern Pacific Border 2 Cascade Mountains 3 Southern Pacific Border 4 Sierra Mountains 5 Columbia Plateau 6 Upper Basin and Range 7 Lower Basin and Range 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 : widely distributed, occurs in most Kuchler Plant Associations within its range SAF COVER TYPES : widely distributed, occurs in most SAF Cover Types within its range SRM (RANGELAND) COVER TYPES : widely distributed, occurs in most SRM Cover Types within its range HABITAT TYPES AND PLANT COMMUNITIES : Tortula ruralis occurs in a wide variety of habitats, including arctic-alpine tundra, wet and dry coniferous forests, grasslands, sagebrush, and deserts [12,22,32,36]. The author has been unable to determine if T. ruralis occurs in southeastern ecosystems or in broadleaf forests. Although T. ruralis can be an important ground cover component, it generally is not a dominant species. However, the following publication classifies T. ruralis as a dominant ground cover species in purple pinegrass (Calamagrostis purpurascens) vegetation types in the Yukon Territory: Vegetation types and environmental factors associated with Foothills Gas Pipeline Route, Yukon Territory [38] Species commonly associated with T. ruralis in sagebrush-grassland habitats include basin big sagebrush (Artemisia tridentata ssp. tridentata), Wyoming big sagebrush (A. t. ssp. wyomingensis), gray low sagebrush (A. arbuscula ssp. arbuscula), bluebunch wheatgrass (Pseudoroegneria spicata), Idaho fescue (Festuca idahoensis), cheatgrass (Bromus tectorum), Thurber needlegrass (Stipa thurberiana), bottlebrush squirreltail (Elymus elymoides), Sandberg bluegrass (Poa secunda), phlox (Phlox spp.), broom snakeweed (Gutierrezia sarothrae), hawksbeard (Crepis spp.), buckwheat (Eriogonum spp.), and lichens [4,12,14,19,41].


SPECIES: Tortula ruralis
IMPORTANCE TO LIVESTOCK AND WILDLIFE : No information was available on this topic. PALATABILITY : No information was available on this topic. NUTRITIONAL VALUE : No information was available on this topic. COVER VALUE : No information was available on this topic. VALUE FOR REHABILITATION OF DISTURBED SITES : No information was available on this topic. OTHER USES AND VALUES : No information was available on this topic. OTHER MANAGEMENT CONSIDERATIONS : Cryptogamic soil crusts, including crusts formed by T. ruralis, are an important component of many arid rangeland ecosystems in the western United States. They are important in the reduction of soil erosion, and facilitate vascular plant seedling establishment by improving water penetration and reducing runoff [22,23]. Soils with high electrical conductivity, high phosphorous, and high salt contents facilitate the formation of cryptogam crusts [1]. Heavy grazing, especially during seasons of low precipitation, high temperature, and persistent winds can seriously damage or destroy crusts formed by T. ruralis and other cryptogams. During these seasons, T. ruralis is most likely dormant and brittle, and very susceptible to trampling by livestock [2]. In Navaho National Monument, Arizona, T. ruralis was reduced from 6.3 percent cover in ungrazed areas to 1.0 percent cover in heavily grazed areas [6]. In semiarid and arid grasslands of Canyonlands National Park, Utah, cryptogams, including T. ruralis, are instrumental in the build-up of organic matter and soil nutrients. Cryptogam cover stabilizes soils eroded by heavy winds and torrential rains, especially in undisturbed areas [27]. Relative abundance of T. ruralis was "significantly reduced" in formerly grazed areas, but was high in undisturbed climax grasslands. The ungrazed areas of the park had an average of six cryptogam species per site, with a total cryptogram coverage of 38 percent. The formerly grazed areas had an average of two cryptogam species per site, with a total coverage of 5 percent. This difference suggests that cryptogam species such as T. ruralis may play a more important role in the stability of desert grasslands than previously recognized. The surface soils of formerly grazed areas had less organic matter, less available phosphorous, and higher calcium content due to slow sheet erosion caused by a lack of protection from an established cryptogam cover [26]. In sagebrush types in Idaho and Oregon, T. ruralis litter and cover may exclude or retard the growth of perennial grasses such as bluebunch wheatgrass, Thurber needlegrass, and bottlebrush squirreltail [37,41]. However, this may be because T. ruralis occurs on sites that tend to have less organic matter accumulation, decreased cation exchange capacity, and decreased total nitrogen compared to sites that support the perennial grass species [41]. Drought and dessication tolerance of T. ruralis has been studied extensively in order to determine if similar physiological mechanisms will improve drought resistance of commercial crops [13,21,34]. T. ruralis can be dried to less than 20 percent of its original fresh weight and will immediately resume protein synthesis upon rehydration [13]. Metabolic activities will resume even after 70 years of dessication [34].


SPECIES: Tortula ruralis GENERAL BOTANICAL CHARACTERISTICS : Tortula ruralis is a short, erect moss that forms fairly large, loose to dense tufts. Stems are usually dichotomously branched and stand 0.4 to 1.6 inches (1-4 cm) high. The conspicuous gametophye is anchored to the soil by rhizoids [10,18,24]. Tortula ruralis undergoes marked morphological and color changes between wet and dry states. When wet, leaves are not twisted around the stem, and it is bright green. When dry, leaves twist around the stem, and it is red-brown in color [10,33,40]. It is generally a xerophytic moss, and is highly drought and dessication tolerant [13,21,33]. RAUNKIAER LIFE FORM : Hemicryptophyte REGENERATION PROCESSES : Tortula ruralis is dioecious, and undergoes a distinct annual reproductive cycle invovling a gametophye and sporophyte generation [8,33,40]: It also spreads vegetatively by forming gemmae. When gemmae detach and disperse to favorable sites, a new gametophyte is formed [24]. Tortula ruralis spororphytes probably also spread by fragmentation. SITE CHARACTERISTICS : Tortula ruralis is found in a wide range of sites including, but not limited to, sandy lake shores, rock, nunataks, and hummocks [5,18,19,20,24]. It grows on rocks and soils of many types, but they are most often calcareous [10,40]. Moisture regimes where T. ruralis occurs vary from arid and semiarid to mesic, and it is found from low to high elevations [16,35,39]. On Ellesmere Island, Northwest Territories, T. ruralis is found on high arctic uplands from sea level to 3,300 feet (1,000 m) elevation [5]. SUCCESSIONAL STATUS : In western Montana, T. ruralis is part of the moss mats that invade interstices between rocks on talus slides. They play an important part in talus succession. The moss mats eventually form a carpet over the rocks, stabilizing the talus formation. The talus then colonized by higher plants [31]. In sagebrush-steppe types in western Colorado, Bonham [4] classifies T. ruralis as a stress-tolerant ruderal. Tortula ruralis tolerates full sun to full shade [16,35]. It is abundant in climax grasslands [22,23,25,26]. It is also found in climax grand fir (Abies grandis)/queencup beadlily (Clintonia uniflora) habitats in the Swan Valley, Montana [30]. Members of the T. ruralis complex form climax moss associations in central Idaho forests [39]. Tortula ruralis had the following relative abundance and cover percentage during successive stages of recovery from grazing in Canyonlands National Park, Utah (first samples were taken 5 years after grazing pressure had been eliminated) [25]: Year Sampled Relative Abundance Cover ______________________________________________________________(%)________ Ungrazed 1967 9130 18.0 Grazed 1967 660 1.35 Grazed 1977 6900 4.0 In Camp Floyd State Park, Utah, T. ruralis constituted 0.3 and 6.1 percent of the cover in areas not grazed for 7 and 20 years, respectively [22]. SEASONAL DEVELOPMENT : In New Mexico, twisted moss branches appear in mid-winter, lengthen slowly through the spring, and rapidly through the summer. Fertilization takes place in early winter. Female gametophytes occur from December to June but male gametophytes are rarely observed [33].


SPECIES: Tortula ruralis
FIRE ECOLOGY OR ADAPTATIONS : Specific information on the fire ecology of T. ruralis was lacking when this Species Review was written (1993). However, poikilohydric plants such as T. ruralis dry quickly during periods of low relative humidity because of their absence of roots and water storage tissue, and low resistance to water loss. Therefore, T. ruralis is likely highly flammable under dry conditions. FIRE REGIMES : Find fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find Fire Regimes". POSTFIRE REGENERATION STRATEGY : Initial off-site colonizer (off site, initial community)
Secondary colonizer (on- or off-site seed sources)


SPECIES: Tortula ruralis
IMMEDIATE FIRE EFFECT ON PLANT : Rangeland fires can severely damage all components of soil crusts, including T. ruralis [22]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : No information was available on this topic. PLANT RESPONSE TO FIRE : Information on the response of T. ruralis to fire was sparse. One study in Scotland reported that T. ruralis was present in burned heaths within the first few years following fire [20]. Following a 1975 fire in a shadscale (Atriplex confertifolia)-black greasewood (Sarcobatus vermiculatus) community in Camp Floyd State Park, Utah, an unnamed Tortula species was absent from burned sites for at least 7 years. The species had an average cover in unburned controls of 7.2 and 6.1 percent in 1980 and 1982, respectively [23]. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : No information was available on this topic. FIRE MANAGEMENT CONSIDERATIONS : No information was available on this topic.

References for species: Tortula ruralis

1. Anderson, David C.; Harper, Kimball T.; Holmgren, Ralph C. 1982. Factors influencing development of Cryptogamic soil crusts in Utah deser deserts. Journal of Range Management. 35(2): 180-185. [5498]
2. Anderson, David C.; Harper, K. T.; Rushforth, S. R. 1982. Recovery of cryptogamic soil crusts from grazing on Utah winter ranges. Journal of Range Management. 35(3): 355-359. [5304]
3. Bernard, Stephen R.; Brown, Kenneth F. 1977. Distribution of mammals, reptiles, and amphibians by BLM physiographic regions and A.W. Kuchler's associations for the eleven western states. Tech. Note 301. Denver, CO: U.S. Department of the Interior, Bureau of Land Management. 169 p. [434]
4. Bonham, C. D.; Cottrell, T. R.; Mitchell, J. E. 1991. Inferences for life history strategies of Artemisia tridentata subspecies. Journal of Vegetation Science. 2(3): 339-344. [16599]
5. Brassard, G. R. 1971. The mosses of northern Ellesmere Island, arctic Canada. Bryologist. 74: 234-281. [21551]
6. Brotherson, Jack D.; Rushforth, Samuel R.; Johansen, Jeffrey R. 1983. Effects of long-term grazing on cryptogram crust cover in Navajo Nationa National Monument, Ariz. Journal of Range Management. 36(5): 579-581. [21581]
7. Clarke, G. C. S.; Duckett, J. G., eds. 1979. Bryophyte systematics. New York: Academic Press. 582 p. [21552]
8. Conard, Henry S. 1956. How to know the mosses and liverworts. Dubuque, IA: Wm.C. Brown Company Publishers. 226 p. [9927]
9. Crosby, Marshall R.; Magill, Robert E.; Bauer, Cheryl R. 1992. Index of mosses: 1963-1989. Monographs in Systematic Botany Volume 42. St. Louis, MO: Missouri Botanical Garden. 646 p. [21063]
10. Crumm, H. A.; Anderson, L. E. 1981. Mosses of eastern North America. New York: Columbia University Press. 663 p. [21553]
11. Crum, Howard A.; Steere, William C.; Anderson, Lewis E. 1973. A new list of mosses of North America north of Mexico. Bryologist. 76: 85-130. [21580]
12. Daubenmire, R. 1970. Steppe vegetation of Washington. Technical Bulletin 62. Pullman, WA: Washington State University, College of Agriculture, Washington Agricultural Experiment Station. 131 p. [733]
13. Dhindsa, Rajinder S. 1985. Non-autotrophic CO2 fixation and drought tolerance in mosses. Journal of Experimental Botany. 36(167): 980-988. [20479]
14. Eckert, Richard E., Jr.; Peterson, Frederick F.; Emmerich, Fay L. 1987. A study of factors influencing secondary succession in the sagebrush [Artemisia spp. L.] type. In: Frasier, Gary W.; Evans, Raymond A., eds. Proceedings of the symposium: "Seed and seedbed ecology of rangeland plants"; 1987 April 21-23; Tucson, AZ. Washington, DC: U.S. Department of Agriculture, Agricultural Research Service: 149-168. [3544]
15. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905]
16. Flowers, S. 1973. Mosses: Utah and the West. Provo, UT: Brigham Young University Press. 567 p. [21554]
17. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; [and others]. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. [998]
18. Grout, A. J. 1903. Mosses with a hand-lens. New York: Grout, A. J. 416 p. [21555]
19. Hironaka, M.; Fosberg, M. A.; Winward, A. H. 1983. Sagebrush-grass habitat types of southern Idaho. Bulletin Number 35. Moscow, ID: University of Idaho, Forest, Wildlife and Range Experiment Station. 44 p. [1152]
20. Hobbs, R. J.; Gimingham, C. H. 1984. Studies on fire in Scottish heathland communities. II. Post-fire vegetation development. Journal of Ecology. 72: 585-610. [19767]
21. Holden, Constance, ed. 1992. Miracle moss. Science. 257: 322. [19697]
22. Johansen, Jeffrey R.; St. Clair, Larry L. 1986. Croptogramic soil crusts: recovery from grazing near Camp Floyd State P Park, Utah, USA. The Great Basin Naturalist. 46(4): 632-640. [21579]
23. Johansen, Jeffrey R.; St. Clair, Larry L.; Webb, Bruce L; Nebeker, Glen T. 1984. Recovery patterns of cryptogamic soil crusts in desert rangelands following fire disturbance. Bryologist. 87(3): 238-243. [1264]
24. Gates, Cyndi A.; Tanner, George W. 1988. Effects of prescribed burning on herbaceous vegetation and pocket gophers (Geomys pinetis) in a sandhill community. Florida Scientist. 51(3/4): 129-139. [21582]
25. Kleiner, Edgar F. 1983. Successional trends in an ungrazed, arid grassland over a decade. Journal of Range Management. 36(1): 114-118. [21578]
26. Kleiner, E. F.; Harper, K. T. 1972. Environment and community organization in grasslands of Canyonlands National Park. Ecology. 53(2): 299-309. [6371]
27. Kleiner, Edgar F.; Harper, K. T. 1977. Soil properties in relation to cryptogramic groundcover in Canyonlands National Park. Journal of Range Management. 30(3): 202-205. [21630]
28. Kuchler, A. W. 1964. Manual to accompany the map of potential vegetation of the conterminous United States. Special Publication No. 36. New York: American Geographical Society. 77 p. [1384]
29. Lawton, E. 1971. Moss flora of the Pacific Northwest. [Place of publication unknown]: Hattor: Botanical Laboratory. [Pages unknown]. [21583]
30. Lesica, Peter; McCune, Bruce; Cooper, Stephen V.; Hong, Won Shic. 1991. Differences in lichen and bryophyte communities between old-growth and managed second-growth forests in the Swan Valley, Montana. Canadian Journal of Botany. 69: 1745-1755. [16295]
31. McCune, Bruce. 1977. Vegetation development on a low elevation talus slope in western Montana. Northwest Science. 51(3): 198-207. [21547]
32. Mishler, Brent D.; Oliver, Melvin J. 1988. Evolution of desiccation-tolerance in the Tortula ruralis complex. I. Di Distribution, habits, and water relationships. American Journal of Botany. 75(6/2): 516. [21548]
33. Mishler, Brent D.; Oliver, Melvin J. 1991. Gametophytic phenology of Tortula ruralis, a desiccation-tolerant moss, in the Organ Mountains of southern New Mexico. Bryologist. 94(2): 143-153. [19613]
34. Okoloko, G. E.; Bewley, J. Derek. 1982. Potentiation of sulphur dioxide induced inhibition of protein synthesis by desiccation. New Phytologist. 91(2): 169-175. [20475]
35. Oliver, Melvin J.; Mishler, Brent D.; Quisenberry, Jerry E. 1993. Comparative measures of desiccation-tolerance in the Tortula ruralis complex. I. Variation in damage control and repair. American Journal of Botany. 80(2): 127-136. [20806]
36. Peterson, Janice; Schmoldt, Daniel; Peterson, David; [and others]. 1992. Guidelines for evaluating air pollution impacts on class 1 wilderness areas in the Pacific Northwest. Gen. Tech. Rep. PNW-GTR-299. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 83 p. [20633]
37. Schlatterer, Edward F.; Tisdale, E.W. 1969. Effects of litter of Artemisia, Chrysothamnus, and Tortula on germination and growth of three perennial grasses. Ecology. 50(5): 869-873. [2078]
38. Stanek, Walter. 1980. Vegetation types and environmental factors associated with Foothills Gas Pipeline route, Yukon Territory. BC-X-205. Victoria, BC: Environment Canada, Canadian Forestry Service, Pacific Forest Research Centre. 48 p. [16527]
39. Steele, Alma. 1978. Bryophyte communities of central Idaho forests. Northwest Science. 52(4): 310-322. [21546]
40. Steere, W. C. 1939. Moss flora of North America north of Mexico. Part 4: Tortula. [Place of publication unknown]: Grout, A. J. 264 p. [21556]
41. Tueller, Paul Teuscher. 1962. Plant succession on two Artemisia habitat types in southeastern Oregon. Corvallis, OR: Oregon State University. 249 p. Thesis. [2366]
42. U.S. Department of Agriculture, Natural Resources Conservation Service. 2016. PLANTS Database, [Online]. Available: /. [34262]
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