Mosses of Nevada On-line
Bryophytes From the Northern Mojave Desert

Lloyd R. Stark¹ and Alan T. Whittemore²

1Department of Biological Sciences, University of Nevada, 4505 Maryland Parkway, Box 454004, Las Vegas, NV 89154-4004

ABSTRACT-- An annotated list of bryophytes from the northern Mojave Desert (inclusive of southern Nevada, southwestern Utah, and northwestern Arizona) is provided from a compilation of literature citations, and brought up to date nomenclaturally. Seventy-five species of mosses and five species of liverworts are known from this region, with roughly two-thirds of these known only from Nevada.

Occupying approximately 129,500 km², the Mojave Desert stretches from Joshua Tree National Monument along the Riverside/San Bernardino County Line in southern California, to the vicinity of St. George, Utah, and from Kingman, Arizona to Goldfield, Nevada. About half of the Mojave Desert lies within Nevada, northern Arizona, and southwestern Utah, with the remainder in California (Figure 1). Within the Mojave, more than 90 mountain ranges exist, with the highest point Mt. Charleston, NV (3623 m), and the lowest point in Death Valley, CA (-86 m). It is home to more than 100,000 archeological sites. One-fourth of all seed plant species in the Mojave Desert are regionally endemic (MacMahon 1985). Defining characteristics of the Mojave Desert include internal drainages, well-developed bajadas and playas, precipitation less than 184 mm/yr, and the creosote bush - bur sage plant community [Larrea tridentata (Sessé & Moc. ex DC.) Cov. - Ambrosia dumosa (Gray) Payne] (MacMahon 1985).

A variety of vegetation zones exists in the Mojave Desert, with elevation and annual precipitation largely determining the composition of vascular plant communities. In the lower elevations up to about 1520 m, the typical creosote bush - bur sage community is dominant. At about 1100-1300 m, joshua tree appears, requiring rainfall of 200-250 mm/yr. From 1520 to 2100 m, pinyon - juniper woodland occurs, where precipitation is between 250-450 mm/yr. Many of the ranges support, with increasing elevation, scrub oak woodland, ponderosa pine woodland, quaking aspen forests, white fir/Douglas fir - Rocky Mountain maple woodland, and bristlecone pine woodland (Clokey 1951).

This vast area in the southwestern United States has gone understudied with respect to the bryophytes. Historically, this probably stems from (i) the assumption among botanists that there are very few mosses in the desert, (ii) the difficulty of collecting and detecting moss populations due to heat, dry conditions, and wind, and (iii) the lack of adequate floras and species checklists for the Mojave. Recent studies have implicated desert crustal species (including lichens, bryophytes, cyanobacteria, and algae) and crustal assemblages as central to a variety of community-wide functions. These crustal populations provide the primary source of nitrogen to desert soils, reduce soil erosion, and enhance water retention and seedling establishment (Meyer 1986, Lesica and Shelly 1992, Eldridge 1993, Evans and Ehleringer 1993, Smith et al. 1997). When removed by off-road activities or overgrazing, the soil crust may take decades to centuries to recover, and its presence is a good barometer to overall community health (St. Clair and Johansen 1993, Belnap 1993). This review of the mosses and liverworts reported from the literature focuses on the northern and eastern region of the Mojave, exclusive of California.

While Bartram (1945) described a new species of Weissia from the Virgin Mountains of southern Nevada, the floristic exploration of the region really dates to the state-wide expedition by Elva Lawton (1958). She visited 12 sites in the Nevada Mojave, concentrating in the Spring Mountains (as "Charleston" Mountains, 7 localities). Thirty-three species of mosses were reported from Clark and Lincoln counties. Hepatics were also collected during this expedition, but none were found in the Nevada Mojave Desert (Clark 1957). At 4 of the 12 localities in the Mojave Desert visited by Lawton outside of the Spring Mountains, three or fewer species of mosses were recovered, indicative of the low diversity present in desert areas. An ecological study of the bryophyte and lichen coverage in hot desert ecosystems was conducted on the Nevada Test Site in Rock Valley (Nash et al. 1977). Their transect analysis yielded two additional species of mosses to the region.

Based largely on collections made by Hugh Mozingo of the University of Nevada, Reno, Lavin (1981) reported 20 new state distributional records for Nevada, adding two species of mosses to the known bryoflora of the Nevada Mojave. Subsequently, Lavin (1982) presented the Nevada distributions of all species in the Grimmiaceae, in which he listed nine species of Grimmia from the Nevada Mojave. Of these, three species were added to the known bryoflora of the region. In addition, Lavin (1982) listed three additional species as comprising elements of the bryophyte vegetation in the Spring Mountains near Las Vegas, including the first report of a hepatic from the Nevada Mojave: Targionia hypophylla L. A checklist of mosses from the Intermountain West (Spence 1988) includes all of Nevada and Utah, and portions of Arizona, Colorado, Idaho, and Oregon. Finally, recent taxonomic and ecological studies in the Mojave Desert resulted in several additional species to report from lowland desert regions (Zander et al. 1995, Stark 1996, 1997, Stark et al. 1998).

Citations from Utah were derived primarily from listings from Washington County. Since only part of Washington County lies within the Mojave Desert, future investigations may result in the exclusion of some species from the list below. The approximate boundary of the Mojave Desert in northwestern Arizona coincides with Mohave County, thus Arizona citations were primarily derived from listings from Mohave County. In this regard, two species listed in Spence (1988) as Arizona Mojavean taxa [Pleurochaete squarrosa (Brid.) Lindb. and Entosthodon bolanderi Lesq.] are not listed here since they were reported from counties in Arizona other than Mohave Co. After converting to the presently accepted nomenclature (Stotler and Crandall-Stotler 1977, Anderson et al. 1990, Zander 1993), this results in 80 species of bryophytes. Clearly, since only the Grimmiaceae have been mapped for Nevada, a study of herbarium material at major regional herbaria will doubtless add to the list presented here.

The authors, along with Brent Mishler of the University of California, Berkeley, are currently involved with an inventory of the bryophytes present in the northern Mojave Desert, and plan to discuss noteworthy collections and additions to this region in upcoming reports.

Aloina bifrons (De Not.) Delg. NV, Clark Co., Lime Ridge (Zander et al. 1995).

Amblystegium serpens (Hedw.) Schimp. in B.S.G. var. juratzkanum (Schimp.) Rau & Herv. NV, Clark Co., Spring Mts, Cold Creek (Lawton 1958, as Amblystegium juratzkanum Schimp.).

Anacolia menziesii (Turn.) Par. NV, Clark Co., Spring Mts, Pine Creek Canyon (Lavin 1981); this collection represents the only known locality of the species in Nevada. UT, Washington Co. (Flowers 1973).

Brachythecium collinum (Schleich. ex C.Müll) Schimp. in B.S.G. NV, Clark Co., Spring Mts, Kyle Canyon, Lee Canyon (Lawton 1958), and probably (author did not specify) Pine Creek Canyon (Lavin 1982).

Brachythecium fendleri (Sull.) Jaeg. AZ, Mohave Co. (Haring 1961, as B. utahense James).

B. rutabulum (Hedw.) Schimp. in B.S.G. NV, Clark Co., Spring Mts, Cold Creek (Lawton 1958).

Bryum argenteum Hedw. AZ, Mohave Co. (Haring 1961). NV, Clark Co., Spring Mts, Lee Canyon (Lawton 1958).

B. capillare Hedw. AZ, Mohave Co. (Haring 1961).

B. turbinatum (Hedw.) Turn. NV, Clark Co., Spring Mts, Lee Canyon; Lincoln Co., Ryan-Kershaw State Park (Lawton 1958).

Ceratodon purpureus (Hedw.) Brid. AZ, Mohave Co. (Haring 1961). NV, Clark Co., Spring Mts, Lee Canyon (Lawton 1958).

Climacium americanum Brid. AZ, Mohave Co. (Haring 1961).

Conardia compacta (C.Müll.) Robins. NV, Clark Co., Spring Mts, Cold Creek (Lawton 1958, as Amblystegium americanum Grout).

Coscinodon calyptratus (Hook. in Drumm.) C.Jens. ex Kindb. AZ, Mohave Co. (Haring 1961, as Grimmia calyptrata Hook.). NV, Lincoln Co., Ryan-Kershaw State Park (Lawton 1958, as Grimmia calyptrata Hook.).

Cratoneuron filicinum (Hedw.) Spruce. NV, Clark Co., Spring Mts, Deer Creek Canyon, Cold Creek (Lawton 1958).

Crossidium aberrans Holz. & Bartr. AZ, Mohave Co. (Haring 1961, as C. spatulaefolium Holz. & Bartr.). NV, Clark Co., near Laughlin (Lavin 1981), Lime Ridge (Zander et al. 1995), River Mts (Stark 1997); Nye Co., Rock Valley (Nash et al. 1977).

C. crassinerve (De Not.) Jur. NV, Clark Co., Lime Ridge (Zander et al. 1995), River Mts (Stark 1997).

C. seriatum Crum & Steere. NV, Clark Co., Lime Ridge (Zander et al. 1995).

C. squamiferum (Viv.) Jur. var. pottioideum (De Not.) Mönk. NV, Clark Co., Eldorado Mts, Eldorado Canyon [Lawton 1958, as C. griseum (Jur.) Jur.], Lime Ridge (Zander et al. 1995), River Mts (Stark 1997).

Crumia latifolia (Kindb. in Mac.) Schof. NV, Clark Co., Spring Mts, Cold Creek (Lawton 1958, as Merceya latifolia Kindb.).

Dicranoweisia crispula (Hedw.) Lindb. ex Milde. NV, Clark Co., Spring Mts, Lee Canyon (Lawton 1958).

Didymodon australasiae (Hook. & Grev.) Zand. NV, Clark Co., Lime Ridge (Zander et al. 1995).

D. nevadensis Zand. NV, Clark Co., Lime Ridge (Zander et al. 1995).

D. tophaceus (Brid.) Lisa. AZ, Mohave Co. (Haring 1961). NV, Clark Co., Spring Mts, Cold Creek (Lawton 1958).

D. vinealis (Brid.) Zand. NV, Clark Co., River Mts (Stark 1997). The absence of this common species from the Mojave Desert regions of other inventories (Flowers 1973, Haring 1961) probably reflects the dearth of intensive collecting within the region.

Distichium inclinatum (Hedw.) Bruch & Schimp. in B.S.G. NV, Clark Co., Spring Mts, Deer Creek Canyon (Lawton 1958). This locality may still represent the southernmost known record for the species.

Encalypta intermedia Jur. in Jur. & Milde. NV, Clark Co., Spring Mts, probably Pine Creek Canyon (Lavin 1982).

E. vulgaris Hedw. AZ, Mohave Co. (Horton 1983). NV, Clark Co., Spring Mts, Deer Creek Canyon, Kyle Canyon, and Lee Canyon (Lawton 1958, as E. vulgaris var. mutica Brid.). This report by Lawton was not confirmed in Horton (1983), although this species is to be expected in southern Nevada.

Entosthodon planoconvexus (Bartr.) Grout. UT, Washington Co. (Flowers 1973).

E. wigginsii Steere. UT, Washington Co. (Flowers 1973).

Eucladium verticillatum (Brid.) Bruch & Schimp. in B.S.G. AZ, Mohave Co. (Haring 1961).

Fontinalis antipyretica Hedw. UT, Washington Co. (Flowers 1973).

Funaria hygrometrica Hedw. AZ, Mohave Co. (Haring 1961).

Grimmia affinis Hoppe & Hornsch. ex Hornsch. NV, Clark Co., Valley of Fire State Park (Lawton 1958); two localities in southern Nye Co. (Lavin 1982).

G. anodon Bruch & Schimp. in B.S.G. NV, Clark, Lincoln, and Nye Cos. (Lavin 1982). This species is likely to be the most widespread plant species in Nevada, and the only species of Grimmia to occur in every county of Nevada (Lavin 1982).

G. laevigata (Brid.) Brid. AZ, Mohave Co. (Haring 1961). NV, Clark Co., Newberry Mts (Lawton 1958). UT, Washington Co. (Flowers 1973).

G. orbicularis Bruch ex Wils. NV, Clark Co., Valley of Fire State Park (Lawton 1958), Lime Ridge (Zander et al. 1995), River Mts (Stark 1997); reported from nine additional localities from Clark, Nye, and Lincoln Cos. (Lavin 1982). Within Nevada, this species is restricted to the Mojave Desert.

G. ovalis (Hedw.) Lindb. NV, Clark Co., Spring Mts, probably Pine Creek Canyon (Lavin 1982); Lincoln Co., Ryan-Kershaw State Park (Lawton 1958).

G. pulvinata (Hedw.) Sm. AZ, Mohave Co. (Haring 1961). NV, Clark Co., Spring Mts, probably Pine Creek Canyon, and two additional localities in Clark Co. (Lavin 1982); Lincoln Co., Ryan-Kershaw State Park (Lawton 1958).

G. alpestris (Weber & Mohr) Schleich. NV, Clark Co., Spring Mts, Kyle Canyon (sensu Munoz 1997; Lawton 1958).

G. trichophylla Grev. AZ, Mohave Co. (Haring 1961). A related species, G. decipiens (Schultz) Lindb., was reported from Mohave Co. (Haring 1961), but plants were subsequently assigned to G. trichophylla (Lawton 1971).

Hygrohypnum luridum (Hedw.) Jenn. UT, Washington Co. (Flowers 1973).

Hypnum revolutum (Mitt.) Lindb. NV, Clark Co., Spring Mts, Deer Creek Canyon, Kyle Canyon (Lawton 1958).

Jaffueliobryum raui (Aust.) Thér. AZ, Mohave Co. (Churchill 1987).

J. wrightii (Sull. in Gray) Thér. NV, Clark Co., Eldorado Mts, Eldorado Canyon; Newberry Mts (Lawton 1958, as Grimmia wrightii Aust.). In Nevada, known only from the Mojave Desert (Lavin 1982, Churchill 1987).

Leptodictyum humile (P.Beauv.) Ochyra. NV, Clark Co., Spring Mts, Cold Creek [Lawton 1958, as L. trichopodium (Schultz) Warnst.].

Mnium arizonicum Amann. NV, Clark Co., Spring Mts, Deer Creek Canyon (Lawton 1958).

Orthotrichum alpestre Hornsch. in B.S.G. UT, Washington Co. (Vitt 1973).

O. anomalum Hedw. AZ, Mohave Co. (Vitt 1973). UT, Washington Co. (Vitt 1973).

O. cupulatum Brid. NV, Clark Co., Spring Mts, probably Pine Creek Canyon (Lavin 1982). UT, Washington Co. (Vitt 1973).

O. hallii Sull. & Lesq. in Sull. AZ, Mohave Co. (Vitt 1973). NV, Clark Co., Spring Mts, near base of Lee Canyon (Lawton 1958, but not mapped in Vitt 1973). UT, Washington Co. (Vitt 1973).

O. laevigatum Zett. AZ, Mohave Co., Virgin Mts, [Vitt et al. 1971, as O. laevigatum fo. macounii (Aust.) Lawt. & Vitt in Lawt.]. NV, Nye Co. (Vitt 1973, although from habit information the report may be from the Spring Mts of Clark Co.). UT, Washington Co. [Vitt 1973, as O. laevigatum fo. macounii (Aust.) Lawt. & Vitt in Lawt.].

O. pellucidum Lindb. NV, Clark Co., Spring Mts, Deer Creek Canyon (Lawton 1958, as O. jamesianum Sull.).

O. pumilum Sw. UT, Washington Co. (Flowers 1973).

O. rupestre Schleich ex Schwaegr. AZ, Mohave Co. (Vitt 1973). UT, Washington Co. (Vitt 1973).

Plagiomnium cuspidatum (Hedw.) T.Kop. AZ, Mohave Co. (Haring 1961, as Mnium cuspidatum Hedw.).

Platygyrium fuscoluteum Card. AZ, Mohave Co. (Haring 1961).

Pseudocrossidium crinitum (Schultz) Zand. NV, Clark Co., Valley of Fire State Park (Lawton 1958, as Tortula aurea Bartr.). This collection represents the only known locality for the species in Nevada, and also the only report from the Mojave Desert as a whole (Spence 1987).

Pseudoleskeella tectorum (Funck ex Brid.) Kindb. in Broth. NV, Clark Co., Spring Mts, Deer Creek Canyon, Kyle Canyon, Lee Canyon; Lincoln Co., Ryan-Kershaw State Park (Lawton 1958).

Pterygoneurum ovatum (Hedw.) Dix. AZ, Mohave Co. (Haring 1961). UT, Washington Co. (Flowers 1973).

P. subsessile (Brid.) Jur. UT, Washington Co. (Flowers 1973).

Schistidium apocarpum (Hedw.) Bruch & Schimp. in B.S.G. AZ, Mohave Co. (Haring 1961, as Grimmia apocarpa Hedw.). NV, Clark Co., Spring Mts, Kyle Canyon, and probably Pine Creek Canyon (Lavin 1982, as Grimmia atricha C.Müll. & Kindb. ex Mac. & Kindb., and again as Grimmia stricta Turn.); Lincoln Co., Ryan-Kershaw State Park (Lawton 1958, as Grimmia apocarpa Hedw.).

Splachnobryum obtusum (Brid.) C.Müll. Reported from the "Virgin Basin, Lake Mead" (Crum and Anderson 1981), which is likely to be in Clark Co., NV near the mouth of the Virgin River.

Syntrichia caninervis Mitt. AZ, Mohave Co. (Haring 1961, as Tortula bistratosa Flow.). NV, Clark Co., Lime Ridge (Zander et al. 1995), Black Mts (Stark et al. 1998); Nye Co., Rock Valley (Nash et al. 1977, as T. bistratosa Flow.).

S. norvegica Web. NV, Clark Co., Spring Mts, Kyle Canyon [Lawton 1958, as Tortula norvegica (Web. & Mohr) Wahlenb.].

S. obtusissima (C.Müll) Zand. AZ, Mohave Co. [Haring 1961, as Tortula obtusissima (C.Müll.) Mitt.]

S. princeps (De Not.) Mitt. UT, Washington Co. (Flowers 1973, as Tortula princeps De Not.).

S. ruralis (Hedw.) Web. & Mohr. AZ, Mohave Co. [Haring 1961, as Tortula ruralis (Hedw.) Gaertn. et al.]. NV, Clark Co., Black Mts (Stark et al. 1998).

Timmia megapolitana Hedw. NV, Clark Co., Spring Mts, Deer Creek Canyon, Kyle Canyon (Lawton 1958).

Tortella tortuosa (Hedw.) Limpr. NV, Clark Co., Spring Mts, Deer Creek Canyon, Kyle Canyon (Lawton 1958).

Tortula acaulon (L. ex With.) Zand. UT, Washington Co. (Flowers 1973, as Phascum cuspidatum Hedw.).

T. atrovirens (Sm.) Lindb. NV, Clark Co., Black Mts (Stark et al. 1998). This species was formerly known as Desmatodon convolutus (Brid.) Grout.

T. inermis (Brid.) Mont. NV, Clark Co., Newberry Mts (Lavin 1981), Black Mts (Stark et al. 1998). UT, Washington Co. (Flowers 1973).

T. mucronifolia Schwaegr. UT, Washington Co. (Flowers 1973).

T. nevadensis (Card. & Thér.) Zand. AZ, Mojave Co. (Flowers 1973, as Pottia nevadensis Card. & Thér.).

Trichostomum sweetii (Bartr.) Stark. NV, Clark Co., Virgin Mts, Yant Pit Canyon (Bartram 1945, as Weissia sweetii Bartr.), foothills of Black Mts, River Mts (Stark 1996).

Asterella californica (Hampe) Underw. AZ, Mohave Co. (Evans 1917).

Mannia californica (Gottsche) Wheeller. AZ, Mohave Co. (Flowers 1961). UT, Washington Co. (Flowers 1961).

Marchantia polymorpha L. AZ, Mohave Co. (McCleary 1961). UT, Washington Co. (Flowers 1961).

Riccia sorocarpa Bisch. UT, Washington Co. (Flowers 1961).

Targionia hypophylla L. AZ, Mohave Co. (Evans 1917). NV, reported without precise locality from southern Nevada (Lavin 1982, as Targonia heterophylla [sic]). UT, Washington Co. (Flowers 1961, Schuster 1992). The systematics of Targionia in the Mojave Desert of California and Nevada is currently under evaluation by the second author. It appears likely that populations of Targionia in the Mojave Desert are not referable to T. hypophylla, but rather represent an undescribed species.

We thank Bruce Allen of The Missouri Botanical Garden for continued assistance with this project, Teri Knight of The Nature Conservancy, Las Vegas, Wesley Niles of UNLV, and Kerry Heise, formerly of the U.S. Forest Service, for helpful information, and Robin Stark for preparation of figure 1. A portion of this research was funded by the National Geographic Society (#5429-95), and the senior author acknowledges a New Investigator Award through the University of Nevada, Las Vegas.

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( Figure 1. The Mojave Desert of Nevada, Arizona, and Utah (major mountain ranges indicated). )

This page is part of the
Mosses of Nevada On-line
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Dr. Lloyd Stark
Plant Ecologist, Bryologist and Assistant Professor
University of Nevada at Las Vegas
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James R. Shevock
California Academy of Sciences, San Francisco
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