Great Basin Naturalist

Volume 53 Number 1 Article 11

4-2-1993

Full Issue, Vol. 53 No. 1

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VOLUME 53 NQN I1 MARCH 1993

BRIGHAM YOUNG university

qmwqmp GREAT BASIN naturalist editor JAMES R BARNES 290 MLBM Brigbrighamliamllamilam young university provo utah 84602

associate editors MICHAEL A BORERSBONVERS BRIAN A malberMALRERMAURER blandy experimental farm university of department ofot zoology bnghainbrigharnBrigharn young university virginia box 171755 boyce virginia 22620 provo utah 84602 J R CMIAIHNCALLAHAN JIMMIEJIMMIF R PARRISH museum of southwestern biologybloBiolog109 universitvuniveisitvuniversity of BIOWESTBIO WEST inc 1063 west 1400 north logan new mexico albuquerque new mexico utah 84321 mailing address box 3140 hemethernet california 92546 PAUpulttueilerL T TU E LLE R department of range wildlife and forestry JEFleejeeJEANNE C CHAMBERS unnersitvuniversitvuniversity of nevada reno 1000 valleyvailey roa USDA forest sebleesen iceieelee research umversituniversity of ne- reno nenevadaada 89512 vada reno 920 valley road reno nevada 89512 ROBERT C WIIITMOREWHITMORE JEFFREY R JHASEJOHANSEN division of Forestforestryrv box 6125 west virginia unium department of biologybloBiolog109 john carrolcarroicarroll university versitversiensi Morganmorgantowntown west virginia 26506 6125 university heights ohio 44118 pulPUIPAUL C MARSH center for environmental studies anonaarizona state universituniversiaUniveruniversitysit 7 tempe anonaarizona 85287

editorial board richard W bainnannbaurnannbaurmannBainBaurnann chairman zoology H duane smith zoology clayton M white zoology jerianlenlerljenjerijerllerlan an T flinders botany and range science william hess botanabotanvbotariv and range science all are at brigham young university ex officio editorial board members include clayton S huber dean college of biological and agricultural sciences norman A darais director university publications james R bamesbarnes editor great basin naturalist

the great basin naturalist foundedhounded in 1939 is published quarterly by brigham young university unpublished manuscripts that further our biological understanding of the great basin and surroundingsurroundingareasareas in v estemwestern north america are accepted for publication subscriptions annual subscriptions to the great bainbasin naturalist for 1993 are 25 for individual subscribers 15 for student and erneeventuserneritusementusritus subscriptions and 40 for institutions outside the united states 30 20 and 45 respectively the price of single issues is 12 all back issues are inm print and available for sale all matters pertaining tot subscriptions back issues or other business should be directed to the editor great basin naturalist 290mlbm290igo MLBM brigham young universityumaumv ersityerdity provo UT 84602 scholarly exchanges libraries or other organizations interested in obtaining the great basin natu ralistcalist through a continuing exchange of scholarly publications should contact the exchange libllibrarianLibi anan harold B lee library brigham young universityunixuniv ersitv provo UT 84602 editorial production staff joanne abel technical editor jan spencer assistant to the editor natalie miles production assistant

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VOLUME 53 31 maroMARCMARCH H 1993 no I1 great basin naturalist 531331531 appp 111 41 introduction TO THE SYMPOSIUM ON SOIL CRUST communities limlarnlard L st clair and jeffrejeffrey R johansen 2

ABSTRYABSIRUTABSIR UTr soisolsoilsoli crust communities in semiaridseiniaridarld and andarid lands around theworldthe world haehave recededreceived increasing attention in the past totwot wo decades A symposium on their ecology was presented at thetlletile annual inelnemeetingeting otof the american bryological and lichenological society held in 1991 in san antonio texas an introduction to the topic and an overviewbenieoenieoebienieble of the papers appearing in the proceedings volume are gienglengiven inm this prologue

keykdjilordnordswords cryptoaiiitccryptogainic enistscrustscrnsts miliohioticinicrobiotic crusts inicianicimicrophyticop1njtic crusts tinitinianclsemiaddanciancl ecosystems alacdiacalgae crust lichen anistcnistcnisf mossMIOSS crust

in many arid and semiarid regions of the ferns and fern allies not components of soil world the surface soil is consolidated into a thin crusts and excludes cyanobacteria and fungi crust by microorganisms lichens and bryo not plants mieroMicromicronoralmicrofloralnorainoralfloral crust loope and gif- phytespaytes soil crusts have varying iiiierotopogramicrotopogra ford 1972 microphticmicrophvticmicrophytic crust west 1990 mi phy some are flattened polygonal and possess crocrobioticbiotic crust and cryptocryptobioticbiotic crust belnap a rough undulating surface others are 1993 are other epithets that have been pro- pediceled all soil crust communities contain a posed cryptogamicCrypto garnic crust will likely persist for microflora of cvanobacteriacyanobacteria bacteria eu some time because of its wide usage and histori- karyotickarykaryotinotic algae aidaldand fungi well developed soil cal precedence of the more recent terinsterms we crusts also contain lichens andor bryophytes feel that microbiotic crust is the most accurate soil crusts of biotic origin have been known and recommend its usage under a variety of names raincrustRaincrust was one of increasing evidence indicates that microbimicrobe the first terms used fletcher and martin 1948 otic crusts play several vital roles in arid and but has been abandoned because of confusion semiarid rangeland ecosystems the most im- with raincrustsraincrusts of nonnonbioticbiotic origin many re- portant role likely is stabilization of soil surfaces searchers designate the crust by its dominant and consequent reduction of soil erosion sup- life form ie algal crust lichen crust or moss port for this hypothesis has been gathered by crust cryptogamic crust a tentenntermn coined by har several workers blackburn 1975 booth 1941 per kleiner and harper 1972 has been the fletcher and martin 1948 loope and gifford most widely used terinterm during the last 20 years 1972 mackenzie and pearson 1979 in some some researchers have been dissatisfied with instances microbiotic crusts improve seedling the term cryptogamic crust because cryptogamscryptocryptogramsgams establishment by providing moist sites in the are plants without seeds a group that includes cracks and complex topography of the crusts

departmentent 413ota413of botimbocimota and rdiltyrang sheswscienceselencee Brigbnghainbrighainhain dolingyoung Unkerlniveisihsity prooprovo utah s4n2 2 dpatapat swe 4r2 dparfinentepartiikntolofffiolopBIHbig john carroll lnixcrsihunixuniv sity Univeruimcrsihuniversitiuniversitysittsiti heightsI1 ohio 44118441 IS

I1 2 greafGREATRE vr BASIN naturalist volumeVoluine 53

eckert et al 1986 st clair et al 1984 soil soils with high electrical conductivitynvereconductivity wereNvere moremope have effects on infiltration and it likely to develop visible crust features A few crusts varying71 is unclear whether they improve or worsen workers have studied recovery ofofcyanobacteriacanocanobacteriaeano bacteria waterwaiter relations in the soil harper and marble lichens and mosses following disturbance an- 1988 west 1990 derson harper and rusliforthruslifortli 1982 johansen probably thetiie second most important role and st clair 1986 johansen et al 1982 1984 microbiotic crusts play is the improvement of these studies indicate that algal recovery occurs

soil fertility both the freeree living and lichenized before lichen and moss recovery I1 and that the cvanobacteriacyanobacteria fix atmospheric nitrogen in sicrsacrsig process of full reconeiarecovery takes many vearsyears st nificantnificant aniamountsounts leffjeffjeffriesfiesmies et al 1992 klubek clair et al 1986 demonstrated that recovery and skujins 1980 rychert et al 1978 west and can be accelerated through addition of crapcryp skujins 1977 furthermore the crusts contrib- crainictogamictotocrainic amendments despite these studies ute to soil organic matter through prinprimaryiarylary pro our current understanding of recovery of micro ducthiducthitvductilityductivitytv of the evanoevanobacteriacyanobacteriabacteria and algae jeffjeffriesfiesmies biotic crusts is very limited et al in cresspress tfiroiurhthrough contributions oforganic microbiotic soil crusts of andaridanidannd and semiarid material and reduced erosion of silts and clays rangelands have received considerable atten cation exchange capacity may be higher in tion in the literature thevhavethey have been the subject crusted soils of no fewer than six reviews in the past four microbiotic crusts in north america are most eaisvearsyearsears dunne 1989 harper and marble 1988 prevalent in the semiarid steppe regions in the isichei 1990 johansen 1993 metting 1991 great basin colorado plateau and columbia west 1990 in the past 50 nearsvearsyears over one hun- basin thesthevchevthey also extend into the hotter more dred papers have been published on various andaridarndannd desertdeserts in the southwestern regions of the aspects of the composition distribution physiphaysi united states these regions differ distinctly cal properties and ecology of aridsenarideenaridsemiaridarndannd semiaridanidanndiiarid soil a from semiarid regions east of the rocky moun- crusts there is growing consensus among7 tains in that they developed without the pres- researchers that these crusts play inan important sure of large herds of grazing ungulaungulatesungulatedtes ie beneficial role in thetlletile ecosystems in which they bison antelope mule deer and elk grazed the occur however unanimity does not exist in semiarid steppes before the arrival of european particular west gutknecht 1991 west 1990 settlers but these herbherbivoresivoresivores did not graze in has questioned the ecological value of microbimicrobeofinicrobi large herds and grazed semiarid areas only dur- otic crusts and called for more rigorous studies ing the colder months of the year the domi- of their ecology physical properties and re nance ofbuncligrassesofbunchgrasses shrubs and microbiotic spensesponse to disturbance crust communities in the intermountain west because of the interest in microbiotic crust reflects the historical grazing pressures present communities and the current debate over their in the region mack and thompson 1982 role in semiarid and andaridanidannd ecosystems a sympo- with the introduction of grazing cattle and sium on soil crust communities was held at the sheep vascular and microbiotic communities annual niemeetingeting ofthe american bryological and have both been impacted evidence indicates lichenological society in san antonio texas 5 that domestic grazing anianimalsaninialsnials seriously damage august 1991 this issuelisuelisne of the great basin the integrity of the microbiotic crust through Naturanaturalistfithitfht contains the proceedings of that ssym-ui traintramplingpling of the crust particularly during dry posium asaswellwellweliweil as other related papers on micron periods of the yearvearyear anderson harper andana biotic soil crusts Holniholmgrengren 1982 anderson haihalharperhalperper and rush the first part of the volume contains various forth 1982 brotherson et al 1983 harper and papers dealing with floristics and distribution of marble 1988 destruction of microbiotic crusts various crust components st clair et al 1993 by off road vehicles and backpackers has deewreewreredendenieNdev the distribution of soil lichens in the arid recently become a concern in many areas and semiarid regions ofthe intermountain west rangefiresRangefires also dandamagelage microbioticinicrobiotic crust coniconleomcom of the united states and address the impact of inmuniunitiesties by killing most of the lichen moss and disturbance on soil lichen communities in this algal constituents johansen et al 1982 1984 region A report on the brvoplintesbryophytes of calcareousofcalcareous factors influencing the development of iimi-ii soils of a semiarid region of south central aus crocrobioticbiotic crusts were studied by anderson har tralla is given by downing and selkirk 1993 per and holmgrenHolnigren 1982 they foufoundnd that silty grondin andjohansenand johansen 1993 give apreliniinarvapreliminary 199319931 introduction TO THE SYMPOSIUM 3

great species list of soil algae from colorado national of southern central australia basin naturalist monument and discuss the spatial heterogene- 531353 13 23 DLNFDUNNE J 1989 Cryptocryptogarniccrvptogainicgarnic crusts in and ecosystems ity of cyanobacteria and coccoid eukaryotic al- rangelands 11 iso180184 182 gae in crusts of the monument wheeler et al ECKERT R LE JR F F PETERSONPETLRSON M S meurisseMEURISSF AND 1993 in a companion paper to the work by J L STEPHENS 1986 E ffectseffects ofsodofsowof soil surfacesurfaceaee morpholmorphos grondin and 1993 on the agyogy on emergence and survival of seedlings in big johansen report sagebrush communities journal of range manage spatial heterogeneity of bacterial populations in mentment3939 414 420 microbiotic crusts finally belnap and gardner ELDRIDGEELDRIDCF D J 1993 Cryptocryptogamscryptogramsgams vascular plants and 1993 present an electron microscopical study soil hydrological relations somesorne preliminary results of microcoleusMicrocoleus natus bauchvauchVauch gomont from the semiarid woodlands of eastern australia vaginatusvagi great basin naturalist 53 484858483858 which role that Micromicrocoleuscoleus plays elucidates the FLETCHERfl FTCHFR J E ANDNDVPNV P MARTIN 1948 some effects of in the microstructure of microbiotic crusts algae and moulds in the rain crust of desert soils the second part of the volume addresses the ecology 29 95 100 ecology of the microbiotic crusts our under- GRONDIN A FE AND J R JOHANSFJOHANSEN 1993 microbial spatial heterogeneity in microbiotic crusts in colorado standing of the relationship between microbi- national monument I1 algae great basin naturalist otic crusts and soil hydrology is furthered in a 53 24 30 study of semiarid woodlands of australia by gutknechtGUIKFCHT K W 1991 desert crusts irreplaceable ve eldridge 1993 contributions of microbiotic neer or ecological frosting utah science 52244522 44 46 HARPFR K T YDAND R L PENDLETONPFNDLFTON 1993 cyanobacteriacyanobactena crusts to soil fertility are discussed by harper and eyadobacterialcyanobactenal lichens can they enhance availavailabilabil and pendleton 1993 who present data indicat- ltv of essentialofessential minerals for higher plants great basin ing that cyanobacteria and cyanobacterial li- naturalistKatura list 53 59 72 A for chens may enhance the availability of several HARPERHARPFK K T ANDndjnadjJ R MARBLE 1988 role nondasnonvas cular plants in management of and and remlandsemiarid range essential minerals for higher plants johansen et lands pages 135 169 in PR T tueller ed application al 1993 present the results of a study of the of plant sciences to rangeland management and inven- impact ofrangefireofrangefirerangefirehirebire on soil algal communities in tory kluwer academic publishers boston the columbia basin and the degree of recovery ISICIIEI A 0 1990 the role ofalgaeof algae and c anobactenacyanobacteria in and lands A andaridarld sodsoilsoll research and rehabdirehabih following fire final review during the two years in the tationtationlon 4 1 17 paper belnap 1993 discusses the use ofofinocuinocuinoch JEFFRIESJEFFRIFS D L J M KLOPATFK S 0 LINK ADAND II11 BOI lants in speeding recovery of microbiotic crusts TON 1992 acetylene reduction of cryptogamic crusts from a blackbrushblackbrush community as related to resatura dehydrationtiondehvdrationtiontiondehydration sodsoilsoll biology and biochemistry 24 CITED 1101 ibaib511051 literature JEFFRIESJEFFRIFS D L S 0 LINK ANDNDJJ M KLOIATEKKLOPATEK inpressingressin press coa fluxes of cryptogamic crusts I1 response to re ANDERSON D C K T HARPERHAKPER AND R C HOLMGRENHOLNIGREN saturation new phytolophytologist9ist 1982 factors influencing development of cryptogamic JOJOIIYNSFNI1 A N S E N J R 1993 cryptogamic crusts of semiarid and sod crusts in utah deserts journal of range manage- and lands of north america journal of phycology 28 ment 35 iso180184 185 139147139 147 ANDERSONANDFRSON D CCKK T HARPFRHARPERnarper AND S R rushforthRUSHFOHIH JOHANSEN J R J ASIILFASHLEY ANDD XV R RAYBURN 1993 1982 recovery of cryptogamic soil crusts from grazing r ffectseffects of rangerangefirefire on soil algal crusts in semianasemiandsemiarid on utah winter ranges journal ofrangeorrangeof range management shrub steppe of the lower columbia basin and their 3535535 355359355 359 subsequent recovery great basin naturahstnaturalist 53 73 88 BELNAP J 1993 recovery rates of cryptocryptobioticbiotic crusts inm JOHANSEN J R A JAVAKUL ANDANDSS R rushfortilrusiifortii 1982 oculantosculant use and assessment methods great basinbasmbaem effects of burning on the algal communities of a high naturalist 53 89 95 desert soil near wallsburgWalls burg utah journal of range BELNAP J ANDTD J S cardnergandnerGARDNERGARDNFR 1993 soil microstructure management 35 598 600 in soils of the colorado plateau the role of the cyanoayano JOHANSENJOIIANSFN J R AND L L slsr CLAIR 1986 cryptogamic bacterium Micromicrocoleusmicrocoleufcoleus vagivaginatusnatus great basin natu- soil crusts recovery from grazing near camp floyd ralist 53 40 47 state park utah USA great basin naturalist 46 BLACKBURNBLACKBURM W H 1975 factors influencing infiltration 632 640 rate and sediment production of semiarid rangelands JOHANSEN J R L L ST CIAIRCLAIR B L WEBB ANDDGG T in nevada water resources research 11 929 937 NFBEKER 1984 Recoreeorecoveryervery patterns of cryptogamicoferyptogainic soil BOOTH W E 1941 algae as pioneers in plant succession crusts in desert rangelands following fire disturbance and their importance in erosion control ecology 22 bryologist 87 238 243 38 46 kleiserKLEINERKLFINER hE FANDF ANDKKTT HARPERHARPFR 1972 enenvironmentoronmentandand canyon brotherson J DQ S R RUSHFORTH AND J R JOHAN community organization in the grasslands of SSENE N 1983 effects of longtermlong term grazing on cryptogam lands national park ecology 53 299 309 crust cover in navajo national monument arizonaanzona klKLUBEKUBEK B ANDandiANDJJ skuliskullSKUJISKUJINSNS 1980 heterotrophic nitrogen journal of range management 36 579 581 fixation in andaridarld sodsoil crusts soil biology and biochemisbiochemicBiochemis DOWNING A J AND P SELKIRKSFLKIRK 1993 bryophytes on the try 12 229 236 calcareous soils of mungo national park an arid area LOOPF WLW L AND G F GIFFORD 1972 influence ofaof a soil 4 creatGREATGRFAT BASIN naturalist volume 53

micromicrofloralniieroflora1floral crust on select properties of soils undertinder mountain area of the western united states great pinvonpingon juniper in southeastern utah journal of soil basin naturalist 53 5 12 andnd waterV ater conseiiationconlen atlon 27 164 167 ST CLIRCLAIR L L J R johseJOHSFJOIJANSFN ANDivdISD B L xn7fbbIBBI BB 1986 mkMACK R N ANDVND J N THOMPSON 19s21982 solutionevolution in rapid stabilizationstabistabl dationLation of fire disturbed sites using a soil steppe mthwith feuheulewfew large hoohooedhobedved inaimnalsitiainmals american crust slumslugslurry inoculation studies reclamation and naturalist 119 757 773 revegetation research 4 261 269 mackenziemackenzifMACKLNIF H JANDJ D HWH W PEARSONpi RSO 1979 preliminaryPreliprellminan sr CIIKCLAIR L L B L webbWFBB J R JOHASFJOHANSEN AMAND G T studies on the potential use of algae in the stabilization nebekerNEBEKFR igaign vptogainiccicniptogamic soil crusts enhance ofot sand xwistesastes andarid wind blowbiowblobio situations british phyph ment of seedling establishment in disturbed and undis cological jourjourrialjournalilalilairial 14 126 turbed areas reclamation and revegetation research mettinMFTTINMFITINC B 1991 biological surface features of semiarid 3 124129 136 lands and deserts pages 257 293 in J skujins ed wesrWFSFWEST N FE 1990 structure anand function of inicroplixticinicrophvtic semiarid lands and deserts sodsoil resource and reclamai soil crusts in wild ecosystemsecos stems of arid to serniasemiaridrid rre tion marcel dekker inc new york gions advances in ecological research 20 179 223 sa 1977 cycleeycieeyele in RYCIIERTluchi ri R C J SKUJINs D SOHSISORFNSEN ADAND D fobpobFORpor wesi-W s1 n E ANDdjJ SKUJINS the nitrogen evele in ecosystems cIFellFLLLLA 1978 nitrogen fixationfi vation bvby lichens and free livli north american coidcoldeold vilterwinter semidesertseini desertdesentdesenn ecosvstemsecosv steins plantarumplantarium 53.53 ing nuciomicroorganismsorganisms in deserts pages 2030 in N fc oecologia Plantarum 12 455345 53 R west and J skujins eds nitrogen in desert ecosysecosss WHEELERwheel m C C VRV R FLECHTNFHfleoFLEC HTNFH adlANDADJJ JOHASFJOHANSEN terns dowden hutchinson and ross inc 1993 microbial spatial heterogeneity in microbiotic stroudsburgStroudsburg pennpennslaniaslaniasianiasylvan ia crusts in colorado national monument II11 bacteria great 53 31 siST clorCLMRCLCLAIRMR L L J R JOIUSEJOHASEN ANDD S R rushfortilRUSIII ORTII basin naturalist 313939 1993 lichens of sodsoilsoll crust communities in the inter Great Basin Naturalist 53(1), pp. 5-12

LICHENS OF SOIL CRUST COMMUNITIES IN THE INTERMOUNTAIN AREA OF THE WESTERN UNITED STATES

1 3 1 Lany L. St Clair ,2, Jeffrey R. Johansen , and Samuel R. Rushforth

ABsrRACf.-Lichens are commoncomponents ofmicrobiotic soil crusts. Atotalof34species from 17generaare reported from soil crustcommunities throughouttheIntermountainArea. Distribution ofterricolous lichens is determinedbyvarious physical and biological factors: physical and chemical characteristics ofthe soil, moisture regimes, temperature, insolation, anddevelopment and composition ofthevascularplantcommunity. Some species demonstrate abroadecological amplitude while others have a more restricted distribution. All growth forms are represented; however, the vast majority ofsoil crust lichens are squamulose (minutely foliose). Fruticose species are least abundant. In exposed, middle-elevation sites vagrant (detached) species are common. This paper describes and discusses terricolous lichen communities of desert habitats of the intermountain western United States. Effects of various human-related activities including grazing, wildfire, air pollution, and recreation vehicles on soil crust lichens are discussed. Gypsoplaca macrophylla (Zahlbr.) Timdal, a rare squamulose lichen which occurs on gypsifersous soils, was recently collected in Emery County, Utah, and is reported as new to the state.

Key words: lichens, cryptogamic crusts, microbiotic crusts.

Lichens are common components of soil St. Clair and Newberry 1991, Schroeder et al. crust communities. In some habitats lichens ac­ 1975, Shushan and Anderson 1969). However, count for a significant percentage ofthe ground very few studies have dealt directly with soil cover, often stabilizing the soil surface and en­ lichens. Anderson and Rushforth (1976) pub­ hancing soil fertility. Over the last 25 years ex­ lished the onlylist oflichens from desert soils in tensive studies have been undertaken in arid the IntermountainWest. Theycollectedlichens and semiarid western North America in an ef­ from 34 sites in three distinct areas ofsouthern fort to better understand the ecolOgical role of Utah. Most ofthe sites (21) were located in the microbiotic soil crusts. Initial studies described Great Basin. Five were in gypsiferous habitats various biolOgical components (Anderson and inWashington County, while the remaining sites Rushforth 1976, Johansen et al. 1981). Other were located in pristine, open grassy areas in studies consideredvarious ecolOgical aspects of Canyonlands National Park. They reported a soil crust communities (Anderson, Harper, and total of17 species in 11 genera; however, 3 oftlle Holmgren 1982, Brotherson and Rushforth species were saxicolous and 6 ofthe remaining 1983, Kleiner and Harper 1977, St. Clair et al. species were misidentified. Nash and Sigal (1981) 1984, Skujins and Klubek 1978). Human-in­ published a checklist ofthe lichens ofZion Na~ duceddamage to soil crust communites has also tional Park in connection with a preliminary been studied (Johansen et al. 1984, Johansen air-quality survey for the park. They reported a and St. Clair 1986). Some research has investi­ total of 159 species in 53 genera from their gated recovery and reclamation/restoration of collections. Nine ofthe species were terricolous damaged soil crust communities (Anderson, lichens from middle-elevation desert habitats. Harper, and Rushforth 1982, St. Clair et al. Two recent monographic works (Thomson 1987, 1986). 1989, Timdal1986) have addedSignificant taxo­ Several lichen floras and checklists for the nomic andecolOgical information about two of IntermountainAreahave beenpublished (Egan the more abundantsoil generainwestern North . 1972, Nash and Johnsen 1975, Newberry 1991, America (Psora and Catapyrenium). St..Clair and

~DeparbnentofBot.my and Range Science, Brigham Young University, Provo. Utnh 84602­ ... Address for reprints. 3Deparbnent ofBiology, John Carroll University, University Heights. Ohio 44118.

5 6 GREAT BASIN NATURALIST [Volume 53

Wanick (1981) repOlted Acarospora nodulosa ful evaluation of the vagrant lichens ofthe In­ (Duf.) Hue. v. nodulosa, a squamulose soilli­ termountain Area shows that Rogers's conclu­ chen collected from gypsiferqus soils in south­ sion is probably accurate for at least some ern Utah, as a new record fot North America. species. For example, Rhizoplaca haydenii is Timdal(1990) describedanewandraresquami­ commonly collected from habitats in which ad­ form lichen genus and family (Gypsoplaca of jacent rocks are coveredwith the attached, um­ the Gypsoplacaceae) from gypsiferous soils in bilicate lichen R. melanophthalma. This southwestern Colorado. Timdal included with situation has led some lichenologists to seriously his description a list of 21 soil crust species question whether R. haydenii is Simply a de­ commonly associated with Gypsoplaca 111£lcro­ tachedform ofR. melanophthalma. Rosentreter phylla. Many species on Timdal's list are com­ and McCune (1992) have also reviewed the monlyfoundon gypsiferous soils throughoutthe status ofthe vagrant lichen Dermatocarpon va­ Colorado Plateau. Recently, Newberry (1991) gans. After carefully comparing D. vagans with characterized the lichen flora of the Uinta D. reticulatwn and D. miniatum, they con­ Mountains ofnortheastern Utah, listing a total cludedthat thevagrant form is simply detached of291 taxa in95 generafrom his collections. As fragments ofD. reticulatum and D. miniatum. part ofhis study he made extensive collections Similar discussions have takenplace concerning of soil crust lichens along the northern border Agrestia hispida, with some lichenologists (We­ of the Colorado Plateau. Rosentreter and ber1967) suggestingthatA. hispida is simply an McCune (1992) described distribution patterns environmental modification of the attached ofvagrant species ofthe foliose genus Dermato­ crustose lichen Aspicilia calcarea. The only ex­ capon in Idaho, Montana, Wyoming, and east­ ception to this general pattern may be Xantho­ ern Oregon and Washington. parmelia chlorochroa, which at least in some The pmpose ofthis paper is to characterize habitats seems to be a truevagrant with no local the lichen component of desert soil crust com­ attached members. However, in other habitats mtmities in intermountain western North vagrant specimens ofX. chlorochroa occur sym­ America. This paperis basedon a careful review patrically with several speCies of Xanthopar­ ofthe literature as well as unpublished 0 bserva~ melia thatgrowover rocks andonto the soil. The tions and collections made by the authors and issue is somewhat confUSing; however, Rogers's others throughout the Intermountain Area over conclusion is probably correct, at least in some the last 14 years. cases. . Squamulose and crustose lichens dominate RESULTS AND DISCUSSION open Great Basin sites with vagrant forms con~ spicuously absent. The three most common li­ Soil Crust Community Structure chen species from the Great Basin are Collema Most soil crustlichens are eithersquamulose tenax, Catapyrenium lachneum; and Caloplaca orfoliose (19%). Squamulose (minutelyfoliose) tominii (Table 1). Development and distribu­ species are particularlycommon. Ofthe 34 spe­ tionofsoil crustcommunities in the Great Basin cies currently known from soil crust communi­ seem to be correlated with the occurrence of ties in the Intermountain Area, 59% are vascular plants. Intershrub spaces are domi­ squamulose, 21% foliose, 12% crustose, and9% 'natedby cyanobacterial and lichen crusts while fruticose (Table 1). Ofthe 20 squamulose spe­ the area immediately beneath shrubs is domi­ cies, 9 are in the genus Psora. nated by various moss and vascular species. A All fruticose and most foliose species are combination of several factors, including mois­ vagrant (unattached), the only exceptions being ture, insolation, and perhaps even allelopathic Xanthoparmelia wyomingica, which is loosely activity (SchlattererandTisdale 1969), seems to adnate overrockysoils, andX. idahoensis, which be dictating this pattern. is often loosely attached to the soil surface. All Gypsiferous soils have the best developed vagrant forms tend to become entangled either licllen communities, often with 100% lichen with vascular plants orwith detritus, and during cover. Species diversity is also very high at wetperiods they may even become temporarily gypsiferous sites, with several rare species attached to the soil. Rogers (1911) suggested (e.g., Acarospora nodulosa var. nodulosa and that many vagrant forms "are Simply detached Gypsoplaca macrophylla) becoming common fragments ofnormally attached species." Care- to abundant. The most abundant species on 1993] TERRICOLOUS LICHENS 7

TABLE 1. Distribution of terricolous lichen species from the Intermountain Region ofwestern United States. Growth form: Cr = crustose, Fo = foliose, Fr = fruticose, Sq = squamulose. Habit: A = attached, V = vagrant. Relative abundance: A = abundant, C = common, R = rare.

Intermountain Area soil lichen habitats

Growth Great Species form Habit Basin Gypsiferous Steppe Upland

Acarospora nodulosa (Dufour) Hue Sq A C-A Acarospora nodulosa vm: reagens (Zahlbr.) Clauz. & Roux Sq A R-C Agrestia hispida (Mereschk) Hale & Culb. Fr V C-A R-C Aspicilia reptans (Looman) Wehn. Fr V R Aspicilia sp. Fr V C-A Buellia elegans Poelt Sq A R-C C-A R Caloplaca tominii Savicz Cr A C R Catapyrenium daedaleum (Krempelh.) B. Stein Sq A R Catapyrenium laclmeum (Ach.) It Santo Sq A C-A C-A CC Collelrut tenm:. (Swartz) Ach. Sq A C-A C R-C R-C Dennatocarpon miniatllm (L.) Mmm Fo V R-C Dennatocarpon retiClllatum Magn. Fo V R-C Diploschistes diacapsis (Ach.) Lumbsch Cr A A Endocarpon pllsillwn Hedwig Sq A R R Flllgensia desertonun (Tomin) Poelt Cr A C Flllgensia fulgens (Swartz) Elenkin Sq A R-C R-C Gypsoplaca macrophylla (Zahlbr.) Timdal Sq A R-C Psora cerebriformis W Weber Sq A R-C C-A Psora crenata (Taylor) Reinke Sq A C Psora decipiens (Hedwig.) HofEm. Sq A C-A CCC Psora globifera (Ach.) Massal. Sq A R Psora icterica (Mont.) Muell. Sq A C Psora luridella (Tuck) Fink Sq A R Psora 1rumtana TImdal Sq A R Psora mssellii (Tuck) A. Schneider Sq A R Psora tllckennanii R. Anderson ex TImdal Sq A C-A C Rhizoplaca haydenii (Tuck) W Weber Fo V C-A Squamarina lentigera (Weber) Poelt Sq A C-A R Toninia caentleonigricans (Lightf.) Th. Fr. Cr A R-C R-C R Toninia tristis (Th. Fr.) Th. Fr. Sq A R Xanthopannelia chlorochroa (Tuck) Hale Fo V C-A R-C Xanthoparmelia idahoenyiy Hale Fo NV R-C XantllOparmelialipchlorochroa Hale & Elix Fo V C-A XantllOpannelia wyomingica (Gyelnik) Hale Fo A C-A

gypsiferous soils are Diploschistes diacapsis and ent but rare in upland sites. Xanthoparmelia Sqitamarina lentigera (Table 1). Vagrantspecies chlorochroa andAgrestia hispida havebeen col- are also missing from gypsiferous sites. lected from this habitat type. Upland, sandy sites in pinyon-juniperwood- Dry; upland steppe sites dominated byvari- lands are dominated by various species of the ous Artemisia spp. often have well-developed genus Psora, including P. cerebriformis, P. tuck- vagrant lichen floras including Xanthoparmelia ennanni, and P. decipiens. Catapyrenium lach- chlorochroa, Rhizoplaca haydenii, Dennatocar- neum and Endocarpon pusillwn are also found pon reticulatwn, and Agrestia hispida. The atthese sites (Table 1). Vagrantspecies are pres- squamulose species Catapyrenium lachneum 8 GREAT BASIN NATURALIST [Volume 53 and Collema tenax are also commonly found in eastern Montana, northwestern Utah, and this type ofhabitat (Table 1). northeastern Colorado. Other species included Some species of soil crust lichens are both in this association, namely Catapyrenium lach­ broadly distributed and generally abundant, oc­ neum, Endocarpon pusillum, Collema tenax, curring throughout the fi,Ill range of soil crust PSQra decipiens, Toninia caeruleonigricans, Ful­ habitats in the Intermountain Area. For exam­ gensiafulgens, Buellia elegans, and Squamarina ple, the squamulose lichens Psora decipiens and lentigera, also occur in the IntermountainArea. Catapyrenium lachneum are common compo­ Some species occurinboththe Great Basin and nents ofall desert habitats in the Intermountain tlle Colorado Plateau, while others are limited Area, occurring on calcareous, gypsiferous, and to one or tlle other. Other taxa included in sandy upland soils. Theyare also common com­ Looman's Parmelietum chlorochroae association ponents ofalpine tundra habitats throughout in­ (e.g., Cladonia pocillum, Phaeorrhiza nimhosa, termountain North America. Both species have andAcarosporaschleicheri) are absent from the also been collected worldwide from Australia, middle-elevation deserts of tlle Intermountain Africa, Asia, and Europe (Rogers and Lange Areabutare commonly found in intermountain 1972). Otherspecies are broadlydistributedbut boreal and!or alpine tundrahabitats. are much less abundant. Such taxa include En~ Europeanlichenologists have described two docarpon pusillum, Toninia caeruleonigricans, lichen associations, Fulgensietum fulgentis and and Buellia elegans. Gelatinous lichens (e.g., Parmelietum vagantis, which are very similar to Collema sp. and Peccania sp.) are abundant but the soil lichen communities ofthe Great Basin not particularly welllmown from western soil and Colorado Plateau (Klement 1955, 1958). crust communities. More careful analysis ofthis Thecombined species list for the two European group, particularly on gypsiferous soils, will in~ associations is virtually identical to Looman's variably yield several additional species. Pres­ Parmelietum chlorochroae association list. ently, the taxon Collema tenax is probably Floristically, the Fulgensietum fulgentis asso­ overused. ciationis more closelyrelatedto tlle GreatBasin One group oflichens conspicuously missing and Colorado Plateau soil lichen communities, from the intermountain deserts is the genus while Parmelietum vagantis more closely re­ Cladonia. Although this genus occurs com­ sembles the short-grass-shrubland steppe com~ monly on the Great Plains, in the boreal forest, munity of western Wyoming, IdallO, eastern and on tlle alpine tundra, it is not represented Montana, nortlleastern Utah, andnorthwestern by a single species in tlle soil crust communities Colorado. The Intermountain Area seems to ofthe Great Basin or Colorado Plateau. have four distinctive soil lichen associations (Ta­ ble 1): calcareous Great Basin soil crusts, sandy Affinities ofIntermountain Area . Colorado Plateau soil crusts, gypsiferous soil Soil Crust Lichens crusts, and northern steppe soil crusts. Some species are found in all four associations, while Very little descriptive or ecological informa­ other species are unique to a given association tionis available for tlle lichen component ofsoil (Table 1). crust communities in the Intermountain Area. Looman (1964), whose work was the first to Annotated List ofSoil Crust Taxa specifically characterize terricolous lichen com­ munities in NorthAmerica, focused ontlle Prai­ Acaro8pora nodulosa (Dufour) Hue var. rie Provinces of Canada and adjacent parts of nodulosa. A squamulose species locally abundanton the nortllern Great Plains. He specifically de­ gypsiferous soils. This species was reported new to scribed the Parmelietum chlorochroae lichen North America by St. Clair and Warrick in 1987. Original collections of this species were made in association, a steppe community with arctic­ Washington County, Utah, with subsequent collec­ alpine affinities. Floristically, this association is tions from similar habitats in Emery County, Utah, verysimilar to the lichen soil crust communities and northwestern Arizona. ofthe Intermountain Area. This association in~ cludes several vagrant species (e.g., Xanthopar­ Acarospora nodulosa (Dufour) Hue var. reagens. A squamulose lichen collected from Mont­ melia chlorochroa, Agrestia hispida, Rhizoplaca rose and San Miguel counties in south~esternColo­ haydenii, and Aspicilia reptans) that are also rado. Locally common on gypsiferoqs soils. This common. components of the short-grass­ species has not been collected in Utah; however, shrubland steppe ofwestern Wyoming, Idaho, more careful examination ofgypsiferous soil habitats 1993] TERRICOLOUS LICHENS 9 in the state will likely confirm its occurrence in Utah collected from middle- to higher-elevation, calcare­ and perhaps even the northwestern comer of ous sites in Colorado, Wyoming, and Utah. Arizona. Catapyrenium lachneum (Ach.) R. Santo This Agrestia hispida (Mereschk.) Hale & Culh. A squamulose lichen is one ofthe most broadly distrib­ fruticose, vagrant lichen that demonstrates tremen­ uted terricolous species in the Intermountain Area. dous morphological plasticity. This species is part of It commonly occurs on calcareous soils in the Great a complicated group ofrelated species thatshould be Basin, middle-elevation pinyon-juniper sites, gypsif­ carefully compared with similar material reported erous soils oftheColorado Plateau, and alpine tundra from comparable habitats in Russia. Ithas been com­ soils. This species has anincredible ecological ampli­ monly collected from middle-elevation, calcareous, tude and demonstrates substantial morphological shrubland habitats dominated by Artemisia spp. variation. and/or Atriplex spp. in Utah, Wyoming, Idaho, and Collematenax (Swartz)Ach.This isidiate, squa­ Colorado. Weber (1967) suggested this species is an mulose lichen is one ofthe mostcommon terricolous environmental modification of Aspicilia calcarea. lichens of the Intermountain Region. It occurs on Roger Rosentreter's work on the vagrant Aspicilia/ calcareous soils inthe Great Basin and also ongypsif­ Agrestia spp. ofwestern NorthAmericawill undoubt­ erous soils on the Colorado Plateau. This species edly show that there are at least several new species , recovers veryrapidlyfollowing perturbation (e.g., fire records for North America from this group. or grazing disturbance). Its capacity for quick recov­ Aspiciliareptans (Looman) Wetm. Afruticose, ery is likely related to the fact that itproduces abun­ semi-vagrant species collected from middle-eleva­ dant vegetative propagules (isidia) (Johansen et al. tion calcareous sites in Idaho and Wyoming and al­ 1984, Johansen and St. Clair 1986). pine tundra sites in Utah. This species is commonly Dermatocarponminiatum (L.) Mann. This fo­ overlooked because it blends in with detritus that liose, usually attached saxicolous lichen occasionally tendsto accumulateintheabove-mentionedhabitats. becomes detached andoccurs as avagrant onthe soil. . This species represents one extreme ofthe Agrestia­ According to Rosentreter and McCune (1992) most Aspicilia complex, a group requiring serious mono­ vagrant Dermatocarpon species are collected from graphic treatment. poorly drained basalt flats dominated by Artemisia Aspicilia sp. A fmticose, vagrant species col­ rigida in western Idaho and eastern Washington and lected from short-grass-shmbland steppe sites in Oregon. After. careful study Rosentreter and Wyoming and Idaho. This species most closely re­ McCune have determined that vagrant species of sembles Agrestia hispida; however, this taxon has a Dermatocarpon are detached fragments ofeitherD. more substantial central thallus with short, blunt reticulatum or D. miniatum; therefore, they recom­ lobes rather than the densely and finely branched mend that the epithet for the vagrant form of this thalli typical of Agrestia hispida. In addition, A. species (Dermatocarpon vagans Imsh.) no longer be hispida has better developed and more prominent used. pseudocyphellae. Thalli ofAs/Jicilia sp. are also more Dermatocarpon reticulatum Magn. This fo­ compact and tend to be more spherical in shape. liose, usually attached saxicolous species is reported Roger Rosentreter's work comparing North Ameri­ as a vagrant soil lichen from northwestern Wyoming, can members ofthis group with similar Russian taxa western Idaho, and eastern Oregon. See related will likely result in an epithet for this species. dicussion under Dermatocarpon miniatum. Buellia elegans Poelt. A squamulose species Diploschistes diacapsis (Ach.) Lunihsch. A with prominentlobes. This species is broadly distrib­ cmstose lichen commonly collected on gypsiferous uted, reachingits bestdevelopmentin. sites protected soils in southern Utah, southeastern Colorado, and from trampling. Ithas been collected from protected northwestern Arizona. This species is the single most sandy soils in Emery County, Utah, as well as gypsif­ abundant lichen collected from gypsiferous soils in erous soils in Colorado, Utah, and northwestern Ari­ the Intermountain Area. zona, and less commonly from protected calcareous soils in the Great Basin. Endocarp,pn pusillum Hedwig. This squamu­ lose species is broadlydistributedbutnotparticularly Caloplaca tominii Savicz. A cmstose, sorediate common. It is found on calcareous soils in the Great species that occurs commonly on calcareous soils Basin and occasionally on undisturbed, sandy soils in throughout the Great Basin. This species was first pinyon-juniper communities. The fact that this spe­ reported by Nimis (1981) as new to North America. cies is rather uncommon may; at least in some meas­ His collections were from the KIuane region ofthe ure, berelatedtoa relativelylowtoleranceforgrazing Canadian Yukon, an area in some respects strikingly and fire disturbance. similar to the open shmblands ofthe Great Basin. Fulgensia deserlorum (Tomin) Poelt. This Catapyrenium daedaleum (Krempelh.) B. granular-subsorediate lichen occurs commonly on Stein in Cohn. A rather rare squamulose species gypsiferous soils in Arizona, Colorado, and Utah. 10 GREAT BASIN NATURALIST [Volume 53

Fulgensia Julgens (Swartz) Elenkin. A squa­ been reported from Arizona and southwestern mulose species with well-developed lobes. It is fre­ Colorado. quently collected from calcareous soils of the Psora tuckermanii R. Anderson ex Timdal. Colorado Plateau, gypsiferous soils, and less com­ This broadly distributed, squamulose soil lichen has monlyfrom undisturbed calcareous soils in the Great beenreportedfor Arizona, Colorado, Idaho, Nevada, Basin. New Mexico, Utall, and WyOlning. P. tuckermanii Gypsoplaca macrophylla (Zahlbr.) TiIndal. shows extensive morphological variation and is com­ This ratherrare squamulose lichen has recently been monly confused with several other species ofPsora. collected from gypsiferous soils in southwestern Ithas beencommonlycollectedfrom calcareous soils Colorado and was reported as newto North America in the Great Basin as well as on soil over rock on tlle by TImdal (1990). Likely, with additional collections Colorado Plateau. andmore careful analysis ofexistingcollections, other Rhizoplaca haydenii (Tuck.) Weber. A va­ species will be added to this genus. This taxon was vv. grant soil lichen collected from calcareous shmbland also collectedfrom gypsiferous soils ontlle San Rafael habitats in IdallO, Utah, and Wyoming. This species Swell in EmeryCounty, Utah, during the fall of1992 demonsh'ates substantial morphological variation and represents a new species record for the state ranging from robust spherical tllalli (typical oflower­ (BRY C21698). elevation sites) to finely branched flattened thalli Psora cerebriJormis vv. Weber. A robust, squa­ (typical of higher-elevation sites). The genus Rhi­ mulose lichenfrequently collectedfrom pinyon-juni­ zoplaca needs attention andis presentlyunderreview per and shmbland habitat in Arizona, Colorado, by Bmce Ryan. IdallO, Nevada, New Mexico, Utah, and Wyoming. Squa'fTW,rina lentigera (Weber) Poelt. This Psoracrenata (Taylor) Reinke. A squamulose squmnulose soil lichen has prominent lobes. S. len­ lichen with prominent, mm'ginal apothecia. This spe­ tigera is one ofthe more abundant lichens collected cies is rare to common on calcareous and gypsiferous from gypsiferous habitats in nOlthemArizona, south­ soils in Arizona, Colorado, New Mexico, and Utall. western Colorado, and southern Utah. It has also been collected from undisturbed, sandy soils in pin­ Psora decipiens (Hedwig) Hoffm. This squa­ yon-juniper habitat. mulose lichen, along with Catapyrenium lachneum, is the most broadly distributed terricolous lichen in Toninia caeruleonigricans (Lightf.) Th. Fr. A western North America. It has been commonly col­ convoluted, cmstose to squmnulose soil lichen com­ lected from calcm'eous soils in the Great Basin, gyp­ monly collected from calcareous soils in tlle Great siferous soils on the Colorado Plateau, and Basin andColorado Plateau. This species is one ofthe high-elevation alpine tundra sites tllfoughout tlle more broadly distributed soil lichens, occurring in Rocky Mountains. Itoccurs less commonlyon soils in Arizona, Colorado, Idaho, Nevada, New Mexico, pinyon-juniper habitat. Utah, and Wyoming. Psora globifera (Ach.) Massal. A squmnulose Toninia trislis (Th. Fr.) Th. Fr. A convoluted, lichen that generally occurs in fissures ofcalciferous squmnulose soil lichen. This species occurs on cal­ rocks. It also occasionally occurs on calcareous soils careous soils and has been collected rm'ely from pro­ in Arizona, Colorado, IdallO, Nevada, Utah, and tected habitats in pinyon-juniper communites. Ithas Wyoming. been reportedfrom Arizona, Colorado, New Mexico, and Utah. Psora icterica (Mont.) Miill. Arg. A squamu­ lose soil lichen commonly collected in the arid areas Xantlwparmelia chlorochroa.(Tuck.) Hale. A oftlle western United States. In the Intennountain foliose, vagrant lichenthatoccurs abundantly onsoils Areaithas beenreportedfrom Arizona, NewMexico, ofshmbland-steppe communities andless commonly and Colorado. Itoften occurs inpinyon-juniperhabi­ in pinyon-juniper communities. The distribution of tat on sandy; undisturbed soils. this species seems to be positively correlated with grazingimpact (McCracken etal.1983). Itcommonly Psora luridella (Tuck.) Fink. A rare, squamu-. occurs with and has been reported lose soil lichenreportedfrom Colorado, NewMexico, Agrestia hispiCW from Arizona, Colorado, NewMexico, Nevada, Utah, Nevada, and Utah. Because P. luriclella is morpho­ andWyoming. Recently, this group has beensplitinto lOgically very similar to P. globifera, the two species are often confused. several closely related species based on subtle mor­ phological and chemical differences (Hale 1990). Psora montana TIIDdai. A squamulose soilli­ Only one oftlle chemical segregates is includedhere chen occasionally collected below timberline.. It is (X. lipochlorochroa). Furtller evaluation of vagrant reported to occur in Colorado, Utall, and Wyoming. Xanthoparmelias in the Intennountain Area will ul­ timately yield several additional species. Psorarussellii (Tuck.) A. Schneider. This squa­ mulose lichen occurs ill arid areas ofthe southwest­ Xnnthoparmelia idahoensis Hale. Aloosely at­ ern United States. In the Intennountain Area it has tached to vagrant, foliose to sub-fmticose lichen 1993] TERRICOLOUS LICHENS 11 collected from calcareous lacustrine ash soils near. oftlle Great Basin and Colorado Plateau. These Salmon, Idaho. This rare butlocally abundant lichen herds always occupiedlower-elevationsites dur­ has been reported only from the type locality. Exami­ ingthewinterandearlyspringmonths whensoil nation of similar habitats elsewhere should reveal a broader distribution pattern. crusts were wet because of seasonal precipita­ tion and thus less vulnerable to the effects of Xanthoparmelia lipochlorochroa Hale & Elix. A rare, vagrant soil lichen that is a fatty acid trampling. However, as the dller summer chemotype ofX. chlorochroa. The two species inter­ months approached and soil crusts became dry mix in the desert shrublands of southwestern Wyo­ and brittle, and thus more vulnerable to tram­ ming. Careful examination of the chemistry of pling, wild grazing animals moved back into the collections ofX. chlorochroa will likely demonstrate mountainous areas of the region. In contrast, a much broader distribution pattern for X. lipo­ chlorochroa. modem humans have maintained larger herds of domestic· animals in greater numbers and Xnnthoparmelia wyomingica (Gyelnik) Hale. have grazed tlle basin and plateau regions well A loosely attached, foliose lichen common on rocky soils in middle- to higher-elevation sites in Colorado, into the summer months or even continuously. Idaho, Utah, and Wyoming. Hale (1990) indicates Furthennore, modem humans have introduced that this species does not occur below 3000 m and alien vascular plant species that now make it does not occur sympatrically with X. chlorochroa. possible to sustain large wildfires in a region However, I (St. Cl.) have personally collected this where wildfire was not particularly common. species from higher-latitude sites below 3000 m (northern Colorado, Idaho, 1md Wyoming); I have Theendresulthasbeenextensive damage to soil also observed that it often occurs sympatrically with crust communities with a concomitant increase X. chlorochroa in many ofthe higher-latitude sites. in soil erosion and decline in soil fertility. Many species oflichens are sensitive to vari­ Human-related Impact on ous types of air pollutants (Nash and Wirth Soil Crust Communities 1988). Recently; they have been used to bio­ Lichens are impOltant components of soil monitor the effects ofair pollution in protected crust communities in the intermountain west­ habitats such as wilderness areas and national ern United States, especially in areas protected parks (St. Clair 1989). Unfortunately; very little from domestic grazing, wildfire, and off-road is known about the effects of air pollutants on vehicle activity. Soil crusts in general and the soil crustlichens. Itis generallythoughtthat the lichen component in particular tend to be very basicsoils ofthe IntermountainAreaameliorate sensitive to human-related perturbation. These acid~gen­ complex, sensitive communities thrived for the effects of air pollution, especially years prior to the advent of modem humans. erating pollutants. Even though no empirical They provide 40-100% ofthe ground cover in evidence supports this hypothesis, researchers an area with relatively sparse vascular plant have shown that lichens growing on calcareous cover. They also effectively reduce wind and substrates do have a higher tolerance for acid water erosion while Significantly increasing soil pollution. Currently, a study is in progress to fertility. However, over the last 150 years a sig­ evaluate the effects of emissions from a toxic nificant portion ofthe soil crust communities of waste incinerator in central Utall on soil crust the Great Basin and Colorado Plateau has been communities. Baseline community data and heavily damaged, mostly due to intensive graz­ toxic element concentrations from the soil'were ing bycattle and sheep. Soil crust communities obtained prior to operation of the incinerator. are generally slow to recover, often requiring Follow-up studieswill showwhetherornottoxic manyyears for full recovery (Anderson, Harper emissions accumulate in the soil and/or nega­ and Rushforth 1982, Johansen et al. 1984). tively impact soil crust communities. Research Soil crust community structure in tlle Inter­ is also needed to accurately evaluate the effects mountain Area evolved without significant im­ pact from large herds of grazing animals (i.e., of acid precipitation on soil crust communities bison) andwith little or no impact from wildfire in general and the lichen componentinparticu­ (MackandThompson1982). Impactfrom herds lar. Failure to document air pollution effects ofdeer, antelope, and elkwas minimizeddue to could further jeopardize a resource that has the smaller size and number of herds and the already sustained significant damage from in~ time ofthe yeartheyinhabitedsemiaridregions tensive grazing and wildfire. 12 GREAT BASIN NATURALIST [Volume 53

LITERATURE CITED NEWBERRY, C. C. 1991. Lichens of the Uinta Mountains and adjacent intermountain North America. Unpub­ lished master's thesis" Brigham Young University, ANDERSON, D. C., K. T. HARPER, AND R. C. HOLMGREN. Provo, Utall. 236 pages. 1982. Faetorsinfluencing development ofcryptogamic NIMIS, P. L. 1981. Caloplaca tominii newto North America. soil crusts in Utah deserts. Journal of Range Manage­ Bryologist 84: 222-225. ... ment 35: 180-185. ROGERS, R. W. 1977. Lichens of hot ;uid and semi-arid ANDERSON, D. C., K. T. HARPER, AND S. R. RUSHFORTH. lands. Pages 211-252in M. R. D. Seaward, ed., Lichen 1982. Recoveryofcryptogamic soil crusts from grazing ecology. Acadmic Press, NewYork. on Utah winter ranges. Journal ofRange Management ROGERS, R. W.,AND R. T. UNGE.1972. Soilsurface lichens 35: 355--J59. in arid andsubarid south-eastern AustraliaI. Introduc­ ANDERSON, D. C., AND S. R. RUSHFORTH. 1976. The cryp­ tion and floristics. Australian Journal of Botany 20: togamic flora ofdesert soil crusts in Utah deserts. Nova 197-213. Hedwigia~8: 691-729. ROSENTRETER, R., AND B. MCCUNE. 1992. Vagrant Der­ BROTHERSON, J. D., AND S. R. RUSHFORTH. 1983. Influ­ matocarpon in western North America. Bryologist 95: ence ofcryptogamiccrusts on moisture relationships of 15-19. soils in Navajo National Monument, Arizona. Great ST. CLAIR, L. L. 1989. Report concerning establishment of Basin Naturalist 43: 73-78. a lichen biomonitoring program for the Jarbidge Wil­ derness Area, Humboldt National Forest, Nevada. EGAN, R. S., 1972. Catalog of the lichens of New Mexico. U.S. Forest Service Technical Report. 22 pp. Bryologist 75: 7--J5. ST. CLAm, L. L., J. R. JOHANSEN, AND B. L. WEBB. 1986. HALE. M. E. 1990. A synopsis of the lichen genus Xantho­ Rapid stabilization of fire-disturbed sites using a soil pannelia (Vainio) Hale (Ascomycotirta, Parmeliaceae). crust slurry: inoculation studies. Reclamation and Smithsonian Contributions to Botany 74: 1-250. Revegetation Research 4: 261-269. JOHANSEN, J.R., S. R. RUSHFORTH. AND J. D. BROTHER­ ST. CLAIR. L. L.,AND C. C. NEWBERRY. 1991. Catalogofthe SON. 1981. The algal flora of Navajo National Monu­ lichens ofUtall. Mycotaxon 40: 199-264. ment, Arizona, USA. Nova Hedwigia 38: 501-553. ST. CLAm, L. L., AND R. B. WARRICK. 1987. Acarospora JOHANSEN, J.R. AND L. L. ST. CLAIR. 1986. Cryptogamic nodulosa (Duf.) Hue. v. nodulosa: a new record for soil crusts: recovery from grazing near Camp Floyd North America. Bryologist 90: 48-49. State Park, Utah USA. Great Basin Naturalist 46: 632­ ST. CLAIR. L. L., B. L. WEBB, J. R. JOHANSEN, AND G. T. 640. NEBEKER. 1984. Cryptogamic soil crusts: enhance­ JOHANSEN, J. R., L. L. ST. CLAIR, B. L. WEBB, AND G. T. mentofseedling establishmentin disturbed andundis­ NEBEKER. 1984. Recoverypatterns ofcryptogamic soil turbed areas. Reclamation and Revegetation Research crusts in desert rangelands following fire disturbance. 3: 129-136. Bryologist 87: 238-243. SCHLATTERER, E. E, AND E. W. TiSDALE. 1969. Effects of KLEINER. E. E, AND K. T. HARPER. 1977. Soil properties in litter ofArtemisia, Chrysothamnus and Tortula onger­ relation to cryptogamic groUIldcover in Canyonlands mination and growth ofthree perennial grasses. Ecol­ NationalPark. JournalofRange Management 30: 202­ ogy 50: 869-873. 205. SCHROEDER, N. E., G. J. SCHROEDER, AND D. E. ANDE­ KLEMENT, O. 1955. Prodromus der mitteleuropaischen REGG.1915. Catalog ofthelichens ofIdaho. Bryologist flechtengesellschaften. Fedde's Repert. Sp. Nov., Bei­ 78: 32-43. heft 135: 5--194. SHUSHAN, S., AND R. A. ANDERSON. 1969. Catalog of the ___. 1958. Die stellung der flechten in der pflanzensozc lichens of Colorado. Bryologist 7~: 451-483. iologie. Vegetatio 8: 43-56. SKUJINS. J., AND B. KLUBEK. 1978. Nitrogen fixation and LoOMAN, J. 1964. The distribution ofsome lichen commu­ denitrification in arid soil cryptogamic crust microen­ nities in the Prairie Provinces and adjacentparts ofthe vironments. Pages 543-552 in Environmental biogeo­ Great Plains. Bryologist 67: 209-224. chemistry and geomicrobiology. VoL 2. Ann Arbor MACCRACKEN. J. G., L. E. ALEXANDER, AND D. W. PubL, Ann Arbor, Michigan. URESK. 1983. An important lichen of southeastern THOMSON, J. W. 1981. The lichen genera Catapyrenium Montana rangelands. Journal of Range Management and Placidiopsis in North America. Bryologist 90: 27­ 36: 35--J7. 39. MACK. R. N., AND J. N. THOMPSON. 1982. Evolution in ___. 1989. Additions and a revisedkey to Catapyrenium steppe with few large, hooved mammals. American in North America. Bryologist 92: 190-193. Midland Naturalist 119: 757-773. TIMDAL, E. 1986. A revision of Psora (Lecideaceae) in NASH. T. H., AND A. B. JOHNSEN. 1975. Catalog of the North America. Bryologist 89: 253-275. lichens ofArizona. Bryologist 78: 7-24. ___. 1990. Gypsoplacaceae andGypsoplaca, a newfam­ NASfI, T. H., AND L. L. SIGAL. 1981. Preliminary study on ily and genus of squamiform lichens. Contributionsto the lichens of Zion National Park. Journal of the Ari­ Lichenology, Bibliotheca Lichenologica 38: 419-427. zona-Nevada Academy of Science 16: 46-50. WEBER, W. A. 1967. Environmental modification in crus­ NASH, T. H., AND V. WIRTH. 1988. Lichens, bryophytes and tose lichens. II. Fruticose growth forms in Aspicilia. air quality. Bibliotheca Lichenologica 30: 3-297. Aquilo Series Botanica 6: 43-51. Great Basin Naturalist 53(1), pp. 13--23

BRYOPHYTES ON THE CALCAREOUS SOILS OF MUNGO NATIONAL PARK, AN ARID AREA OF SOUTHERN CENTRAL AUSTRALIA

A. J. Downingl and P. M. Selkirkl

ABSTRACT.-Bryophytes were found to be abundant as components of microbiotic soil crusts on the calcareous soils of Mungo National Park, an arid area ofsouthern central Australia. Six sites thatreflected differences in soils, topography, and vascular plant vegetation were studied. At each site bryophytes were abundant, both in tenns ofnumber ofspecies present andpercentgroundcover. Numberofspecies presentdidnot differ- Significantlybetweensites, butpercentbryophyte cover was lower at a site on sanddunes in malleewoodland anda site on a silcrete ridge than atthe four other sites. Environmental factors (soil texture, pH, conductivity, nutrient status, vascular plant vegetation, light level, leaflitter, and fire frequency) appear to playa Significant partin detennining bryophyte distribution. Mosses that occur at Mungo are also widespread on calcareous substrates throughout arid southern Australia. Many of the bryophyte species present at Mungo also occur on limestones in high rainfall areas of eastern Australia. Environmental factors favoring bryophytic soil crusts in arid Australia are also present on limestones in high rainfall areas ofeastern Australia andmay account for tilepresence ofmany aridzone bryophyte species onlimestones. InAustralia there appears to be a relationship between rainfall and the ratios of acrocarpous to pleurocarpous mosses, and tllallose to leafy liverworts. Recognition of calcareous soils, widespread in arid areas of southern Australia, may be pOSSible by assessing a combination ofcharacteristics ofbryophyte assemblages.

Key words: aricllands, Australia, bryophytes, calcareous soils, limestones, microbiotic crusts.

This investigation of bryophyte distribution holding known as Mungo was ptirchased by the at Mungo National Parkwas undertaken as part New South Wales National Parks and Wildlife of a wider study to detennine whether there is Service. In 1979 Mungo was offiCially declared a suite of bryophytes conSistently associated a national park and was extended in 1984 when with calcareous soils inAustralia, andwhetherit Zanci, the adjOining property to tlle north, was is pOSSible to detennine the calcareous nature added to the park. Sheep and cattle were re­ of a substrate by the associated bryophytes. moved from Mungo in 1978; their absence was Many bryophyte species that occur on lime~ important in our selection of a study area, as stones in the high rainfall areas ofeastern Aus­ observations of bryophytes on limestones and tralia are more usually associatedwith arid areas siltstones at Attunga, in eastern Australia indi­ of Australia (Downing 1992, Downing et al. cate tllat the presence of sheep changes the . 1991). This study considers tlle relationships nature of bryophyte assemblages (Downing between bryophytes of arid areas of southern 1992). Graetz and Tongway (1986) have shown Australia and bryophytes on limestones in the that removal ofmicrobiotic soil crusts by heavy relativelyhigh rainfall areas ofeasternAustralia. grazing causes changes in soil structure and . Mungo NationalPark (33°45'S, 142°59'E, 91 chemistrywhich are significantforplantgrowtll. m.a.s.l.) lies withintheAustralian aridzone (Fig. Overseas studies (Brotherson etal. 1983, Johan­ 1) and is well known for its Significance in Abo­ sen and St. Clair 1986, Kleiner and Harper riginal prehistory; with AbOriginal occupation 19(2) have shownthatsevere tramplingby graz~ dating back to 40,000 y.b.p. Prior to 1922 the ing animals canbe tlle most damaging hazard of present Mungo National Park was part of a microbiotic crusts in North American deserts. property of203,000 ha carrying apprOximately Lake Mungo lies on the flat plains of the 50,000 sheep. The property was subsequently Murray Basin, a shallow sedimentary basin divided into smaller holdings; and, in 1978, the created by subsidence at the beginning of the

lSchool ofBiological Sciences. Mncqllnrie University. Sydney. NSW 2109. Au,tndia.

13 14 GREAT BASIN NATURALIST [Volume 53

18° S ......

33° S

arid zone boundary • botanical regions

• sites described in text

Fig. 1. Map ofAustralia showing location of Carnarvon and NuUarbor Botanical regions and sites described in the text in relation to the Australhm Arid Zone.

Tertiary, now consisting of Quaternary aeolian Phosphorus content is always low «0.001%) sediments overlying Tertiary marine sequences and is correlated with soil texture (Northcote (Geological Survey of NSW 1971). Fluvial and 1980, Stafford Smith and Morton 1990). lacustrine sediments were deposited during the Lake Mungo, a vast, dry lake, is the principal Pleistocene, and these are overlain by Quater­ landform ofMungo National Park. The lunette nary sandy to clayey calcareous sediments, the orcrescent-shaped ridge that flanks the eastern most widespread surface geological formation shore of Lake Mungo and dominates the land­ ofthe Murray Basin (Northcote 1980). scape is visible froll the airas far as 100km away. Calcareous materials are a significantfeature The base ofthe IUQette is composed ofclay, silt, ofthe area, and calcium carbonate (CaC03) can and sand, topped bymobile siliceous sands. be present (3-30%) in soils as "fine earth car- The lake floor consists ofyellow-grey crack­ . bonate." Calcium carbonate canbe present also ing clays. To the west of the lake and to the as calcrete nodules, which are soft to very hard southeast of the lunette are open plainS of concentrations of calcium carbonate cemented brown calcareous earths. On the northeastern soil (15-45% calcium carbonate) in a usually side of the lunette are subparabolic dunes of loose calcareous soil mix (Goudie 1983). The siliceous sands (Eldridge 1985). A low silcrete surface horizons of most soils in the area are ridge runs through the northwestern section of alkaline, and most ofthe calcareous earths have the lake. This is the only site within the park pH values between 8.6 and 9.5. Deep sands are where rock (excluding calcrete rubble) is found, the exception and may have pH values close to and the hard silcrete rock has been a valuable neutral. Sodium chloride content can be as low resource for stone tool manufacture byAborigi­ as 0.02% in sands, 0.05% in all other soils, with nalpeople. the maximum salt appearing in horizons where Mungo lies within the arid zone ofAustralia there is the greatest visible accumulation of as defined by Meigs (1953), with the southern calcium carbonate. Nitrogen is low, 0.05--0.06% boundary ofthe Australian Arid Zone follOwing in the A horizons ofall SOils, indicatingverylow the 250-mm rainfall isohyet, the eastern and levels oforganicmatter. Potassium levels arelow northern boundary following the 375-mm iso­ in'sands and higher in sandy loams and loams hyet, and the Indian Ocean forming the bound­ «0.001%; Buckley 1982,· Northcote 1980). ary in the west (Fig. 1). 1993] BRYOPHYTES OF MUNGO NATIONAL PARK 15

TABLE 1. Climatic data. Mean monthly maximum and l~inimum temperatures (OC) for Mildura, 34°14'S, 142°05'E, 95 km SW of Mungo National Park: m.max = mean maximum temperature, m.min. = mean minimum temperature (Bureau of Meteorology 1975). Average monthly rainfall in mm for Pooncarie, 33°22'S, 142°35',56 km NW of Mungo (Bureau of Meteorology 1975). Rdays = number of rain days.

J FM A M J J A S 0 N D Yearly Temperahlre m.max(oC) 32 31 28 24 19 16 15 17 20 24 27 30 24 m.min.oC) 17 16 14 11 7 5 4 5 7 10 12 15 10 Rainfall Mean (mm) 21 20 17 18 26 27 21 24 21 26 20 21 262 Median (mm) 9 8 9 10 18 19 18 20 15 18 10 11 246 Rdays 3 2 2 3 4 5 5 5 4 4 3 3 43

Mungo National Park has hot, dry summers Site A: 33°43'45/fS, 143°Ol'15/fE, 98 m.a.s1 andcoolwinters (Table I-records for Mildura, "western shore of Lake Mungo. An eroded area closest temperature-recording station). Taking close to Mungo homestead and shearing shed. into account its more northerly location, mean Calcareous brown earth soils. Chenopod monthlytemperatures at Mungo are likelytobe shrubland with Maireana pyramidata (Benth.) I-2°C higher than those of Mildura. Frosts are P. G. Wilson (black blue bush) and occasional common during the cooler months ofthe year. Casuarina cristata Miq. ssp. pauper (F. Muell. Rainall averages for Pooncarie, 33°22'S, ex Miq.) L. A. S. Jolmson (belall). 142°35'E (closest rainfall recording station to Site n: 33°44'44/fS, 143°07'24/fE, 68 m.a.s.l. Mungo; Table 1), recorded over 103 years indi~ ·lake floor, SW of Walls of China. Yellow-grey cate an evenly distributed rainfall (Bureau of cracking clays. Chenopod shrubland with Meteorology1989a). However,in aridAustralia, Maireana pyramidata and sparse grasses. where there is considerable variation in rainfall Site C: 33°45'10"S, 143°07'30"E, 70 m.a.s.l. from yeartoyear, the meanis oftenmuchhigher lunette, eastern lake shore, N ofGrand Canyon than the median, and thus the median rainfall track. Yellow-grey cracking clay but with more valueis a more reliableindicatorofa typicalyear sand on the surface than at site B. Chenopod (Bureau of