Full Issue, Vol. 58 No. 4

Great Basin Naturalist

Volume 58 Number 4 Article 13

10-12-1998

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GREATR EAT BASINBAS I1 naturalistNAtuRatiALIaty ST

rebIALmeb m A

VOLUME 58 NQ 4 OCTOBER 1998

ML BEAN LIFE SCIENCE MUSEUM

BRIGHAM YOUNG university GREAT BASIN naturalist httpwwwlibbyuedunms FAX 8013783733801 378 3733

editor assistant editor RICHARD W BAUMANN NATHAN M SMITH 290 MLBM 190 MLBM PO box 20200 PO box 26879 brigham young university brigham young university provo UT 84602020084602 0200 provo UT 84602687984602 6879 8013785492801 378 5492 8013786688801 378 6688 emailE mail baumannbyueduricharclbaumannbyueduriehardrichard emailE mail nathan smithbyuedu associate editors

JAMES C CALLISON JR PAUL C MARSH department of environmental technology center for environmental studies arizona utah valley state college state university tempe AZ 85287 orem UT 84058 JERRY H SCRIVNER BRUCE D ESHELMAN department of biology department of biological sciences university of ricks college wisconsin whitewater whitewater WI 53190 rexburgredburgRexburg ID 83460110083460 1100

JEFFREY R JOHANSEN STANLEY D SMITH department of biology john carroll university department of biology university heights OH 44118 university of nevada las vegas las vegas NV 89154400489154 4004 BORIS C kondratieff department of entomology colorado state ROBERT C WHITMORE university fort collins CO 80523 division of forestry box 6125 west virginia university Morganmorgantowntown WV 26506612526506 6125

editorial board richard A heckmann chair zoology berranjerran T flinders botany and range science duke S rogers zoology bruce A roundy botany and range science richard R tolman zoology lanylarry L st clair botany and range science H duane smith monte L bean life science museum all are at brigham young university ex officio editorial board members include steven L taylor college of biology and agriculture and richard W baumann editor great basin naturalist the great basin naturalist founded in 1939 is published quarterly by brigham young university unpublished manuscripts that further our biological understanding of the great basin and surrounding areas in western north america are accepted for publication subscriptions annual subscriptions to the great basin naturalist for 1998 are 25 for individual sub- scriscribersbers 30 outside the united states and 50 for institutions the price of single issues is 12 all back issues are in print and available for sale all matters pepertaining to subscriptions back issues or other busi- ness should be directed to the editor great basin naturalist 290 MLBM PO box 20200 brigham young university provo UT 84602020084602 0200 scholarly exchanges libraries or other organizations interested in obtaining the great basin naturalist through a continuing exchange of scholarly publications should contact the exchange librarian 6385 HBLL poboxPO box 26889 brigham young university provo UT 84602688984602 6889

editorial production staff joanne Y abel technical editor emailE mail jyaemailbyuedu copyright 0 1998 by brigham young university ISSN 001736140017 3614 official publication date 12 october 1998 109810 98 750 27394 the great basin naturalist PUBLISHED AT PROVO UTAH BY ML BEAN LIFE SCIENCE MUSEUM BRIGHAM YOUNG university

ISSN 001736140017 3614

VOLUME 58 31 OCTOBER 1998 no 4

great basin naturalist 584 0 1998 appp 295 311

ALGAL composition OF microbiotic CRUSTS FROM THE CENTRAL DESERT OF BAJA california MEXICO

valenevaleriemalene R fiechtnerFlechtFlechtnernerlneri1 jeffrey R Johansenjohansenljohansen1johansenn1 and william H dark2clark2clarka

ABSTRACT A total of 66 algal species representing 32 genera were recovered from soils of 10 sites in the cataviacatavifiacatavma region of the central desert of baja california mexico the most common species encountered were the cyanophytes nostoc commune and schizothnxschizothrix calcalcicolecalcicolacicola the chlorophyte myrmecia astigmatastigmaticaastigmaticalica and the diatoms hantzschiaHantzschia amphioxusamphioxys Hantzhantzschiaschia amphyoxys f capitatecapitatacapitata luticolaLuticola cosmicohniicohmibohnii Luluticolaticola luticamutica and PinnulaPinnpinnulanapinnulariaulanariaflafia borealis var scalaris nine species not found in any previous studies of north american desert soils were present in our study sites including 3 taxa new to scienceselence cyhndrocystiscylindrocystis brebissoniibrebissonnbrebissonn var desertidebertideserti var nov elakatothnxelakatothrix obtusata sp nov and fasciculochlorisfasciculochlons rnexicanamexicansmexicanamexicana sp nov attempts to correlate species composition with soil chemical and physical parameters were unsuc- cessful apart from a ph effect on cyanobactenalcyanobacterial distribution overall composition of the soil algal community in the cataviacatavifiacatavma region is distinct from other desert sites we have studied although some cosmopolitan desert soil taxa were present

key words algae soil mexico baja california central desert cataviacatavifiacatavma cryptogamic crusts cylmdrocystiscylindrocystis brebis soniijoniisonn var desertidebertideserti elakatothnxelakatotbrix obtusata fasciculochlonsfasciculocbloris mexicansmeximexicanaeanacana microbiotic crusts

microbiotic crusts also called cryptogamic hchemzedlichenized cyanobactenacyanobacteria fix atmospheric nitro- crusts cryptocryptobioticbiotic crusts and microphytic gen and can be significant contributors to crusts are common in many andaridarld and semiarid desert nitrogen budgets rychert and skujins areas in the western united states st clair 1974 evans and Ehlenehleringernger 1993 belnap and johansen 1993 they consist of lichens 1996 crusts are susceptible to damage by live- mosses algae and fungi which form water stock backpackers and off road vehicular traf- stable surface aggregates that have been demon- fic which disrupt the crust compact the soil strated in some regions to be important in sta- and if frequent enough kill algal lichen and bilizingbilizing soil and preventing erosion harper moss components of the crust klemerkleinerkiemerkielner and and marble 1990 johansen 1993 williams et harper 1972 anderson et al 1982 rangefireRangefire al 1995 williams dobrowolski and west can also seriously impact the crust community 1995 furthermore many of the free living and johansen et al 1982 1984 1993 natural

depaitmentdepartment of biology john carroll university 20700 north paikpark blvd university heights OH 44118 cmajrmajJ smith museum of natural history albertsonaibAlbelboneibon college of idaho 2112 cleveland blvd caldwell ID 83605443283605 4432

295 296 GREAT BASIN naturalist volume 58 recovery from disturbance can take from a few does exist cameron 1969 all work on hot to many years anderson et al 1982 johansen deserts emphasizes the cyanobactenalcyanobactcrial compo- etetalal 1984 1993 callison et al 1985 nent of the soil community intelinterestest in the algal component of these the objectives of the present study were communities has increased in recent years twofold first we wished to carefully charac- early investigators of these algae were in-in terize the algal community from the central trigued by the occurrence of a group of organ- desert of baja california mexico a hot desert ismsiams generallygeneigenel allyaily thought to be aquatic in such region previously unstudied with regard to its an extremely and environment fionFlonfloristicsticstie work soil algal flora second we wished to test for has demonstrated the presence of a number of correlations between algal composition and algal geneiagenera some of which also occur in soil chemical and physical properties in sites aquatic habitats and many of which are con- with highly similar climate although soil fined to terrestrial ecosystems most fresh- chemistry has been considered important in water divisions are represented cyanophyta determining soil algal distribution starks et chlorophyta eustigmatophyta chrysophyta al 1981 correlations between taxa in desert xanthophyta bacillariophyta and euglenaeugleno soils and soil chemistry have not been made phytapayta in previously published work researchers tended to focus on I1 or 2 taxonomic MATERIALS AND METHODS divisions the cyanobacteriacyanobactena are probably the study area best studied group because identifications can be made based on morphomorphotypestypes in moistened ten study sites were established in the of of soils although there is considerable disparity cataviacatavifiacatavma area the central desert baja between cyanobactenalcyanobacterial floras based on mixed california mexico fig 1 a mid peninsular cultures and those based on uniumalgalunialgalalgal isolates location 9 km northwest of rancho santa ines diatoms are usually superficially treated al- 2846n IMGX 550 m elevation the geol- though rushforth and researchers trained in his ogy iss dominated by weathered cretaceous in paraguay grastilgrastie laboratory have found considerable diversity granite tonalite of the Jarjaraguayaguay block Grastil et al 1975 the decomposition of which forms in this group by direct preparation of soil dia- a sandy textured soil blom and toms anderson and rushforth 1976 johansen coarse clark 1984 mean annual has been re- et al 1981 1984 ashley et al 1985 johansen precipitation ported to range from 46 mm blom and clark and st clair 1986 green algae although 1984 to 101.7101 7 mm garcia 1981 mean annual abundant and ubiquitous are much less thor- iol1017 is 18 with a summer mean oughly studied due to the necessity of work- temperature is 19c of 25 ac a winter mean of 13 ac hastings with uniumalgalunialgalalgal isolates in which details of 258c8c 132c2c ing in 1964 hastings and humphrey 1969 garcia life history i e zoospore morphology color ie 1981 and occasional freezing temperatures changes with senescence etc are required vascular plant community is dominated for correct placement in genus and the in species by larrea tritrldentata sesse and moc coville johansen et al 1993 made some effort to tn and ambrosia chenopodifohachenopodifolia benth payne green algal isolates and demonstrated identify with of opuntia being the most com- the potential diversity of coccoid chlorophytes species mon cacti at each of the 10 sites we recorded in soils in presence of all perennial vascular plant species most papers dealing with taxonomy recent adjacent to the collection site table 1 micro- and distribution of desert soil algae study the biotic crusts formed significant cover in inter semiarid steppe cool deserts the shrub and in shrubintercactishrubmtercacti spaces figs 2 3 the 10 sites great and colorado plateau basin provinces will be referred to as sites 1 10 in this paper see johansen 1993 for a review of these they correspond to WH clark field collec- papers most work on soil algae of hot deserts tion numbers 9573 9582 specific locations was done in the early 1960s these studies for each sample site are as follows include both the sonoran cameron 1960 an 1964 etc and mojave durrell 1962 shields site 1 29047042n2904704 2n 114046111wFW iuliimllimal aw and drouet 1962 hunt and durrell 1966 site 2 29046573n294657 3 IUGIZ114046124w4w deserts to our knowledge the chihuahuan site 3 29047078n294707 8nan 114046049wIM 9xax desert soil algal flora has not been studied site 429047105n4 294ri0 5xax 114046105w1144610 5xax although a single report of soil algal density site 5 29047177n2904717 7nan IUGIG114 46162w2waw 19981 microbiotic CRUSTS OF baja california 297

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figs 1 3 catavifia area in baja california mexico 1 general topography and vegetation 2 microbiotic crusterust at site 10 3 detail of algal crusterust raised by slight disturbance sealescale 12 cm

site 6 290470872904r087nN WWluwwludwlU114 sample collection site 7 294r09rn29047090n 114046279w sample areas all within a 2 km area were site 8 29046444n 114046039v114046039w chosen to represent various soil types and habi- site 9 29046478n 114IMGX46048v tats present we obtained the precise location site 10 2904r039n29047039n 114046084w of each site with a sony PYXIS IPS 760 global 298 GREAT BASIN naturalist volume 58

tabletailleTABLL 1 perennial vascular plant species present in each of the 10 cataviacatavifiacatavma sites sites species 1 2 3 4 5 6 7 8 9 10 ambrosia chenopodifoliachenopodifoha benth payne X X X X X X ambrosia dubosadumosa gray payne X X X atriplex polycarpa torr S wats X Ceratceratoidesoides lanata pursh howell X encelia califorcalicallcailfornicamcamoa nutt X X X eriogonumfasiculatumenogonumfasiculatum benth X larrea tntritrldentata DC cov X X X X X X lycium califorcalifornicumcaliformcumnicum nutt ex gray X X opuntia cholla weber X opuntia ganden wolf rebman & pinkava X X opuntia molesta brandegee X prosopis glandulosaglandulose var torreyanatorreyana L benson johnston X Simmondsimmondsiasia chinenchinensissis link schneider X solanum hindsianum benth X Viguiefiguieraviguierara laclaciniatalacimatalaciniateluclacilacliaciniataimata A gray X X X X viscainoaViscainoa genigemgeniculatagemculatageniculateculata kell greene X X positioning system vascular plant cover and scope with nomarskidomarskiNomarski DICDIG optics and pho- percent visible coverage by microbiotic crust totographed using kodak PKL 135 film identi- were noted for each plot and recorded photo- ficationfication was made on the basis of cell and graphically composite crust samples consist- colony morphology using standard authorita- ing of 10 cores top 3 cm were taken in a 2 x tive references geitler 1930 32 Desikdesikacharyachary 2 m area at each site additional samples for 1959 kantz and bold 1969 for identification soil chemistry analysis were collected from the of non diatom eukaryotic algae individual iso- center of each plot samples were dry when lates were picked into 5 ml liquid BBM and collected 29 30 april 1995 and we trans- incubated 2 4 wk until good growth had been ported them to the laboratory within I1 wk of obtained identification was made on the basis collection where they were stored under of life history and morphological criteria using refrigeration until analysis standard authoritative references komarek and fott 1983 ettiettl and cartnergartner 1995 because of characterization many cyanophytes grow poorly on artificial non diatom algae media additional identification of cyanophytes composite samples were crushed and was made directly from wet mounts of wetted mixed to produce a homogenous sample A i- soil samples incubated 48 72 h in the light g aliquot was removed and added to 99 ml of a we prepared type materials in 2 ways 070.70 7 saline solution as an osmotic protectant holotype material was prepared by filtering a for a 102 dilution of the original sample young healthy culture onto glass fiber filters Aliquoaliquotsts of 0 1 or 020.20 2 ml were spread in tripli- that had been ashedached and subsequently han- cate on agar solidified Z 8 medium carmichael dled with forceps to minimize the possibility 1986 for quantitationquantiquantizationtation of cyanophyta and on of eukaryotic DNA contamination the filters bold s basal medium BBM bold and wynne were allowed to air dry placed in glassine 1978 for quantitationquantizationquantitation of non diatom eukaryotic envelopes attached to herbarium cardstock algae cultures were allowed to dry overnight and then wrapped in herbarium envelopes before inversion sealed with paraparafilmfilm and these materials were then deposited in the incubated in constant light at 20 23c until herbarium of nonvascular Cryptocryptogamscryptogramsgams at the good growth had been obtained 3 6 wk we monte L bean museum brigham young uni- then counted the number of colony forming verversitysity provo utah Uniunialgalalgal cultures having units on each plate for identification of cyan isotype or paratype status were grown on agar ophyta wet mounts prepared directly from slants of BBM and deposited in the UTEX individual isolates on Z 8 plates were exam- culture collection at the university of texas ined using an olympus BH 2 photomicro austin texas 199811998 microbiotic CRUSTS OF BAJA california 299

TABLE 2 summary of species abundance and richness data for each of the 10 study sites in the cataviacatavifiacatavma region of the central desert of baja california mexico abbreviations used cfugchug colony forming units of algaegramalgae gram soil CYANOB number of cyanobacterialcyanobactenal taxa EUKARY number of non diatom eukaryotic taxa DIATOM number of diatom taxa TOTAL total number of algal taxa VASCUL number of perennial vascular plant taxa CRUST estimated percent cover of microbiotic crust site cfugchug CYANOB EUKARY DIATOM TOTAL VASCUL CRUST

1 2 x 104 4 6 6 16 2 100 2 6 X 103 2 4 6 12 2 90 3 2 x 104 7 8 6 21 2 80 4 2 X 104 9 11 6 26 3 ND 5 8 x 103 7 4 8 19 4 55 6 5 x 104 6 6 6 18 6 100 7 5 x 104 6 10 8 24 5 100 8 2 x 104 6 8 7 21 5 100 9 1 x 104 4 14 6 24 6 100 10 2 x 104 3 4 9 16 1 100

characterization of physical parameters finally canonical corre- the diatoms sponspondencedence analysis CCA was used to simul- taneously ordinate sites species and environ- subsamples of each composite sample were mental variables braak 1986 1987 nor- removed acid cleaned washed and mounted ter mally quantitative data are used for CCA into permanent diatom slides following johan- analyses although had quantitative data sen et al 1982 we then examined diatoms we for diatoms we had none for all other algae using a zeiss axioskop photomicroscope with resolution of the CCA recorded high resolution Nonomarskidomarskimarski DICDIG optics relative to increase we 0 for 1 for a single isolation from a density of species was determined using counts absence and 2 for 2 isolations from a site of 100 frustulessample site or more for diatom taxa we recorded 0 for absence I1 soil chemistry for relative abundances of 1 15 and 2 for soil chemical and physical analyses were relative abundances 15 CCA was run with conducted by the soil testing laboratory at full species data sets and then subsequently brigham young university using standard run with shortened sets the taxa eliminated methods soil survey staff 1962 soil conser- in short sets were those which did not vary in vation service 1972 analyses included per- number in 9 or more sites cent gravel soil texture gravimetric method ph saturated paste electrical conductivity RESULTS and percent organic matter walkley black floristics method nitrate N calcium magnesium and sodium levels were determined from soluble the total concentration of algae in the 10 extracts phosphorus and potassium were ex- locations within the study site ranged from 6 x tracted using sodium carbonate via the olsen 103 to 5 X 104 cfugchug soil table 2 microbi- method standard for alkaline soils sodium otic crust cover was obvious at all sites with absorption ratio SAR was calculated using the majority of sites showing 80 100 cover levels of calcium magnesium and sodium perennial vascular plant diversity was low with only 1 6 species recorded from each area statistical analysis table 2 several different biometric methods were A total of 66 algal species representing 32 used to detect patterns in the data the 10 genera were recovered from these sites table sites were clustered based on jaccard s simi- 3 some widespread taxa were found in 8 or larity goodall 1978 utilizing the unweighted more sites these common taxa included the group average method of cluster generation cyanophytes nostoc commune and schizothrix pielou 1984 centered standardized princi- calcicolacalcicolecalcicola the chlorophyte myrmecia astigmat pal component analysis pielou 1984 was used ica and the diatoms Hantzhantzschiaschia amphioxusamphioxys to ordinate sites based on soil chemical and Hantzhantzschiaschia amphyoxys f capitatacapitatecapitata luticolaLuticola 300 GREAT BASIN naturalist volume 58

TAISITABU 1 3 algal distributiondistl ibution in 10 sites fromflom the cataviacatavifiacatavma rexionlegionregion of the Cergercentralgentralitral desert ofofbajdbaja california mexico cat- egoriesegoriesorles I1 1 isolate flom the site 2 2 01or moiemolemore isolates from the site relative density is given for diatom taxa and clnysophytechrysophyte cysts absence is indicated hyby a blank site species 1 2 3 4 5 6 7 8 9 10 cyanobactfiriaCYANOISAC II11 kia Anaanabaenahaenahuenabuena sp I1 linlxalyngbyelyngbya diguetidiguati coincomgoingom I1 linnalinhalinno pittputpittealisputeulisealiseuliseulls mont I1 Micromiciololeusmicrocoleuscoleus steenstrupsteenstrupnii boye pet I1 Micromicrocolettsmtcrocoleuhcoletts vanivagiuaffnatusvaginatusnatus valtvallmanmauvancheivanchenvaitchermanchelmauchelVanVait cheicherchel goingom 1 1 1 1 Myxomtiowrlinamyxosarcinasarcina hurmenburmenburmensishurmensississos skuja 1 myxosarcinaMyxomixoaiMixoaioalsarcinaonauna pectabilisspectabilis ceitlergeitleiceigler 1 Myxomixofurctnumyxosarcinasarcina sp 1 nostoc commune Vamauchellicherticher 1 2 1 1 1 2 1 1 1 nostocinuscoruinNIOC muwrum ag 1 nostocnoytocNO toetoo piscipiscinalenale kutkiltz 1 1 1 1 nostoc punctpunchpunctforineforinefo i me koitzkoltzkutz llanothariotharlot 1 1 1 1 plectoneinaplectoncma tongtomasinianurninianum vaivalvarvan gracile hansgiiansghanig 1 Plecplectonematonema sp 1 1 1 scliizothischizothrix i arenaria belkbeikberk gom 1 1 1 schizothrixchizothncalcicolaafgomcalcaicalcicolacalcicolecicola ag corngorn 2 2 2 2 1 2 2 2 2 2 sutSulscytonemaaneina ocellatuinocellatumocellatuintamtum lynglilyngblyngby I1 scytonenzaclclonemaonema sp 1 1 1 c111011ophytaC iiiokoihyia Apatoapatococcuscoccus constipatuscoiutipatus printz brintnimtzprintpimtz 1 2 1 Bracteabracteacoccusbracfcacoccuicoccus aggreatus teiteregderegoeioel eg I1 Bracteabrdcteacolliiibracteacoccuscoccus cohaerens bischott & bold I1 Bracteabracteacoccusbracteacocctificoccus grandis bischoff & bold 1 1 1 bracteacoccusbrticteucocwsBracteacoccus minormonotmonor chodat petrocipetrovipeti ova1t 1 1 1 2 1 bracteacoccusbralteacBracteacoccusLCU ininutusmvnutu schwaischwalschwarz 1 1 1 bracetacoccusbi alelacoccu& peudominoipseudoininor bischoff & bold 1 1 chlorellaChloiella ellipsoidellipsoideaelhp&oideaea cerneck 1 chlorella vulgaris bellelbeijerinckmck 1 2 2 chloichlorococcumococlwn minutuinminutumminutuin staistalstarr i I1 chlorowrlinopwchlorosarcinopsis areaggregatagata aicealcearcearee & bold 1 chloichlorosarcinopsisowrcinop&ifi arenicolalreiremcola croovergroover & bold 1 1 chiotchloichlorosarcinopsisowrcinops auxotrophica croovergroover & bold 1 1 chloichlorosarcinopsisosarcinopsis bastropiensisba&tropiensif croovergroover & bold 1 1 chlorosarcinopsischloi oiarlinopiis gelageiagelattmsagelatinosatinosa chant & bold 1 2 2 chlorosarcinopsischloro&arcinopsis 8ernipervirenssermperoirens croovergroovergi ooveikovei & bold 1 cylinbcijhndi ocif&tiocystisoocystis brebissoniibielw&onn vaivar dedesertideserticde&ertiserti sp nov 1 cohniibohnii luticolaLuticola luticamutica and Pinnulapmnulanapinnulariatiaria bore- cylindrocystiscyhndrocystis brebissoniibrebissomibrebis somi alis var alarisscalarsscscalaris most taxa were rare with 30 var desertidebertideserti var nov of the 66 species identified appearing in a sin-sin figs 4 9 gle site although rare min ouioulour sites most taxa colonialcoloniaeae cellularcellulaeCellulae solitanaesolitariaesolitariansolisolit tanae cylinaylin isolated are species that commonly occur in Coloni brasiniprasini ariae tricaedncaedricae extremisextremesex rotunrotundatusdatus 10 15 gmim latae desert soils notable exceptions iei e taxa not tremis 14 56 urngmumm longaelongac paries cellularcelcellulaelulae tenuis bellupellu formerly found in desert soils of north amer- in cidusaidus nucleus cencentraliscentralestralis Chloroplastchloroplastschloroplastusus elonga ica include cylindrocystiscyhndrocystis brebissoniibrebissomibrebis somi var tus morcisporcis longitudinallongitudinalibusibus lobatis constconstrictusconstnctusrictus desertidebertide serti elakatothnxelakatothrix obtusata elhptochlonselliptochloris ad centrum partes daaeduae formantes unaquaeque subsphaencasubsphaerica fasctculochlonsfasciculochloris mexicanamexicansmextcanamexicana lobo pyrenpyrenoideoide dimidiadimedia inm celluliscelmellulislulis aliquaalaqua axiaaxialibuslibus sphaeropsis lobophoralobophora lobosphaera tirolensis Pyrepyrenoidesnoides plu minusvemmusve distincta gragrgranulisgrabulisgramilismilisanulis luticolaLuticola muticmuticoidesmuttcoidesoidesoldes and VzVischeriaschena helvetica amelisamylis raro vagina amalaamyla evievidentievidentedenti Zygozygosporazygosporespora these taxa unusual in desert soils are de- non observataobservata scribed below three of them are new to typuscypus die april 1995 a solo dedesentiserti locus 7 science WHC 9579 lat bor 2947090294709.00 long oce 199819981 microbiotic CRUSTS OF BAJA california 301

TABLETABLES3 continued site species 1 2 3 4 5 6 7 8 9 10 dictyochloropsis splendida geitler I1 diplosphaeradiplosphaeru species I1 elakatothrixelakatothnx obtusata sp nov I1 I1 I1 elliptochlonselliptochloris subsphaerica reis ettl & gaitgalt 1 ettha bilobata vinatiervinatzerVinatzer komarek I1 I1 etthaettliaattlia cohaerens grooverGrooveivel & bold ettl & gart 1 1 fasciculochlonsfasciculochloris mexicansmexicanamexicana sp nov I1 klebsormidium dissectumdissectum gay ettl & cirtchrtgirtgartgarngann 1 1 klebsormidiumflaccidumklebsormidium flaccidflaccidumum koitzkoltzkutz sil matt & bl I1 1 lobosphaera tirolensis reisigl 1 1 1 lobosphaeropsisLobosphaeropsis lobophoralobophora andr ettl & gart 1 munellamuriella decolor vischer I1 muriella terrestristerrestris boye pet I1 myrmecia astigmatastigmaticaastigmaticalica vinatiervinatzerVin atzer 2 2 1 2 2 1 myrmecia biatorellae tsehaschtsch & plessl boye pet I1 1 1 1 1 Myrimyrtmyrmecianecianecla globosaglobose printz 1 myrmecia incisaencisa reisigl I1 Mynmyrmecianeclanecia macronucleata deason andr 1 1 1 spongiochloris minor chant & bold 1 1 stichococcus bacilbacillarisbacillanslansianslarislafis nageli 1 eustigmatophyta ViVischeriaschena helvetica vischer & pascher hibberd 2 1 1 2 bacillariophyta DIATOMS Hantzhantzschiaschia amphioxusamphioxys ehr grunow 11 14 16 8 5 10 10 13 25 50 Hantzhantzschiaschia amphioxusamphioxys f capcapitatacapitatecaptcapttatatataitataoata 0 mullermilllermulier 5 5 10 6 1 13 2 6 9 luticolaLuticola cohnn hilse mann 34 24 41 33 24 13 33 19 25 19 luticolaLuticola luticamutica kiltzkutz mann 30 38 32 33 38 32 32 33 35 20 luticolamuticoidesluticolaLuticola muticmuticoidesoides hustedt mann I1 nitzschia hantzschhantzschianaiana rabhlabh I1 nitzschia punctatepunctatapunctata var minor temp & paragperag 2 PinnulaPinnpinnulariapinnulanaulanaria borealis ehr 1 2 1 1 PinnulaPinnpinnulanapinnulariaulanariaflafia borealis var scalaris ehr labhrabh 1 6 10 1 1 1 3 staurosiraStaurosira construeconstruensns ehr williams & round 1 chrysophytachbysophyta chrysophyte cysts 20 18 34 15 21 31 21 28 5 7

imga114462791144627.9 9 regio catavifiaeCatacatavmaevmae desertumDesertum cen type collected inm april 1995 from desert soil tralis california Ininfernainfernalferna mexicum holotypusHolotypus surface site 7 WHC 9579 29470909947090294709.00 N BRY C 48041 herbarium Cryptogamcryptogamorumorum latitude 114462791144627.99 W longitude cataviacatavifiacatavma nonvasculanumnonvascularium bnghambrighambangham young university region central desert baja california mex- provo utah isotypus in statustamu vivo BC 989 8 ico holotype BRY C 48041 herbarium of UTEX congeriesCongeries culturarumCulturarum university of nonvascular Cryptocryptogamscryptogramsgams brigham young uni- texas austin texas versityversity provo utah living isotype BC 989 8 colonies vivid grass green cells solitary UTEX culture collection university of cylindrical with rounded ends 10 15 pragragnam texas austin texas wide 14 56 pragraim long cell wall thin clear this taxon is very similar to the nominate nucleus central chloroplast elongated with variety of C brebissoniibrebissonnbrebissonusonn meleghmenegh in terms of longitudinal lobed ridges constricted in the its chloroplast morphology and general shape center to form 2 halves each with a pyrenoid figs 5 7 it differs in its smaller size it is these halves appearing axial in some cells most similar to C brebissoniibrebissomibrebis somi var minor west Pyrenpyrenoidsoids more or less distinct with starch et west which has a size range similar to var grains rarely with an evident starch sheath desertideserttdeberti but differs in its slightly different Zygozygosporesspores not observed chloroplast structure the chloroplast of C 302 GREAT BASIN naturalist volume 58

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figs 4 11 cihndrocystiscyfindrocystis brebissoniibrebissonnbrebis sonnsonu var desertideberti and elakatothnxelakatothrix obtusata scale 10 pmirntrn figs 4 7 C brebissoniibrebissomibrebis somi vaivalvar desettidesettiserti 4 vegetative cell division note starch sheath around pyrenpyrenoidsoids 5 and 7 chloroplast showing lobed ridges and I1 pyrenoid 6 persistent cell walls following cell division arrows and oblique cell division right figs 8 11 E obtusata 8 9 short chain of vegetative cells 10 11 vegetative cell division note triangular shape of cells 199819981 microbiotic CRUSTS OF BAJA california 303 brebissoniibrebissomibrebissomi var minor is illustrated as being occasionally in short chains uninucleate 5 656.56 5 stellate dillard 1990 plate 24 fig 6 while lmtm in diameter 6 14 rinpinpm long cell wall thin that of var desertideberti is clearly elongated with chloroplast parietal elongate filling half to longitudinal lobed ridges fig 7 howeverHoweveiver in entire cell occasionally fragmented pyrenoid the smallest cells the chloroplast of var desertideberti indistinct reproduction only through auto- can appear stellate fig 4 and so the varieties spore production could be confused the most compelling argu- type collected april 1995 from desert soil ment for recognizing the new variety is the surface site 4 WHC 9576 2947105294710.55 N very distinct habitat differences cylindrocystiscyhndrocystis latitude IMIO114461051144610.5 5 W longitude cataviacatavifiacatavma brebisbrebissoniibrebissomisomi var desertideberti occurs in neutral to region central desert baja california mexico alkaline desert soils while other varieties of holotype BRY C 48042 herbarium of non- this species are in acidic aquatic habitats the vascular Cryptocryptogamscryptogramsgams brigham young univer- distinct habitat differences have caused us to sity provo utah living isotype BC 646 4 place relatively more importance on the minor UTEX culture collection university of texas morphological differences between the vari-varivarl austin texas paratype location site 5 eties sexual stages which are important in 2947177294717.7T N latitude 1144616114461621144616.2 2 W longi- delineating cylindrocystiscyhndrocystis taxa were not tude cataviacatavifiacatavma region central desert baja observed only a single isolate was obtained california mexico paratype BRY C 48043480437 herbarium of nonvascular Cryptocryptogamscryptogramsgams brig- elakatothnxelakatothrix obtusata sp nov ham young university provo utah living paratype BC 717 1 UTEX culture collection figs 8 11 university of texas austin texas colomacolonia flavovirens mucus extracellulosus our species differs from other species in mollis difdiffususdiffusesfusus copiosuscopiosus in culeuleuicultunsculturisculturistturis vetustior the genus in having much shorter cells which ibus cellularcellulaeCellulae generahtergenegcneraliterrahter decrescentisdecrescentesdecrescentes ad ex- are not as clearly tapered as is typical for the trema ovalesdovales ad leviter triangulatnangularestriangularesres raro prope genus E obtusata is most similar to E gelati sphaencisphaerici natura triangulatnangulantriangulariri maxime evievidentievidentedenti nosa which also possesses an amorphous confestimconfestim post divisionemdivisionem interinterdumdum curvataecurvatae mucilage and has cells somewhat longer and solitanaesolitariaesolitariansolisolitariaetanae vel binabmatimbinatimtim infrequenter in catenis thinner 13 18 pmgm long by 3 6 pragraum wide com- brevibusbreviousbrevibus unmucleataeuninucleatae 5 656.5gs6 5 gmumu m latae 6 14 pared to 6 14 tmum long by 5 656.56 5 tmum wide in E pragraurn longaelongac paries cellulosuscelluloses tenuis chloro obtusata because of its relatively small length elastusplastus parieparlepanepanetalisparietalisparietalestalistails elongatuselongateselongatus hemicellulam vel to width ratio E obtusata is much stouter and cellularcellulamcellulam comcomplensplens interinterdumdum fractuscractus prye less tapered than all other species min the genus noides indistincta reproductioreproduction non nisi per fabrifabricanfabncamfabricamcameam autosporarum elliptochloriselkptochlons subsphaericasubsphaenca reisigl cypus typus die april 1995 a solo desertidedebertiserti locus 4 ettl & cartnergdrtnergartner 1995 WHC 9576 lat bor 29471059947105294710.55 long oce p 424 fig 127a127 a c 1144610114461051144610.5 5 regio catavifiaeCatacatavmaevmae desertumDesertum cen tralis california infernaIninfernalferna mexicum holotypusHolotypus colony spherical slightly mounded grass BRY C 48042 herbarium Cryptogamcryptogamorumorum green even at 6 mon cells cylindrical when nonvasculanumnonvascularium brigham young university young occasionally slightly bent up to 151.5isls1 5 times provo utah isotypus in statustamu vivo BC 646 4 as long as wide 3 8 tmum wide 5 10 lmtm long UTEX congeriesCongeries culturarumCulturarum university of becoming ellipsoidal to spherical with age texas austin texas paratypus BRY C 48043 spherical cells 8 18 tmum in diameter chloro- herbarium cryptogamoruinCryptogamcryptogamorumorum nonvasculanumnonvascularium plasts typically single parietal with a central brigham young university provo utah paiapalapara pyrenoid usually touching the edge of the cell typuscypus in statustamu vivo BC 717 1 UTEX congeriesCongeries at only a few points becoming lobed or dis- culturarumCulturarum university of texas austin texas sected into several plasticsplastidsplastids with age starch colony yellow green extracellular mucilage positive when treated with iodine reproduc- soft diffuse copious in older cultures cells tion only through autospore production generally tapered at ends oval to somewhat this chlorophyte is distinctive because of triangular rarely nearly spherical with triangu- its production of small cylindrical cells that lar nature most evident immediately after divi- eventually round out to become larger spheri- sionslon occasionally curved in singles or pairs cal cells 304 GREAT BASIN naturalist volume 58

fasciculochlorisfasciculochlons mexicansmexicanamexicana sp nov with 1 2 contractile vacuoles 24242.4 323.2 4 imlm diameter 5 8 pmgm long flagella of approxi- figs 12 16 in mately equal length stigma linear median colomacolonia scaberascabery sicca subsubstratestrato adhaerens to anterior in position chloroplast parietal atrovindisatroviridis cellularcellulaeCellulae dispositae in massis fasci- median nucleus posterior culus cubiciscubicis vel tetratetratibustibus isobilateralisobilaterahbusisobilateralibusibus type collected in april 1995 from desert parieparlepanepanetibusparietibustibus arete adpressadpressisis in culeuleuicultunsculturisculturistturis juven soil surface site 5 WHC 9577 29471772904rl77294717.7 tibus in cultunsculturiseulculeuiculturistturis vetustionbusvetustioribus fere sphaeres N latitude 114461621144616.2 W longitude catavinacatavifiacanavinaCatavina centes et in muco inclusae unmucleataeuninucleatae region central desert baja california mexico 353.53 5 6566.55 limlm latae 6 858.58 5 lm longaelongac paries cel holotype BRY C 48044 herbarium of non- lulosuslukosus tenuis firmus spissescens ad 4 gmim in vascular Cryptocryptogamscryptogramsgams brigham young univer- cultunsculturisculeuleuiculturistturis scenescentibus Cytoplasma franugranu sity provo utah living isotype BC 767 6 UTEX lansianslaris vacuovacuolisvacuoleslis contractilibus durbusduabus visibilibus culture collection university of texas austin in celluliscelmellulislulis juvenjuventibustibus chloroplastschloroplasus parieparlepanepanetalisparietalisparietalestalistails texas cellularcellulamcellulam comcomplensplens ubi malurusmaturus Pyrepyrenoidesnoides the original description of the genus fasci excentricexcentricaexcentncaexcentricala grgranulisgianulisanulis grangrandibusdibus amaliamyli consoniconsoci culochlorisculochloris mclean and trainor 1965 cites ata solitanasolitariasolitanosolisolit ariaarlatana in celluliscelmellulislulis juvenjuventibustibus aliquot in packet formation by vegetative cell division in celluliscelmellulislulis vetustioribusvetustionbus zoosporae 4 16 in quo 2 or 3 planes presence of an extracellular que cellulacallula materno ellipsoideae parieparlepanepanetibusparietibustibus gelatinous matrix surrounding individual cells flagellisflaflagellistgellis binis circa aequalibus 1 2 vacuovacuolisvacuoleslis and cell packets and production of walled contractilibus 24242.42 4 323.23 2 4 imtm latae 5 8 imlm zoosporeszoo spores with the average size of 4 x 7 imtm longaelongac stigma lineanslineauslinearislinearisneans media ad anticum and unequal flagella I1 species FE boldiiboldia is Chloroplastchloroplastschloroplastusus parieparlepanepanetalisparietalisparietalestalistails in medio zoosporazoosporezoo spora included in the genus our isolate displays the nucleus posticusposhcusposhkusposticus generic characteristics of cell packets formed typuscypus die april 1995 a solo desertidedebertiserti locus 5 by vegetative cell division mucilage sur- WHC 9577 lat bor 2947177294717.77 long oce rounding both individual cells and cell pack- 1144616114461621144616.2 2 regio catavifiaeCatacatavmaevmae desertumDesertum cen ets and production of walled zoosporeszoospores with tralis california infernaIninfernalferna mexicum holotypusHolotypus flagella that are longer than the body length BRY C 48044 herbarium Cryptogamcryptogamorumorum and at least in some cases slightly uneven in nonvasculanumnonvascularium brigham young university length 10 difference comparison of agar provo utah isotypus in statustamu vivo BC 767 6 grown cultures of our isolate with FE boldiiboldia cul- UTEX congeriesCongeries culturarumCulturarum university of ture 1451 obtained from the UTEX culture texas austin texas collection revealed several differences the colony rough dry adherent dark green most notable of which were mucilage produc- cells arranged in masses cubical packets or tion and zoospore morphology mucilage pro- isobilateral tettetradstetradisrads with tightly appressedappiappl essed walls duction in our isolate was copious and evident in young cultures becoming nearly spherical surrounding both individual cells and cell and encased in extracellular mucilage in older packets figs 12 13 average cell diameter of cultures uninucleate 353.53 5 656.56gs 5 gmumum wide 6 858.58 5 vegetative cells was 646.4 lmum not including the lmlim long cell wall thin firm becoming thick- mucilage envelope and 898.9sg umgm including the ened to 4 almjlmtraihaihn in senescent cultures cytoplasm envelope the diameter of envelopednonnonenveloped cells granularglangianular with 2 contractile vacuoles visible in was slightly smaller than that reported in the young cells chloroplast parietal filling the cell literature mclean and trainor 1964 cells when mature pyrenoid eccentric associated from UTEX culture 1451 were approximately with large starch granules solitary in young 7 gmum in diameter mucilage production was cells multiple in older cells Zoozoosporesspores 4 16 per less pronounced than that observed in our iso- mother cell biflagellate walled ellipsoidal late or evident in photographs in the original

figs 12 25 see facing page fasciculochlorisfasciculochlons mexicansmexicanamexicana and ViVischeriaschena helvetica scale 10 gmum figs 12 16 FE mexi cana 12 packet formationfoihoiror mationmatlon in vegetative cells 13 cells embedded in extracellular matrix 14 16 zoospore formation and morphology figs 17 25 V helvetica 17 18 vegetative cells showing lobed chloroplast and prominent pyrenoid 19 autospore formation 20 mature cells showing carotenoid accumulation 21 zoosporeszoospores note prominent stigma nowarrowa 22 young vegetative cell 23 mature cell in early stages of orange pigment accumulation 24 zoosporangium 25 mature cell note irregularn i egulai cell shape 19981 microbiotic CRUSTS OF BAJA california 305

m4ma

MMM A

4 306 GREAT BASIN NATnaturalistURALIST volume 58

publication the most striking difference be- habitat from the wet rocks and mosses in the tween our isolate and FE boldnboldu is inm zoospore austrian alps from which this species was morphology the original description of FE described we are concerned that with the boldnboldu zoozoosporesspores includes the presence of 2 disparity of habitats our taxon may actually contractile vacuoles a median to posterior have a genetic identity quite different from stigma flagella of unequal length and a size of the type of the species 4 X 7 gmjmJ m characteristics of the UTEX 1451 culture are generally consistent with the origi- luticolaLuticola muticmuticoidesoides hustedt nal description with the exception of size mann in round et al 1990 zoosporeszoospores of our subculture of the type mater- hustedt 1961 6659866 598 fig 1602 ial averaged 3 X 6 gmjm zoozoosporesspores of our 3x6 the valves broadly elliptical lanceolate with isolate had an average size of 3 X 535.35 3 gm and a im rounded ends 10 18 gmum long 6 7 gmujnuan wide clearly anterior stigma fig 16 length of the raphe filiform proximal ends clearly deflected 2 flagella appeared approximately equal on to one side axial area broadened toward the some cells but flagella which differed in center isolated punctum in a marginal or near length by approximately 10 ss555.55 5 nmgm vs 66.060go 0 55 60 marginal position on side opposite the side nmgm were also observed exam- after extensive toward which raphe ends are deflected cen- ination of glutaraldehyglutaraldehyde de fixed cells we feel tral area transverse but generally not reaching confident in describing the flagella of our iso iso- the margins striae distinctly punctate 22 24 late as uneven but less obviously so than the in 10 umum flagella of FE boldn tm the elliptical lanceolate shape strongly deflected raphe ends and marginal punctum lobosphaeropsisLobosphaeropsis lobophoralobophora separate this taxon from luticolaLuticola mutica and andreeva lutica its varieties our specimens were more coarse- ettl & cartnergartner 1995 ly striated than those observed by hustedt p 418 fig 123a123 a 1961 66 cells spherical to subsphencalsubsphericallsubspherically uninucleate 4 12 gmnm in diameter cell wall thin in ViVischeriaschena helvetica pascher young cells thickening to I1 usngm in older cells vischer & hibberd 1981 chloroplast parietal becoming lobed or some- ettl & gdrtnergartner 19952401995 240 times filling the entire cell with a clear pyre- fig 61giegle c e noid droplets oil present some cells with a figs 17 25 slight orange pigment reproduction only through production of 2 4 autosporesautospores colony spherical yellow green to olive cells the genus lobosphaeropsisLobosphaeropsis was separated spherical to oval infrequently irregular 8 22 from chlorella based on its lobed chloroplast gmim in diameter cell wall thin chloroplast parietal 1 reisigl 1969 it is distinct from lobosphaera sometimes covering only I side of the in that it possesses a pyrenoid cell often lobed with a square cut pyrenoid orange pigment often obvious starch not pre- lobosphaera tirolensis sent A large vacuole with brownian move- reisigl 1964 ment of contents often present oil droplets evident in some cells ettl & cartnergartner 1995 in reproduction through production of 2 p 415 fig 122d122 d 4 autosporesautospores or through zoospore production Zoozoosporesspores flask shaped iniini- colony green to yellow green cells spheri- tially elongated metabolic 323.23 2 8 gmim wide cal or less often oval in small groups unauuninuuninnunmu 8 16 gmumu m long rounding up quickly chloro- cleate 6 16 lmjm in diameter cell wall thin plast band shaped covering 13 or less of the chloroplast parietal lobed without a pyrenoid cell stigma eustigmatophyceaen anterior starch granules visible when stained with prominent outside of chloroplast iodine reproduction only through autospore characteristics of our strains are very similar production to the description of eustigmatos magnus JBJ B our isolates fit the morphological descrip- petersen hibberd 1981 E magnus is char- tion for this species quite well although the acterized as having spherical cells with a flexi- desert soil of baja california is a very different ble smooth cell wall lobed parietal chloroplast 199819981 microbiotic CRUSTS OF BAJA california 307

TABLE 4 soil chemistry analysis of 10 sites in the central desert of baja california mexico OM organic matter EC electrical conductivity lmhoscmjmhoscm SAR sodium absorption ratio percent sand silt and clay were calculated after removal of gravel mineral nutrients in ppm sitesrte

parameter 1 2 3 4 5 6 7 8 9 10 mean ph 74 63 73 73 75 75 77 65 69 66 71 gravel 100 160 220 210 350 110 90 110 150 90 159 sand 808 771 666 733 576 781 879 801 791 751 756 clay 82 114 144 117 224 82 64 99 89 102 112 silt 110 115 191 150 201 137 57 100 120 147 133 OM 05 06 06 08 04 06 04 05 07 04 06 n03NO 37 31 26 23 33 27 21 26 28 29 28 P 108 227 99 107 312 74 83 177 179 94 146 eachexch K 64 77 102 69 58 58 27 58 102 94 71 sol ca 724 187 781 792 1019 480 754 222 526 216 570 sol mg 99 90 142 62 155 74 54 141 122 101 104 sol na 496 160 262 54 374 248 16 178 96 123 201 EC 07 03 06 04 08 05 03 04 04 03 05 SAR 14 08 07 02 09 09 01 07 03 06 07

large vacuole and angular pyrenoid vischeriaVischeria pairs had similarity indices of 29 44 because helvetica shares the lobed chloroplast angular the sites clustered so poorly this cluster is not pyrenoid and in most cells the thin smooth shown cell wall and spherical shape however hib- soil chemistry analyses reveal that the soils berd 1981 states that at least some cells in a are sandy have little organic matter 08os 080.8 culture of spherical smooth walled cells are and have a ph in the neutral to slightly acidic irregular or polyhedral his illustration of V range table 4 the greatest difference among helvetica hibberd 1981 fig 10 is identical to soils is seen in the amount of phosphorus our cells the sizes of the 2 species are also potassium calcium and sodium comparison similar with our cells being more similar to of the relative similarity of sites using PCA the size range reported for E magnus flagel- analysis based on soil chemistry showed little lated cells in vischeriaVis cheria are unmentioned by agreement with the clustering dendrogram hibberd 1981 but illustrated in ettl and based on floristic analysis this finding sug- gartner 1995 we did not observe the dis- gests that differences in algal composition tinct ridges figured in the line drawings of V seen among the sites cannot be explained by helvetica ettl and gartner 1995 we decided the 14 soil variables examined to place our strains into V helvetica based on to further explore the relationship between the consistent presence of 3 cornered angular soil chemical and physical parameters and algal cells and on the similarity in appearance to species distribution we performed several photomicrographsphotomicrography of the type culture hib- canonical correspondence analyses CCA most berd 1981 if we are correct our photomicro were informativeununinformative since we had more envi- graphs of the zoosporeszoospores are the first to be pub- ronronmentalmental variables than sites it was not lished fig 21 these taxa need further char- appropriate to perform CCA with all environ- acterization and we are not certain the distinc- mental variables forward selection of envi- tion between eustigmatos and vischeriaVis cheria will ronronmentalmental variables was conducted and the persist when more strains of both are isolated following were selected organic matter K and characterized electrical conductivity percent gravel ca mg and sodium absorption ratio short- relationships between algal the ened species list was used CCA distribution and soil variables this did not agree with either the cluster analysis or the clustering of sites on the basis of algal PCA based on soil chemistry we interpret this species revealed a low level of similarity among to mean that correlations between combined the sites probably due to the extent of the rare soil chemistry parameters and algal species taxa present sites 6 and 7 were most alike distribution are poor subsequent analyses uti- showing 52 similarity the remainder of site lized single environmental variables which 308 GREAT BASIN naturalist volume 58

dispbisp chelchei cybrdcybry 7 mude mute chauchvu 0

clau elsueasu scyt 0 copinopi 2 niha clarciar aniha 10 bibopibo A 613rps accoapco myxo nocci cigvtco komunomu 0 cyst plt09 albosplbos haamc 08 5010 lu nopullill lumuslumu& shaam&haam was cibaalbaclba dihevihe kid laslaa 0myasamyas 6 3ce31rr I1 brcoarco 6 brrnr 4vrgr lapulypu 0samispmi lydi mabimybi bratbrmt loti0 anisumanipum etbj6 oaymaomyma scar mxspm E 9 plecpiec lbloablo clapclag ayinmyin eccoetco

siba 4 cise0

fig 26 canonical correspondence analysis CCA of the 10 catavinacatavifiacanavinaCatavina sites when the axes were constrained to ph alone all species used but not all shown axis 1 is negatively correlated with ph r 09620.9620 962 iei e more alkaline samples aiealeare to the left moiemolemore acidic samples to the right eigenvalues for ist and and2nd axes 02500 250 and 03860 386 respectively sites hollow hexagons cyanophytes solid squares chlorophytes solid circles diatoms and chrysophytes solid anglesangiestrianglestu eustigmatophytes solid hexagon species codes consist of the first 2 letters of the genus first 2 letters of the species and fustfirsthirstbirst lettelletterietter of the subspecific taxon if given with unknown species the first 4 letters of the genus are given potentially confusing codes follow brmrarmr bracteacoccusBracteacoccus minormtnorminon bratbrmt bracteacoccusBracteacoccus minuminutesminutustus ch moreliachlorella cc chlorococcum cl chlorosarcinopsis cyst chrysophyte cysts lo10 lobosphaera lb lobosphaeropsisLobo sphaeropsis lumu luticolamuticaluticolaLuticola luticamutica my Myrmyrmecianiecia MX myxosarcinaMyxo sarcina sc schizothrixschizothnx si stichococcus st staurosiraStaurosira showed substantial variability among sites note that the myrmecia species were found these have been presumed to be important in clustered together suggesting they have simi- the literature ph percent silt sodium absorp- lar ph requirements canonical correspon- tion ratio of these only the CCA constrained dence analysis is less effective with the few to ph was informative fig 26 cyanobacter sites used in this study so these results should ial taxa occurred in soils of higher ph while be interpreted with caution however CCA chlorophyte and diatom taxa were distributed does appear to be a promising ordination across the entire ph range it is interesting to method for soil algae apart from ph we can 199819981 microbiotic CRUSTS OF BAJA california 309

see no trends that clearly connect algal species our contention that composite samples are a distribution in the catavinacatavifiacanavinaCatavina sites to soil chemi- necessity given spatial heterogeneity in crypto- cal andor physical parameters gamic crust communities grondin and johansen 1993 discussion acknowledgments this study demonstrates that when a variety of methods are used to characterize the our thanks to david M ward jr who community a high level of algal diversity can assisted with fieldwork and susan okuley be detected in desert soils we identified a who assisted with algal plate counts klaus total of 66 species in past studies of desert fritsch graciously helped with difficult ger- soil algae many investigators have concen- man wordspassageswords passages the zeiss axioskop micro- trated their efforts on identifying cyanobacte scope was purchased in part with funds from ria and diatoms cameron 1960 1964 durrell NSF BIR 9319239 continuing support for 1962 shields and drouet 1962 hunt and supplies and publication costs was provided durrell 1966 anderson and rushforth 1976 by john carroll university john carroll uni- johansen et al 1981 1984 ashley et al 1985 versity also provided release time for research johansen and rushforth 1985 johansen and to both flechtner and johansen st clair 1986 the absence of careful chloro phytepayte and xanthophyte characterization is likely literature CITED due to both the difficulty of their identification which requires uniunialgalalgal culture and the lack ANDERSON DC AND SRS R rushfortiirushfortil 1976 the cippciypcrapcryp togamic flora of desert soil crusts in southern utah of comprehensive taxonomic treatments prior nova hedwigishedwigiaHedwigia 2869128 691 729 to 1980 in general when non diatom eukary- ANDERSON DCD C KTK T HARPER AND SRS R rushforth 1982 otic algae were identified it was only to the recovery of cryptogamic soil crusts from grazing on genus level martin 1939 cameron 1960 cain utah winter ranges journal of range management 3535535 355 359 1964 friedmann al 1967 et archibald and ARCHIBALD PA AND HCH C BOLD 1975 notes on the bold 1975 metting and rayburn 1979 cam- edaphic algae of the galapagos soil science 120 eron 1964 attempted to identify non diatom 400 402 eukaryotic algae from the sonoran desert in ASHLEY J SRS R rushfortii AND JRJ R JOIIANSENJOHANSEN 1985 soil algae of crusts from the uintah southern but identified only 9 cryptogamic basin arizona species utah USA great basin naturalist 4543245 432 442 johansen et al 1993 observed a total of 72 BELNAP J 1996 soil surface disturbances in cold deserts algal taxa from a single site in the lower effects on nitrogennitrogenatenitrogenasease activity in cyanobactenalcyanobacterial lichen columbia basin washington which was sam- soil crusts biology and fertility of soils 2336223 362 367 BLOM phobetus pled seasonally for 12 mon observed 47 PE AND WH CLARK 1984 Phobetus desertusdesertus a they new melolonthine scarabaeidae coleoptera from chlorophytes and 9 xanthophytes many of the central desert of baja california mexico pan which were identified to species it is interest- pacific entomologist 6030460 304 312 ing to note that the latter study employing BOLD HCH C AND MJM J WYNNE 1978 introduction to the methods identical to those used in this study algae prentice hall inc englewood cliffs NJ 706 appp had nearly identical numbers of taxa even CAIN J 1964 A preliminary survey of the algal flolafloiaflora of though the floras demonstrated striking differ- soils of certain areas of texas southwestern natural- ences in composition ist 9166 170 this study is the most comprehensive floris- CALLISON J JDJ D brotherson AND JEJ E BROWN 1985 the effects of fire on the blackbrushblackbrush coleogyne tic examination of soil algae from a geographi- ramosissima community of southwestern utah jour- cally restricted hot desert community the nal of range management 3853538 535 538 large number of species observed is due both CAMERON R E 1960 communities of soil algae occurring to the variety of assay techniques used and the in the sonoran desert in arizona journal of the ari-an- fact that 10 subsitessubsubsidessites within the area were stud- zona academy of science 1 85 88 1964 terrestrial algae of southern arizona trans- ied the observation that over 50 of the taxa actions of the american microscopical society 83 were identified from a single site demonstrates 212 218 compositional heterogeneity within desert 1969 abundance of microflora in soils of desert soils this finding also causes us to speculate regions technical report 32 1378 jet propulsion laboratory pasadena CA 16 appp that more sampling extensive in any one of our carmichael WW 1986 isolation culture and toxicity subsitessubsubsidessites would yield more taxa and reinforces testing of toxic freshwater cyanobactenacyanobacteria blue green 310 GREAT BASIN naturalist volume 58

algae pages 1249 1262 min V shilovchilov editor funda- JOHANSEN JRJ R AND SRS R RUSHFORTH 1985 cryptogamic mental research in homogenous catalysis volume 3 crusts seasonal variation in algal populations in the cordongordon and breach new york tinticgintic mountains juab county utah USA great DESIKdesikacharydlsikaciiaryACHARY TV 1959 cyanophyta indian council agri- basin naturalist 451445 14 21 culturalal research new delhi 686 appp JOHANSEN JRJ R AND LLL L ST CLAIR 1986 cryptogamic DILLARDdil LARD GEG E 1990 freshwater algae of the southeastern soil crusts recovery from grazing near camp floyd united states part 3 chlorophyceae zygnematales state park utah USA great basin naturalist 46 zygnemataceae mesotaeniaceae and desmidiaceae 632 640 section 1 bibliotheca phycologicaphycological 85185 1 275 JOHANSEN JRJ R J ashleyandwrASHLEY AND WR RAYBURN 1993 effects durrellDURIU LL LWL W 1962 algae of death valley transactions of rangerangefirefire on soil algal crusts in semiarid shrub of the americanamerlean microscopical society 8126781 267 278 steppe of the lower columbia basin and their subse- great eillETTL H AND G GARTNERGARDNER 1995 syllabus der boden quent recovery basin naturalist 537353 73 88 luft und flechtenalgenflecbtenalgen gustav fischer verlag JOHANSEN JRJ R A JAVAKUL AND SRS R RUSHFORTH 1982 stuttgart 721 appp effects of burning on the algal communities of a high EVANS D AND JRJ R ehleringer 1993 broken nitrogen desert soil near wallsburgWallsburg utah journal of range cycles in andandlandsaridarldaridlandssandlandslands evidence from 15nN of soils oecolodecolo management 3559835 598 600 gia berlin 9431494 314 317 JOHANSEN JRJ R SRS R RUSHFORTH AND JDJ D brotherson FRIEDMANN I1 Y LIPKIN AND R OCAMPO PAUS 1967 1981 subaerialSuba enal algae ofnavajoof navajo national monument desert algae of the negev israel phycologia 6 arizona great basin naturalist 4143341 433 439 185 195 JOHANSEN JRJ R LLL L ST CLAIR BLB L WEBB AND GTG T GARCIA G 1981 modificacionesmodificaci6nes al sisterna de clasificacionclasificaci6n NEBEKERNEBLKER 1984 recovery patterns of cryptogamic climaticaclimdticaclimaticalclielleilmatleamatica de koppen enriquetaEnnqueta carciagarciagareia de miranda soil crusts in desert rangelands following fire distur- mexico df219ppDF 219 appp bance bryologist 8723887 238 243 GEITLFR L 1930 32 cyanophyceae koeltz scientific KANTZ T AND HCH C BOLD 1969 morphological and taxo- books konigstein germany 1196pp1196 appp nomicnomie investigations of nostoc and anabaena in cul- mycologicalphycological studies IX of texas GOODALL DW 1978 sample similarity and species cor- ture university publication 6924 67 pp relation pages 99 149 in RHR H whittaker editor ordi- ap KLEINER E AND K T HARPER 1972 and nation of plant communities W junk publishers EFF KX environment the hague community organization in the grasslands of canyon lands national park ecology 532995389953 299 309 GRASIILGRASTIL RG RP PHILIPS AND EC ALLISON 1975 KOMAREK AND B ponFOTT 1983 chlorococcales pages reconnaissance geology of the stalestate of baja califor- J funpoh 1 1043 in G huber pestalozzi editor das phyto- nia geological society of america memoir 140 170 plankton des SasubwasserssawassersSuBwassers volume 7 Schweizerschweizerbartbart PP stuttgart GRONDIN A AND JRJ R JOHANSEN 1993 microbial spatial MARTIN TL 1939 occurrence of algae in some heterogeneity in microbiotic crusts in colorado the in virgin in in utah soils proceedings of the soil science society national monument I1 algae great basin naturalist of america 42494 249 250 532453 24 30 MCLEAN RJR J AND FR TRAINOR 1965 fasciculochlorisfasciculochlons a HARPERHARPLR K 1 AND R MARBIMARBLE E 1990 A role for nonvas- KTI JRJ new chlorosphaeracean alga from a connecticut soil cular plants in management of and and semiarid in phycology 41454 145 148 rangeland pages 135 169 min PT tuellerthellertheiler editor veg- MEMETTINGITING BBANDWRAND WR RAYBURN 1979 algal communities etation science applications for rangeland analysis science and soil micro environments in eastern washington and academic publishers management kluwer silt loam soil science 12774127 74 78 dordrechtDordrecht germany PIELOU ECE C 1984 the interpretation of ecological data HASTINGSHAS 1 INGS JR 1964 climatological data for baja califor- john wiley and sons new york 263 appp of nia university arizona institute of atmospheric REISIGL H 1964 zur Systemsystematiksystematicatik und okologieOkologie alpiner physics technical report 14 132 appp bodenalgenBodenalgen osterreichische6sterreichische Botabotanischebotamschenische zeltzeitschnftZeitzeitschriftschrift HASTINGSHAS 1 INGS JR AND RRR R HUMPHREY 1969 climatological ililii111402111 402 499 data and statistics for baja california technical report 1969 bodenalgen studienstudden II11 osterreichische 18 university of arizona institute of atmospheric Botabotanischebotamschenische zeltzeitschnftZeitzeitschriftschrift ilg116492116 492 506 physics 96 appp ROUND FE RMR M CRAWFORD AND DGD G MANN 1990 hibberihlliblrdhibberdHIBnerinerlBERi DJ 1981 notes on the taxonomy and nomen- the diatoms cambridge university press cam- clature of the algal class eustigmatophyceae and tntritrl bridge 747 appp bophyceae synonym xanthophyceaexanthophxanthopeXanthoph ceabceae botanical RYCHERT RCR C AND J SKUJINS 1974 nitrogen fixation by journal of the linnean society 829382 93 119 blue green algae lichen crusts in the great basin HUNIHUNT CD AND LW DURRELL 1966 distribution of desert proceedings of the soil science society of fungi and algae USU S geological survey professional america 3876838 768 771 sog paper 50955509 55 66 ST CLAIR LLL L AND JRJ R JOHANSEN 1993 introduction to HuSTEHUSICDIDr FE 1961 1966 die kieselalgenKiesel algen deutschlandsdeutschlandeDeutschlands the symposium on soil crust communities great osterreichsosteireichsOsterreichs und deidelder schweiz volume 7 part 3 in L basin naturalist 53153 1 4 rabenhorst editor kryptogamen flora von deutsch SHIELDS LML M AND F DROUET 1962 distribution of ter- land OstosteneichOsteosterreichoesterreichneichnelcherreich und der schweiz reprint 1977 otto rerestrialstrial algae within the nevada test site american koeltz science publishers konigstein germany journal of botany 4954749 547 554 816 appp SOIL conservation SERVICE 1972 soil survey laboratory JOHANSENJOIIANSCN JRJ R 1993 cryptogamic crusts of semiarid and methods and proceduresproce duiesdules for collecting soil samples and lands of north america journal of phycology USDA SSIR 1 US government printing office 2914029 140 147 washington DC 63 appp 199811998 microbiotic CRUSTS OF BAJA california 311

SOIL SURVEY STAFFSTAFE 1962 soil survey manual USDA agri- WILLIAMS JDJ D JPJ P dobrowolski NEN E WEST AND DAD A culture handbook 18 reissue USU S government GILLETE 1995 microphytic crust influence on wind printing office washington DC 503 appp erosion transactions of the american society of starkstlleSTARKS TLLETL LE SHUBERT AND FR TRAINOR 1981 ecol- agricultural engineers 3813138 131 137 ogy of soil algae a review phycologia 206520 65 80 WILLIAMS JDJ D JPJ P dobrowolski AND NEN E WEST 1995 TER BRAAK CJFC J F 1986 canonical correspondence analy- microphytic crust influence on interninterrillll erosion and sis a new eigenvector technique for multivariate infiltration capacity transactions of the american direct gradient analysis ecology 67116767 1167 1179 society of agriculturalofagricultural engineers 3813938 139 146 1987 the analysis of vegetation environment relationships by canonical correspondence analysis received 2 september 1997 vegetatiovegetation 696969 69 77 accepted 25 march 1998 great basin naturalist 584 0 1998 appp 312 327

RESPONSE OF understory SPECIES TO CHANGES IN PONDEROSA PINE STOCKING LEVELS IN THE BLACK HILLS

daniel W ureskluresk1uresky and kieth E SeverSeversonsoni1

ABSTRACT the objective of this study was to test the hypothesis that therethele are no differences in understory production by species due to stocking levels of pinus ponderosa ponderosa pine understory production was estimated by species on 3 replicates each of 8 growing stock levels ranging from clearcutsclearcuts to unthinned stands in both sapling and pole sized pine stands 48 plots over 3 nonconsecutive years all stands were approximately 70 yr old when thinning treatments were applied production of many herbaceous species especially agropyron sppapp wheatwheatgrassesgrasses and carex sppapp sedges declined as growing stock levels measured in terms of basal area of ponderosa pine increased while trends in total production were similar there were specific differences between sapling and pole stands sedges and oryzopsis asperasperfoliafolia roughroughleafleaf ricricegrassncncegrassegrass produced more in sapling stands whereas danthoniaDanthoma interintermediaintermedialmedia timber oat grass was more abundant in pole stands shrub production dominated by arctostaphylos udauwuva ursi bearberry was relatively consistent across all stocking levels except unthinned although the total number of species declined as pine basal area increased a few species such as linnaea borealis twinflower and shepherdia canadensis buffaloberrybuffaloberry were found only under relatively dense pine canopiescanopies while floristicflonstic species richness was greater at lower stocking levels of ponderosa pine the total number of species would be greater if all stocking levels were present

key words understory ponderosa pine black hills plant species biomass production stocking levels floristicflonstic diversity

understory overstory relationships have and ecosystems and the ecological processes been studied extensively in pinus ponderosa that connect everything in ecosystems practi- ponderosa pine stands in western north cal application of principles of ecosystem man- america ffolliott and clary 1982 these agement will require more specific data in studies have reported the response of forage the case of understory overstory relationships classes grasses and forbs to overstory para- this means a better understanding of responses meters such as canopy cover and basal area by plant species rather than groups of species most have concluded that herbaceous under- such as grasses forbs and shrubs story vegetation exhibits greater growth when the purpose of this paper is to describe the competing overstory is reduced recent how individual plant species responded to work on understory overstory relationships in changes in basal area of pinus ponderosa and the black hills has related production of to discuss how these responses could affect gramingraminoidsoids forbs and shrubs to various grow- plant species richness in the understory of ing stock levels of pinus ponderosa in both pinus ponderosa forest in the black hills this sapling and pole stands uresk and severson study was designed to test the null hypothesis 1989 that understory production does not differ by however conventions governing manage- species due to changes in pinus ponderosa ment of public lands are changing emphasis growing stock levels is being placed on ecosystem management this approach blends social physical eco- STUDY AREA nomic and biological needs and values to assure productive healthy ecosystems kauf- the study was conducted on the black hills mann et al 1994 key features of ecosystem experimental forest about 30 km west of management are the protection and restora- rapid city south dakota the overstory tion of diversitybiodiversitybio which kaufmann et al within the experimental forest is dominated by 1994 define as the variety of life and its pinus ponderosa understory shrubs2 include processes including variety in genes species arctostaphylos uva ursi bearberry prunus

anteienterlcenterscenterentei forgreathllfoiforhorbor creatGreat plains ecosystem research rocky mountain research station school ofminesof mines campus rapid city SD 57701 needling trees weiewelewere classified as shrubs for this study

312 199819981 PONDEROSA PINE understory 313 virginiajavirginianavirginiana common chokecherry berberis 1976 but budget restrictions resulted in sam- rapensrepens oregon grape amelanchier alnifolia pling only 5 levels 0 5 14 23 m2ma halhahai l1 and saskatoon serviceberry and symphoricarpos unthinned in 1974 and 1981 sppapp snowberry common herbs include production of understory vegetation was oryzopsis asperasperfoliafolia roughroughleafleaf ricricegrassegrass measured during august 1974 1976 and 1981 danthonia interintennediaintermediaintermedialmedlamedia timber oatoatgrassmatgrassgrass carex on six 15ism m randomly placed transects per sppapp sedges poa pratensispratensis kentucky blue- plot twelve 30 X 61glemgiemcm quadratsquadquadransrats were ran- grass lathyrus ochroleucusochroochroleucousleucus cream leavinepeavinepeavine domly located along each transect in 1974 and and campanula rotundifoliarotundifolia bluebell most 1976 these data indicated that an increase in of the experimental forest is dominated by the number of quadratsquadquadransrats would provide a better pinus ponderosaarctostaphylosponderosaponderosalarctostaphylosArctostaphylos uva ursi habi- estimate of minor plant species therefore in tat type hoffman and alexander 1987 or HU 1981 25 circular plots measuring 01250.125 m2ma 5 pinus ponderosaponderosasymphoricarposponderosalsymphoricarposSymphoricarpos albaarcalbalarc each were systematically located along 5 of the tostaphylos uva ursi as described by thilenius transects current annual growth of all herbage 1972 was harvested at ground level for each species the experimental forest encompasses about all leaves and terminal portions of twigs to the 1375 ha and elevations range from 1620 m to first node were clipped on shrubs also by 1800 m average annual precipitation is 600 species material was oven dried at 60cgoc60 C for mm 70 of which falls from april through 48 h and weighed weights were averaged and september soils are primarily gray wooded expressed as mean per plot for data analysis shallow to moderately deep and derived from heterogeneous variances were present so metamorphic rock boldt et al 1983 data were transformed using a 109loglo I1 trans- formation analyzed with a one wayglanalysis of METHODS variance years and stands analyzed separately and means separated by tukey s HSD proce- eight growing stock levels GSQGSL of pinus dure in those cases where heterogeneous vari- ponderosa including clearclearcutscuts and unthinned ances persisted means were separated using controls were sampled in both sapling and dunnett s t3ta dunnett 1980 statistical infer- pole sized stands these GSLs were numeri- ences were made at a probability level of 010.1oi cally designated as 0 5 9 14 18 23 28 mm2ma for type I1 error to decrease type II11 error ha 1 and unthinned growing stock indicates plant nomenclature follows great plains all living trees in a stand growing stock level flora association 1986 and van bruggen is the basal area mam2 haglhagiha 1 of a stand adjusted 1985 to account for differences in the average size of trees left in the stand after thinning there- RESULTS AND discussion fore the numerical designation of GSL approximates but does not necessarily equal several variables for this study were con- the actual basal area of the stand basal area of trolled to reduce variability effects of tree size 0 m2ma ha 1 indicated clearcut plots average class were separated all plots were replicated 3 basal areas of unthinned pole stands ranged times on the same soil type and had similar site from 37 to 40 m2ma ha 1 in 1981 while unthinned indices all stands were approximately 70 yr old sapling stands varied from 27 to 33 m2ma ha 1 when treatments were applied and data were three replicated plots were randomly assigned collected over 3 separate years high natural from a total of 48 plots and installed for each variability in the understory among replications GSL from 5 to unthinned in sapling and pole was common despite this control As a result sized stands in 1963 stem diameters were statistical tests often revealed nonsignificant 8 10 cm in sapling sized stands and 15 18 differences despite widely separated means cm in pole sized stands when treatments were the common occurrence of heterogeneous installed three replicated clearcut plots in variances resulted in analysis via dunnett s t3ta each of the sapling and pole sized stands were a conservative test therefore the standard selected and treated in 1966 cut material was error is as important as the mean values in the removed from all plots each of the 24 plots in tables especially values for individual species sapling stands was 010.1oi ha and the 24 pole stands relationships between overstory and general were each 020.2 ha all GSLs were sampled in categories of understory gramingraminoidsoids forbs 314 GREAT BASIN naturalist volume 58

TABLE 1 understory productionpioproduction kg ha 1 oven dried basis for 1974 in sapling sized ponderosa pine stands growing at different tree stocking levels black hills south dakota numbers are means standard errors values within a row followed by differentdiffeidiffee ent letters are significantly different at the 0100.10olo0 10 probability level growingbrowmgrowm g stock levels mam2nr haba 1 clearcutsClearcuts 5 14 23 unthinned

GRAMINgraminoidsghaminoidsoldsoinsOIDS agropyron cabinumcaninumcamnum 204 100 73 43 7744 1 bromus margimarginatusnatus 1111 bromus porten 2222 carex sppapp 239 1231 335 232a 25 sij513 54b5 ab4b 42134 ab2b Dandanthoniathoma interintennediaintermediaintermedialmedia 19 7 47 20 47 2 11 5 212 1 elymus sppapp 222 2 festuca octofloraoctooctqflorafloravlora 1 Kokoelenakoeleriajoelenaeleria pyramidpyramidatapyramidateata 8844 oryzopsis asperiasperlasperifoliaasperifohaasperifoliaefolianolianoila 144 4141ab 304 441 124 8888abb 62 4813 30 53 oryzopsis pungentpungens 333 3 poa pratensispratensis 60 42 42 23 poa interior 333 3 555 5 I1 schizachne purpurascens 1111 total 67667611101lioalloa 806 23p231a 212 glab 818141b41b 364h36 ab4b forbsFOKBS achillea millefoliummillefohummillefolium 57 21 30 14 2211 1111 I1 Agoagoserisagosensserisserls glancaglauca I1111 anaphalis margarimargaritaceatacea 2222 antennaria negneglectsneglectalecta 3322 1111 1111 1 apocynum androsaemifolium I1 I1 1 artemisia sppapp 1 aster sppapp 1 aster laevislaevin I1 1 1111 astragalus adadsurgenssurgens 5555 astragalus aepinus 8844 12 6 1111 2222 astragalus tenellus 1111 campanula rotundifoliarotundifolia 535 3 111144 1 I1 shrubs and total aboveground biomass pro- danthonia interintermediaintermedialmedia produced more partic- duction were previously reported uresk and ularly in clearcutsclearcuts As in sapling stands signif- severson 1989 icant differences by species among GSLs were rare only carex sppapp were found to produce graminoid response aminoidGr more at lower GSLs in 1981 table 6 carex sppapp oryzopsis asperasperfoliafolia and agro- the general responses of gramingraminoidsoids in both pyron caninuscaninum were the most productive sapling and pole stands were similar to other species under sapling stands tables 1 3 black hills studies that included data on species only carex sppapp and oryzopsis asperasperfoliafolia pro- pase 1958 wrage 1994 pase 1958 limited duced significantly more herbage at GSL 14 descriptions to the most important species m2ma ha 1 and below boapoa prapratensistensis and dantho- the rest were combined into an other cate- nia interintermediaintermedialmedia were also common under gory but wrage s descriptions 1994 were saplings but biomass of both was lower only complete pase s study included a variety of oryzopsis asperasperfoliafolia and danthonia intermeintermew sites throughout the black hills and wrage s dia were common in GSLs above 18 m2ma ha 1 study area on soils similar to ours but a differ- trends noted in pole sized stands were some- ent habitat type was about 105ios10.5 km SE and what similar tables 4 6 one of the most 350 m lower in elevation than the experimen- important differences between sapling and tal forest where this study was conducted pole stands however was that carex sppapp and both studies reported responses similar to oryzopsis asperasperfoliafolia produced relatively less those herein but some important differences in pole stands than in sapling stands whereas were evident pase 1958 found that significant 199811998 PONDEROSA PINE understory 315

TABLE 1 continued

browingrowinggrowing stock levels mamm22 ha 1 clearcutsClearcuts 5 14 23 unthinned cirsium sppapp 36 36 delphinium bicolor I1 erigeron sppapp 3333 I1 I1 fragaria vesca 9955 9977 434 3 2 1 I1 21 1 galium boreale 3333 I lactuca sppapp 1 111 lathyrus ochroleucusochroochroleucousleucus 5ala la 27 ab5b 12 taitab 74ab7 aab4ab ap1 11 linnaea borealis 5la 17 17 1p1 oxytropis sericea 1 solidago sppapp 1 taraxacum 0officinaofficinalekooi cinalebinalele 1 1 I1 vicia americana I1 I1 I1 I1 viola sppapp 8 1 12 10 1 1 1 I1 I1 I1 zizia aptera 81I1 11 11 11 total 134 3981 102 16ab16 37 M14a1 34 22lb221b 84b8 41

SHRUBS amelanchier alnifolia 3322 1 I1 arctostaphylos uva ursi 244 74 205 56 293 78 2551148 50 7 berberis rapensrepens 8888 populus tremultremuloidesoides I111I1 1 I1 1 I1 rosa woodsiiwoodsie 7733 757 5 4433 4 1 111 I1 rubusidaeusrubus idaeusidakus 28 23 23 7 545 4 shepherdia canadensis 9999 spiraea betulifoliabetulifoha 222 2 1 I1 I1 1 424 2 111 1 11 1 1 1 symphoricarpos sppapp 18 8 772112 4433 313 vaccinium scopariusscoparium 21 21 11 total 303 10881 244 571 308 7611 305 20a20s1 54 61

GRAND TOTAL 1113 lisailsa1 1152 196196s 557 138b138db 420 571 98 licHG

amounts of oryzopsis asperasperfoliafolia persisted under common gramingraminoidsoids in pine cover class 0 19 moderately dense canopy cover 40 59 19 27 0 9 m2ma ha I1 basal area wrage 1994 listed inm2ma2 ha 1 basal area and dense canopy cover boapoa carex sppapp and the exotic bromus inennisinermisinermia 60 71 28 33 m2ma ha 1 basal area which smooth brome as being most common in open compared with our findings pase 1958 and stands 30 pine cover 14 m2ma hatlhahaci1 basal wrage 1994 used canopy cover in their area we found agropyron caninuscaninumcaninum carex papers we converted these to basal area esti- sppapp and oryzopsis asperasperfoliafolia most abundant mates using bennett s 1984 model canopy in 0 and 5 GSL sapling stands and danthonia cover 051baft2 ac 1 1941.94 r2ra 0830.83 carex sppapp and agropyron caninuscaninum most wrage 1994 did not find oryzopsis asperasperfoliafolia abundant in pole stands in his plots both pase and wrage noted that we considered 11 species as uncommon in danthonia oatoatgrassmatgrassgrass was most abundant under sapling stands defining uncommon as produc- intermediate pine canopiescanopies we found the ing 5 kg ha 1 in 3 stands over all years same relationship under sapling stands but in tables 1 3 included among these were bro- pole stands danthonia was more abundant in mus marginatemarginatamarginata mountain brome schizachne clearcutsclearcuts than in other GSLs poa the most purpurascens false melic oryzopsis pungentpungens abundant grass component in open stands mountain ricricegrassegrass and the exotic agropyron studied by both pase and wrage was common cristatumcristatum crested wheatwheatgrassgrass there were 10 but subdominant in this study pase 1958 uncommon grgraminoidaminoid species in the under- listed poa prapratensistensis sporobolus heteroheterolepislepis story of pole sized stands tables 4 6 two of prairie dropseed and carex sppapp as the most these bromus marginatusmarginatus and oryzopsis 316 GREAT BASIN naturalist volume 58

tat3 illtiltix y j dakota a 1 t- v 219 2401 oi IV 01 unthinned 7 11 2 2 60 S 1 1 1 1 1 1 1 1 1 1 v 1 c f 1T 1 I1 I1 1 oi 1 a 14i 42 tlIV tlIV 1057 V V 2 I1 IV 2 7 south

hills

& black 111 191 01 T i 01 01 10 00 7f 2 2 2 5 1 1 1 1 1 1 1 28011 0 1 1 1 10 CO 260 390 01 1 0 71 10 IV 01 3 01 co 2 V 6 V 0

levels

stocking 10 18bc00 141 1 101 1 1 1 111 1 151 0 i 10 CO i i CO 4f 5 4 3 1 level 2301 1 1 1 1 1 1 1 1 I1 1 H 1 tree 1 00 1 1 i 00 0o 1 41 67 1 10H 127 0 1 1 1 20 IV 8 t 0 V lh0l 01 8 6 6 IV IV V oi

probability different 1 74abe1 aalmazoi t 00 CO tat1 ha 1 1 1 at 171 43 I i 28 01 co i CO i 1- eat010ewt 00 905 01 4 201oi0.1 t- 1 2ol01 1 3 1 7 18 1 1 1 1 1 1 1 1 1 1 I1 1 1 1 m2ma 1 00 00 10 t co iroCO i 1 01 i 1 00 01 1 tat0 ol 0 1 18 28 55 1 36 growing thezaa i i tsi m IV 107 4 3 216i 6 1 2 1 8 2 CO 01 1 01 aia i at levels

stands different 0 stock0 01 187abt c3ca CO 00182 00 1 10 131 1 1 14 1 200 0 1 i CO 1 1 i 0 bd 1 1 0lal01 3 7 2oi 6O 3co 2oi 9cn 14I1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 pine a I1 I growing 1 1 05 t 00 1 01 1ta 6D co oi co 0 1 77 f 10 171 23 1 26 49 1 3 I 01 i oi298 V 3 45410 0i 2 IV 2 oi 2 01 t1 significantly 0 f ponderosa

areaaeaau 17 oi ID in 79ab03 OT f 1 141 22 37 36 45 1 1 99 IV oi CO CO t f 00OO r 03 ol01 01 01 01 sized 1 9 8 4 1 9 2 3 2 2 9oi 1 1 1 1 1 1 1 1 1 1 I1 1 1 1 1 1 1 1 letters 10 f- co 01 ti010.1 CO CT 1 0 1 01OI 01OI CO 01OI 55 87 83 121 30 29 99 83 2 1000 10 eco00 00 2 i rhe0340ewhtl 0CO 4 3 01 V 05 IV 2 9 4 00 2 01 1 tco sapling

different in 00 22801 01 100160 81a 05 00 i 1 160 rl 19 111 1 64 22 1 28 amle1976 by 201 10 01 01 ihi H 3CO I 00 H 201OI 01 60 r 1i 510 510 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 i 1 i 1 1 I1 CO 1 0 1 1 1 01 10 CO eormol 67 i 50 37 41 12 1111 i for 2 zelwel101 26110 10 311raai 3 io0 IV 696796oi co 4 V 5 113i 1 1 1 followed 0i 01 CO 1

basismemmo

lokrow dried a

within 247at i 1 6908902900 t i 1 oven cuts 03 0 00 05 05 1 1 191 81 200 46 i 18 39 i 0 i t 31CO t 1 00 01 01 f 1 i 01 CO 4 6O 201 4 4 clearcutsc3ca 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 obearClearW 1 10 to ihi H 01 en i- co fi c 1 c v 1 1 1 191 12I1 15I 31 15 36 71 1 34 tl 144 595 2 867 133areCO 4 i CO V 2 i CO i values V co i V I amil tt 0 0 G i 10 00 i 1 kg

errors production g

standard a S folia

androsaemifolium 1 capitatum media elzataelw 1 S folia leucus1 a rotundifolia lecta surgens11 b ifera 1 intermedialintermedia 1 folium adsurgens asperifoliaasperifoliae bicolor caninuscaninum neglectsneglecta 1 rotundi ochroleucousochroleucus understory 1 1 subtrinervis 1 admasmeans pungentpungens g SP millefolium stoloniferas S 5 i asperif5 inter pyramid grassmaxom aumneg stolon ll ochro il vesca ad alcsrrssr boreale 1 8 mille appspp tensis ste appspp arezoezoo camatacomata its g siisit S s appspp lllj&s s pratensiss s s laevislaevin OIDS ili y & e ff g g 2 c 0 a a s a chenopodium g 0 pra campanula delphinium agropyron eleria ble antennaria apocynum danthonia oryzopsis unknown astragalus 10 1 oryzopsis siti lathyruslo slcerigeron solidago 1 koeleria fragaria graminoids agrostis 1 achillea cirsium 1lillj1lt1 galium j 1 1 TABLE I sili 1 carex 111 maunz j stipa 11aster numbers totalaczel GRAmIN poa l aaa i giKo FORBS 0laiiitlaillaiilaimitit lliiaqi3 q0qa 00 PONDEROSA PINE understoryq

ja u t 1b t 00 3 114 138cCO 1 ab 1 ill 1 1 9cac 1 OS rh 7 7 1 unthinned 1 1 4 1 1 1 S 1 1 1 1 1 1 a i i 15 1 0 1b os 1 1 1 I7 660260 1 2 ab 1 4 10269mem 341 oq V 01 CO

i 1b oi 0 327c1ca ab 329oq 342bT 8cac00 CO V H CO CO 00 1 1 oi28 1 1 1 1 1 1 19OT 1 1T i 1 1 7 V 68778700oo IV 1 2 79105 850g i I t 00

1 327abc1 libalibc 344 1 t 330CO0 01 1 f 10 1 14 i CO CO 0i ab2b01 I1 i CO CO CO 3 7 1 il 1 1 1 23 1 1 1 1 1 1 1 0011 1 CO CO 1 CO t 01 43 101 14I IV 4 553 0i 6 7 3 1 597 10 oi 1202 10 10 c1ca0

bca haalha7l 16abe lb laracCO & 00185 178 113agr i 1 1 1 s i i CO i i V ab4b co 00 1 3 1 3 181 1 1 1 1 1 1 1 1 1 m2ma 1 1 1 CM 1 1 01 1 OS 0 CO i D 10 I1 011 1 2 I 5910 6 1 7 6 5 0 1243 I 126710 1542 01 01 10f 1 1 1 1 levels L

0 stock0 bc 0 S 16abc 0 n 00 0.0 c1ca326 bM 3 331CO 128co i 1 bc CO i co co 10 ab4b CO 1 tat1 2cl 3 3 5io 1 1 5 ff 14I 1 1 1 1 1 1 I1 1 1 1 growing 1 N CO 10 co 1 oi t r 2 3 64 5 053753 V 9 4 7 668778 0 ID 10 1296ammeCT i 1 s 01 0 1

rt 10 280ab mew n ID 65525510 habmama a 10 233mremee 6aaa 85 1 510 510 800oo 4 00 60 01 3CO CO to QO 1i 01 1 1 1 OS 1 1 1 1 1 1 1 1 1 1 1 9 1 1 io 10 c1ca 0 OS I1 CO 1 10 10 10 CO 10 5 5 22 4 306ehmreq 9 3 0i 1 45 85 23 1901 wema00 IV 01 M lema1081 00 101255 CO0 f os0 01 1 1 1 1 01 1

H M 38ab00i 4111 ama01 leabl2ab01 ci010.1 1 70a l 1 1 121 54 g1 3CO 510 201 1 CO 1 i i i 10 1 1 1 1 1 in 1 1 1 1 1 1 5 1 CO 0O 1 10 1 ln 00 0 OS 111 0 0 1 14 3 6 1 p 181 29 i 5 246 i V 1095lemmwemmos oi 115610 2199os 01 01 1 c1ca

rt tat1 Q d 1 1 196ab cuts 1510 mmampaCO 35410 i 1 aab3aba geagem 10 411 3 i 81 01 151 9os 2ci CO CO 00 CO 201 5 clearcuts 1 1 1 1 is 1 1 1 1 1 1 1 1 Clearai 1 1 i co 0 1T ao os ci 10 20 17T 1 1 V tlV 010 13 696296 V 660760gmeva0 98 9 1700 24f 2 964mew 0 v3 03 01 212601 G 01 1- 0

ursiurmiarmi2 2 2 5 S d lei 2 0 S s appsppa s uvaagaagu oides canadensis a alnifolia 1 1 S scoparium 1 scoparius officina g continued e 0 nianaS S a jaj3 1 tremuloides ri i rapensrepensi I shrubmalum TOTAL forbs virginiajavirginiana rapensrepens 1 tremul 0 americana continued i virgi idakusidaeus 0 aduncaaldunca hisdjsers woodsiewoodsii Q t1ta symphoricarpos ci 1 1 arctostaphylos 1 amelanchier iliiiiji 2 1 1 11 1 GRAND shepherdia 1 taraxacum t M vaccinium unknown i unknown trifolium t i berberis 1 populus prunus TABLE 1 1 1 rubus1 1 vicia viola2.2 3total 11 rosa total 0 h SHRUBS 3 FORBSg E R iai2 ES sjst on cc 318 GREAT BASIN naturalist volume 58

tableTABLL 3 understory production kg ha 1 oven dried basis for 1981 in sapling sized ponderosa pine stands growing at different tree stocking levels black hills south dakota numbers are means standard errors values within a lowrow followed by different letters aledledieare significantly different at the 0100.10olo0 10 probability level growingg stock levels mi2 ha 1 clearcutsClearcuts 5 14 23 unthinnedunthmned cramingraminoidsgraminoid&GRAMIN oldsOIDS agropyron cristatum 11 agropyron spicaspicatumtum 9999 agropyron caninuscaninum 282 48 74 19 24 24 646 4 bromus margimarglmarginatusrnarginatusnatus I1 carex sppapp 508 23223211 429 igoa 105 5656ab 26 1311 15 1311 danthoniaDanthoma interintermediaintermedialmedia 48 8 1411412727 49 16 15 11 434 3 festuca ovina I1 I111I1 Kokoeleriakoelenueletiaeleria pyramidpyramidatapyramidateata I1 1 3333 oryzopsis asperiasperlasperifoliauspenfohaasperifoliaefolia 17611saisab811 393 491 237 121a121st 70 17b 59 2412411 oryzopsis pungentpungens 2222 111 foaboaeoapoa pratensispraprutensisprutensis 252 67 111 I1 foaeoapoa interior 27 18 16 7 1 unknown grassgi ass I1111 23 0 2222 I1 total 129712972631263 1089 1421 424 177 b 11815h118 isb 79 351

FORBS achillea millefoliummillefolium 88 42 52 17 949 4 333 3 221I1 Agoagoserisagosensserisserls glaucaglanca 14 6 antennaria negneglectaneglectslecta 222 2 5533 1111 212 1 1 I1 111 apocynum androsaemifolium 1 323 2 212 1 545 4 11I1 astragalus canadensis 3322 astragalus adsurgensadsurgens 20 4 17 6 4433 3333 222 2 astragalus sppapp 1111 I1 astragalus tenellus 12 6 6633 5555 5555 555 5 campanula rotundifoliarotundifolia 25 5 8822 111 I1 I1 cirsium vulgare 17 13 111 1 I1 delphinium bicolor I1 erigeronengeron sppapp 555 5 1 I1 1 I1 fragaria vesca 33 2 2411isab15 77511ab5b 33111ib 22111ab1b pungentpunpungensgens also occurred in sapling stands oth- agropyron cristatumcristatum and phleum prpretensepratenseatense ers included Kokoelanakoelariaelaria cristatacnstatacristanacristata prairie june occurred only in a single stand each boapoa grass which was common in sapling stands prapratensistensis was not considered exotic because it boapoa arida plains bluegrass bromus porten occurs in both naturalized and native forms in plains nodding brome and the exotic phleum this area great flora association 1986 wrage reported 3 exotic grasses agro- prpretensepratenseatense timothy all uncommon species 1994 pyron rapensrepens agropyron intermedium and occurred in pine stands stocked at 14 m2ma ha 1 basal area wrage 1994 appendix D bromus inennisednisermis altogether we found and identified 22 dif- had 8 grass that contributed 1 species ferent gramingraminoidsoids in the pine understory in foliar 3 stands over all years and all cover in this study thirteen were common to sapling pinus classes agropyron ponderosa cover and pole sized stands while 3 and 2 species A intermedi- rapensrepens quackquackgrassgrass intermedium were unique to each respective size class pase ate wheatgrasswbeatgrasswheatgrass boutelouaBoutelotta curtipendula side 1958 reported 38 grgraminoidaminoid species and oats grama calamagrostis rubescentrubescensrubescens pine wrage 1994 listed 23 reedreedgrassgrass muhlenbergia torresitorreyi ring muhly an average of 9 grgraminoidaminoid species occurred schizachne purpurascens sporobolus asper in GSLs 0 5 and 14 m2ma ha 1 over all years rough dropseed and stipa spartenspartea porcupine this was higher P 0050.05oos than the 6 and 5 grass species found in GSL 23 and unthinned stands three exotic grasses were found under the respectively uresk and severson 1989 the pine overstory one agrostis stoloniferastolonifera red- mean number of species present in pole top was common but scattered the others clearcutsclearcuts 11 and GSL 5 m2ma ha 1 10 was 199819981 PONDEROSA PINE understory 319

TABLE 3 continued

growinggrowirgrowerig stock levels mamm22 ha 1 clearcutsClearcuts 5 14 23 unthinned galium boreale 18 13 11I1 1 222 2 I1 hieracium canacanadensedense I1 111 1 insirising missouriensismissounensts I1 lathyrusLathy riisytis ochroleucusochroochroleucousleucus 40 20 11728117 28 68 33 39 13 10 6 linnaea borealis 1 222 2 I1 lupinus sppapp 3333 7744 monardafistulosamonardomonarda fistulosefistulosa 11I1 I1 I1111 polygonum persicanapersicariapersicanopersicana 11I1 I1 smilacinaSimlacina stellata 222 2 solidago sparsiflora 7744 6655 1 11 11 I1 taraxacum officinale 111I1 I1 trifolium rapensrepens I1111 444 4 11 11 I1 vicia americana 57 35351 12 213 99139 ab9b 2 221 1llhliblahib viola aduncaaldunca 25 1 18 3 222 2 111I1 I111I1 total 369 99 276 663 117 5252abe 92 lo10lobjob 26 16c

SHRUBS amelanchier alnifolia 212 I1 I1111 111I1 1 arctostaphylos uva ursi 604 179 820 95 872 427 821 333 205 111 berberis rapensrepens 888 8 444 4 populus tremuloidestremultrernuloidesoides 1111 333 3 rosa woodsiiwoodsie 25 2 21 12 13 8 11116ilg 6 333 3 rubus idakusidaeus 130 117 61 24 22 13 1111 333 3 shepherdia canadensis 999 9 777 7 spiraea betulifolia 1111 1111 727 2 848 4 424 2 symphoricarpos sppapp 38 10 12 4 20 13 535 3 323 2 vaccinium scopariusscoparium I111I1 total 802 307 915 126 938 428 866 326 230 107

GRAND TOTAL 2450 1681 2280 102a 1478 596ab596 1076 330330111lly111uly 334 146114611

greater FP 0050.05oos than the 5 5 and 3 species tables 1 3 but few significant differences found in GSLs 14 23 and unthinned stands by species among GSLs were noted lathyrus respectively uresk and severson 1989 wrage ochroleucusochroochroleucousleucus produced more herbage in mid 1994 counted 20 19 and 16 grass species in level GSL sapling stands 5 23 in 1974 table his open tree cover 30 14 m2ma hahaihal 1 basal 1 and fragaria vesca woodland strawberry area intermediate 30 60 14 27 m2ma haihahal I1 and vicia americana american vetch produced and dense 60 27 m2ma ha 1 pinus ponderosa more in clearcutsclearcuts in 1981 table 3 but no dif- stands respectively ferencesferen ces were noted in pole stands pase 1958 reported trifolium repensrapens white clover fra- response forb garia vesca and achillea millifoliummillifolium to be forb response relative to GSLs was simi- abundant in open stands but only lathyrus lar to that of gramingraminoidsoids except that forbs pro- ochroleucusochroochroleucousleucus and solidago sppapp goldenrod per- duced about 30 less forage tables 1 6 no sisted under moderately dense pine canopiescanopies single forb species dominated any GSL of artemisia ludovicianaludoviciana white sagewortsagewort gly- either size class however achillea miuifoliummillifoliummillifolium cyrrhiza lepidotelepidotalepidota wild licorice psoralea argo common yarrow campanula rotundifoliarotundifolia and phylla silverleaf scurfscurfpeapea and monardafistumonarda fistubistu lathyrus ochroleucusochroochroleucousleucus were the most common losa wild bergamot were the most common forbs found in pole stands tables 4 6 achil- forbs in wrage s 1994 open stands but only lea millifoliummillifolium and lathyrus ochroleucusochroochroleucousleucus were glycyrrhiza lepidotalepidotelepidota persisted into intermedi- the most abundant forbs in sapling stands ate stands forbs were scarce in his dense 320 GREAT BASIN naturalist volume 58

TABLE 4 understory piproductioneduction kg ha 1 oven dried basis for 1974 in pole sized ponderosa pine stands growing at differentditteldittei ent tree stocking levels black hills south dakota numbers are means standard errors values within a row followed by different lettelletterss are significantly different at the 0100 10 probability level

growing3 stock levels mam2m 2 ha 1

clearcutsClearcuts 5 14 23 unthinned graminoidsGRAMIN OIDS agrostis stoloniferastolonifera I1 agropyron spicaspicatumtum 3322 1 agropyron caninimcammacammm 155 89 3322 I1 bromus margimarglmarginatusmargmargmatusnatusmatus I1 bromus porteriponteri 222 2 I1 carex sppapp 747 4 15 9 212 1 10 5 I1 danthonia interintermediaintermedialmedia 322 21 135 31 46 17 5522 Kfestuca octofloraoctoflora 20 10 I1 festuca ovina 1 221I1 koeleriaKo eleria pyramidpyramidatapyramidateata 3333 oryzopsis asperiaspenasperifoliaaspenfoliaasperifoliaefoliajolia 9988 29 24 17 15 4433 poa appspp 10 4 poa pratensispratensis 7744 4444 total 53853876176a 193 45111 48 lsiisiisit1511515 32 17be17 5 414 korbsFORBS achillea millefoliummillefolium 39 11 6633 1111 1111 I1 antennariaantennataantennanaAntennana negneglectsneglectalecta 7744 9999 9966 3322 apocynum androsaemifoliumandrosaemifohwn 1 1 1 1 1 I1 astragalus alpinusalpmusaepinus 13 3 171111 I1 I111I1 astragalus tenellus 4433 333 3 3333 astragalus sppapp 3333 1 111 I1 I1 campanula rotundifoliarotundifoharotundifolia 35 22 6622 I1 cirsium arvensearmense 14 14 cirsium vulgare 15 8 erigeronengeron sppapp 2211 4411 111 I1 I1 fraganadraganafragaria vesca 3322 1 1 1 stands those most common were callumgalium vulgaris butter and eggs was found only in boreale northern bedstraw and smilacina pole clearcutsclearcuts the other 4 polygonum persi- stellata false solomonSolomorsmois s seal caria lady s thumb tragopogon dubiusdubics goats many forb species occurred under the pine beard sisymbrium altissimum tumbling mus- canopy but they contributed little to total tard and rumex acetacetosellaosella sheep sorrel aboveground biomass we categorized 22 were rare A large amount of chenopodium species as uncommon using uncommon as capitatum strawberry blite was found in I1 we defined it for gramingraminoidsoids wrage 1994 plot of 1 replicate of sapling GSL 9 m2ma ha 1 in also noted many forbs that were generally 1976 table 2 it is unknown whether this scarce if we arbitrarily classify uncommon species is indigenous or exotic great plains species in wrage s study as those having 1 flora association 1986 wrage 1994 listed 4 foliar cover in 3 pine cover classes over all exotic species cirsium arvense convolvulus years 18 forb species would be considered arvensearmense field bindweed taraxacum officinaofficinalele uncommon wrage 1994 appendix D and tragopogon dubicsdubius all were classed as nine of the forbs found in the pine under- uncommon except convolvulus venseararvensearmense which story are exotic species only 2 however were was common in open stands common taraxacum officinaofficinalele common dan- although our aboveground biomass esti- delion and trifolium rapensrepens these species mates of forbs were less than for gramingraminoidsoids are both ubiquitous over north america two there were more forb species than grgraminoidaminoid others both noxious weeds cirsium arvensearmense species fifty three forbs including 15 identi- canada thistle and C vulgare bull thistle fied only to genus grew in the understory 25 were common only locally another linaria identified species were common to both 199811998 PONDEROSA PINE understory 321

TABLE 4 continued

growingbrowingrowin g stock levels mamm22 ha 1

clearcutsClearcuts 5 14 23 unthinned

galium boreale 1 1 iris missouriensis 11I1 I1 I1 lactuca sppapp 525 2 313 1 8877 I1 lathyrus ochroleucusochroochroleucousleucus 767 6 linaria vulgaris I1 oxytropis sericea 6644 979 7 1 potentilla sppapp I1 solidago sppapp 333 3 1 1111 taraxacum officinaofficinalele 1 1 Tnfoliumtrifolium rapensrepens 555 5 23 23 1 vicia americanaamencanaamencana 222 2 I1 I1 1 viola sppapp 51 323 2 total 161 1581 86 35ab 28285bab5b 227b22 ab7b al1111l

SHRUBS amelanchier alnifoliaalmfoliaalmaimfoliapolta 1111 555 5 1 arctostaphylos uva ursi 209 7 305 94 306 72 135 35 66 58 berberisberbens rapensrepens 19 19 betula papyrpapyriferaifera I1 juniperusJumperus commoniscommuniscommunis I1 populus tremultremuloidesoides 21 21 I1 rosa woodsiiwoodsie 10 8 1 2222 I1 I1 rubusrubusidaeusidakusidaeus 49 17 10 6 shepherdia canadensis 10 10 spiraea betulifoliabetuhfolia 11I1 1 323 2 1 I1 1 11 1 symphoricarpos sppapp 777 7 333 3 222 2 2222 total 299 48 347 77 3113117070 149 43 67 58

GRAND TOTAL 998 84a84s1 626 4613 387 87bt87be 203 34b34bt 72 58

saplings and poles while 6 and 7 species were shrub response to the sapling and pole unique underunderstoriesstories mean shrub production was similar across pase wrage respectively 1958 and 1994 all pinus ponderosa sapling stocking levels in noted 63 and 41 forb species in their respec- 1976 and 1981 and in 1974 pole stands in the tive studies remaining sapling 1974 and pole 1976 1981 we found an average of 18 forb species stands all growing stock levels produced more over all years in clearcut sapling stands which shrubs than unthinned stands in general shrub was significantly higher P 0050.05oos than the production tended to be somewhat higher at 15 13 and 13 species found in GSLs 5 14 mid levels than in clearcutsclearcuts and much higher and 23 m2ma ha 1 respectively eleven species than in unthinned stands although differences were found in unthinned stands fewer P among lower GSLs were not significant tables 0050.05oos than in clearcutsclearcuts uresk and severson 1 6 arctostaphylos uva ursi accounted for shrub production all stands 1989 trends were similar in pole sized stands most in regardless of size class or stocking levels but differences twenty one species were found in clearcutsclearcuts were not significant except in pole stands in and 18 in GSL 5 which was higher P oos0.05 005 1981 table 6 production of arctostaphylos than the 12 and 13 found 14 and 23 in GSLs uva ursi was 75 99 of total shrub production fewest species 5 were found in unthinned in sapling stands and 70 99 in pole stands pole stands uresk and severson 1989 wrage lower percentages tended to occur in clearcutsclearcuts 1994 counted 36 29 and 24 forbs in pinus higher proportions of rubus idaeusidakus red rasp- stands with open intermediate and dense berry and symphoricarpos sppapp snowberry canopy covers were found in clearcutsclearcuts rosa woodsiiwoodsie and 322 GREAT BASIN naturalist volume 58

g num- U 1i i V z ff co fe 1 1 unthinned 3 2 1 1 1 1 1 1

1 T s i s 1 1 1 10 1 1 01 dakota Iv 4 4 0 Iv IV 2 nj p Q 5 south3 0 c1ca

ls 1201 lmo 1 1 1 11 1 1 IV 00 1 10 01 hi 1 1 1 5 28 1 1 1 2 01 1 1 1 1 1 1 1 1 1 1 1 1 3201 1 10 i i 1 00 1 1 U V 1 V CO V 35CO 1 1 IV 8 IV V 100 blacknj S

1levels jg oi 2waw& 191 bc 1 3CO 510 01 10 10 1 4 1 1 5 1 5 23 1 1 1 stocking a 1 1 1 1 i i 1 i i 10 1 00 10 10 60 37CO V 48 1 1 1 1 1 10 5 5 I V 5 V 5 IV IV IV 01 1 sw evel u u treealaealaou ty

s abili Q probabili awil 1 1 1 1 different n 00 46bctat0 1 1 23 28 1 01 H CM CO V V 1 1 12i alomprobarom c1ca co i N 00 6 2 7 3 18 1 1 1 4 3 1 1 1 1 1 1 1 1 1 1 1 1 1 9 a m2ma jt in 00 CO p 1 1 1 1 25 170 0 1 at 1 I 101 6 0 8 ai 0 1 I N 2 3 114 1 IV 1 IV IV 6 4 IV 1 7 35co 1 1 I bce levels 5 0 growing the 6 &b atni 0 ab stocks 1 t 18bfe 01 i 1 17 1 1 V 1 121 CO 1 1 1 11 stands bc 1 N 11 1 i 1 different 3 4 1 1 1 1 4 14 1 1 4 U 1 ai 1 1 1 1 1 1 il 22 I 1 kfcgkocg growing 0 01 10 10 t 1 10 00 D o 0 1 5 05196 205805bem 4 7 5 0124 1 1 18 t 1 11 0 pinegacoa U 01

s significantly ponderosa I1 CM bbb& co co 1 27 121 13i 13 a 800 7 0 3CO 60 01 1 01 i 01 1 2 1 0 1 1 1 1 1 1 1 1 2 & U 05 1 1 1 1 ii i i 1 1 1 mgo are 9 1 1 1 14 00 0 CM 1ii 1 CO 60 01 1 01 co n 1 tat3 V 8 116aze1 12 V 3 162 4 2 20 1 1 30 O 0 oii V V CI 3 CO 1 1 01 sizeda s 1 K D letters achapoleacmai0 & s

inS 1 1 120ab different 22 21 5800 00 0 CO 10 M 38 25 01 15 12 1 1 co CO 23 r 101 10 IV a N in 1 co 3 3 01 6 4 0 1i 0 1 1 4 1976 1 1 1 1 1 1 1 1 1 1 1 1 510 1 I 1 1 1 1 v 1 2 M co 01 00 10 i CO 1 t 1 t go 52 32 gmt 38 27 10 10 1 1 1 by 1 10 CO 001800 2 CO 5 01 3 ehl341 2701 15 7 4 0124 8 I7 1 7 co36 7 folforuzl0 11 CO 0

basis3 followed cfi afiufiU dried 2lovrovrow C ai cutsadem3 11 20 69oi 70 30a 1901 01 1 21 1 10 34 10 0 0CO 1 12 1 24 01 CO CO 0i onanowanoven 20 M 5in t 0 0 3 3 CO 10 20 00 01 clearcuts 1 1 1 1 1 1 1 2 1 1 1 1 1 1 5 8 1 n3na 1 1 1 1 1 within y ClearU CO i 1 01 CO co H CO CO 71 0 10 1 10 t 00 1 10 1 52 70 23 0 3 0 0 1 633233 25 37 88 20 10i 25 26 1 10 406 f 001 co 12101 a 5 3 co 00oo 010 IV 0i 3 01 01 3 1 0t 10 1 maiamiamia Q D kgbc values3

C 0 w gerrors production 0 a 1 2 0 &i320 0

androsaemifolium folia standard media t S 1 folia leucus 5 0 rotundifolia 1 azulelecta so intermediaintermedial i S folium 2 ifera surgensadsurgens w caninuscaninum asperifoliaasperifoliae neglectsneglecta bicolor ci rotundi ochroleucousochroleucus subtrinervis understory pungentpungens 1 1stolonifera millefolium S asperi 1 1 inter t grass 1 neg s 1 S ji .1 glancaglauca i cl stolon 1 1 vulgare ochro U 1 ab&b e ad vescaCO boreale vulgaris a c ovina mille 0 1 tensis meansegaasi s S S 3 y S pratensis i 0 s omma S S laevinlaevis OIDS CO i B S oalmala a 111 S E a a t3ta 3 j 1 a astragalus j si a seris S delphinium 5 S agropyron pra campanula 0 danthonia 0 antennaria apocynum geria M U oryzopsis oryzopsis unknown 2 1 H erigeron lathyrus alearemlo graminoids agrostis agoseris 1 fragariafrageria fc S p festuca 3 achillea 0 cirsium s 1 1 1 g p galium i e linariaS TABLE i CQ S SP ie carex 1 stipa 1 total 0 aster TS GRAmIN 1 cl S poagaagea ruoAgo a b ulzFraS s is 1 1 cs cs FORBS s la13l3 urri ca l 1 1 1 lri i Q hah3 c5 11 l 11 bers t U U ll 3 0 fo PONDEROSA PINE understoryz CID t1 CI

0 m D y fi 94b ic v 094 05 960 unthinned 1 1 1 1 1 1 1 H CO i 1 D 1 oq CO fi 3 96 V 2 1000 113wap1 1 & 05 0

.0 0 0.0 CO 1 143b CO i 10 zerbzerm ta145t1 CO w f 1 1 1 F ab5b10 V 1 11 1l1 1 1 1 2800 1 1 1 CO al11 1 rlr l N 1 tat0 10 1 24ci ci422 IV 2 426CO 485co tat1 t f

0 10 155ab 1 155 io 127b 0o 10 CO p ab8b00 1i 1 i 1 1 1i 1 r 4 1 1 1 00 1 1 1 1 23CO tl 1 t t 10 17 1 c i i 1 0 4 27N 2 675elm1 V IV 2 1 00681 jo656756 0 0 f

1 S A 0 16pib11 129be05 maz 23bCO 18600 tat0 CO aal 1 1 260 1 pha t1 i i CO M 11 1 00 CO gl 4ta 1 3 2cl 3 1 1 1 1 1 N 18 1 1 1 1 m2mac3ca i R 1 i 1 1 m 1 0 2610 10 IV 5 IV 1056 6 507 1 4 N 5 100550 6207200CO 0 10 1 33 10 levels a

0.0 stock 0.0 1 30&b& iw 1 pami 1 gli2771 1t l 7 CO i CN D co co0 ml 1 0 1 4 2 6 1 1 6 1 rt 14 1 1 1 1 1 1 1 1

1 i 1- growing 05 1 CO 1 CO t CO 00 0 9 1 173 934COmew 3 7 948t CO 0 Iv 2 Iv 4 1226 0 0 05 CO

to OT 306ab rfiafi 265a10 mrmppmCO 279mim1 52c1ca 1 1 V CO CO 3CO N 10 10 2 CO 8 7 2 5 1 1 1 1 1l1 1 1 1 1 05 1 1 1

9 1 1 1 1 1 1 cr t 0o 1 81 co 0 a 174 6 6 oo 3 co0 7 11 2 IV 00 1220 10 alV 1124ci co 1456ahme ii i

l r 252abCO 00 291arl 36ab10 428CO 10 10 CO 05 1 65 I1 55 23 i 1 0 00 b CO 0 1r 3CO lo10 CO 201OI CO 0 1 6 1 6 8 7 1 1 1 1 1 10 1 1 1 1 1 1 1 1 1 1 1

1 5 1 1 I i 1 CO i 1 D co 00oo 10 r CO CO lo10 05 n13 111 65 84 231 8 zeh164 i I 827 1 3 5 CO 1 6 3 leb0 ol01 to 00 001 1019 1523 1 00 10 0 1 1

0.0

1 220z1b CO en a 132a i 187 CO i 1 0 CO cuts c1ca82a 00 co 131 1 141 1 21 111 I i 11i i CO 1 td V 03 c1ca N 00 3CO 1 905 9 1 1 2 1 1 1 6 1 1 1 1 1 1 2l1 1 1 1 clearcuts1 ctrct3 al21 1 1 Cleara 0 c1ca 1i CD 0 1 00 1 05 82C 1 14 161 21 46 41 1 1 emi687 9 NI i 821 6 V 2 477 8 00oo 00oo V 1931 tf 10 00 05 S 1 1

a ursi2 a n3na 1 1 le appsppa altissimum a oides canadensis & wl alnifolia 1uvaavaawa 1 3 missouriensis dubicsdubius 2 a officina5 S 2 Q g tremuloides2 continued i i TOTAL J rapensrepens golmgorm S g- 1 forb S a lemmas a 1 g i tremul ommmap americana sa 1 continued11c S & g S 2 g g J idakusidaeus e aduncaaldunca aispotentillafruticosa woodsiewoodsii Q iis symphoricarpos u j g g arctostaphylos

amelanchier GRAND 10 sisymbrium tragopogon i 1 shepherdia 1 taraxacum s 3 1 5 unknown 11 i trifolium solidago i berberis solidago populus igssa a s rubus TABLE P vicia viola bolez rosa all totalackan m total 3 0 i 3 SHRUBS FORBS is E aj tityk li c1calltgtt11itl 324 GREAT BASIN naturalist volume 58

tailleTABLE 6 understoryundeidundei story production kg ha 1 oven dried basis for 1981 in pole sized ponderosa pine stands growing at different tree stocking levels black hills south dakota numbers are means standard errors values within a row followed by differentdiffeidiffee ent letters are significantly different at the olo0100 10 probability level growinggrowirgrowerig stock levels mam2m ha 1 clearcutsClearcuts 5 14 23 unthinnedunthmned graminoidsGRAMIN OIDS agropyron caninuscaninum 41 9 26 3 I1 1 aristida purpurpurpureapurpureanea I1 bromus porteriponteri 2222 carex sppapp 83 26a26s 60 231 7 55bcnbc 14 411 licalc1 IC Dandanthoniathoma interintermediaintermedialmedia 651 95 365 60 95 17 30 15 llcI1 festuca ovina 17 8 1 333 3 koeleriaKoeleria pyramidpyramidatapyramidatepyrarmdataata 10 10 4422 I1 oriizopsisoryzopsisyzopsis asperiasperlaspenasperifoliaaspenfoliaasperifoliaefoltafolia 11811 8 24 13 737 3 15 15 oryzopsisoryzopsis pungentpungens 5555 2222 phleum prpretensepratenseatense 111 1 boaroapoa pratensispratensis 89 37 I1 boapoa interior 5511 1111 1 unknown glassgrass 2222 1 I1 1 total 916 811 484 63b633 113 26 61 30cd30ed 43d4 ad3d forbsFOKBS achillea millefoliummillefohummillefolium 72 18 15 3 5533 1 I1 Agoagoserisagosensserlsseris glaucaglanca 5544 1 antennariaantennataantennanaAntennana negneglectsneglectalecta 23 15 777 7 333 3 313 1 1 I1 apocynurnapocynum 1 1 1 111 apoctnum androsaemifolium 11I 5544 2222 I astragalus sppapp 17 9 212 1 111 I1 I1 astragalus adadsurgenssurgens 4422 21 10 25 13 5544 astragalus canadensis 17 14 34 28 9999 I1 astragalus tenellus 5533 7766 5555 13 6 campanula rotundifoliarotundifolia 393988 525 2 2222 I1 cirsium vulgare 111 1 delphinium bicolor 1 erigeron subrinsubrinervissubnnerviserdiservis 10 9 I1 I1 1 fragaria vesca 12 7 7722 111I1 I1 I1 rubus idaeusidakus produced more at the 3 lower to sapling stands and was common in higher sapling GSLs in 1976 table 2 while arc- GSLs tables 1 3 none of the shrub species tostaphylos uva ursi was more abundant in under saplings were categorized as uncom- pole clearcutsclearcuts in 1981 table 6 pase 1958 mon three species all unique to pole sized noted the same patterns of arctostaphylos stands were classed as uncommon juniperus uva ursi abundance but this species did not communiscommoniscommunis betula paperpaperiferaifera paper birch occur in wrage s 1994 study area symphori- and potentilla fruticosafruticose shrubby cinquefoil carpos occidentoccidentalisoccidentalistalis was the most common pase 1958 counted 18 shrub species while shrub noted by wrage 1994 under all pinus wrage 1994 noted only 6 canopy levels juniperus communiscommoniscommunis common there were no differences P 0050.05 in juniper rare on our study site and on wrage s number of shrub species among sapling GSGSLsLs 1994 was abundant on sites studied by pase they ranged from 6 to 8 across all levels with 1958 this species is a predominant con- no apparent pattern in pole stands 8 shrub stituent of pinus ponderosa understory on species occurred in clearclearcutscuts but only 4 species limestone derived soils thilenius 1972 hoff- were found in GSL 5 uresk and severson man and alexander 1987 1989 wrage 1994 did not note any differ- fourteen shrub species occurred under the ences he found 4 shrub species under open pinus canopy 11 of which were common to 5 under intermediate and 5 under dense pinus both sapling and pole sized stands one species canopiescanopies none of the shrubs found in the vaccinium scopscopariusscopahumscopariumaHumanum grouseberry was unique understory of our study or wrage s were exotic 199819981 PONDEROSA PINE understory 325

TABLE 6 continued crowingrowingg stock levels maiwm22 ha 1 Clearclearcutscuts 5 14 23 unthinned

galium boreale 1 I1 1 hedysarum alpinum I1 insiris missouriensis I111I1 lathyrus ochroleucusochroochroleucousleucus 57 27 48 15 34 13 21 11 222 2 linaria vulgaris 11I1 I1 monardaftstulosamonarda flstulosa I1 polygonum pefperpersicariasicana I1 rumex acetacetosellaosella I1 solidago sparsiflora 212 1 424 2 5522 I1 taraxacum officinaofficinalele 242 4 I1 trifolium rapensrepens 91 33 15 12 vicia americanaamencanaamencana 222 2 555 5 7766 323 2 I1 viola aduncaaldunca 19 6 12 5 I1 1 1 total 385 litilaiialia111 189 12b 100 8cac 44 ladl5d15d 663l633631ad3d

SHRUBS amelanchier alnifoliaalmfoliaalmaimfoliapoliapolta 11I1 I1 1 3322 arctostaphylos uva ursi 1110 loga 864 144144abb 673673115blisb 522 gil9119111 26 1818 berbensberberis rapensrepens 34 34 1 juniperusJumperuspenas communiscommoniscomcommumsmunis I1 populus tremultremuloidesoides 93 93 I1 prunus virginiajavirginianavirginiana 11I1 I1 rosa woodsiiwoodsttwoodstn 22 84 222 2 636 3 1 222 2 rubusrubusidaeusidaeusidakus 96 56 36 12 shepherdia canadensis 215 148 777 7 spiraea betulifoliabetulifoha 31 6611 1 1 I 31 21 111 symphoricarpos sppapp 165111 4411 8855 3322 331I total 1344 433 945 ilg116116ab 910 144ah144db 535 9719711 31 19 GRAND TOTAL 2644 35 1620 86b 1123 132b 640112c640 112c 41 ald2ld21d

total understory response son 1989 found more species P 0050.05oos in sapling and pole clearcutsclearcuts 34 and 40 respec- total understory production was variable tively than in mid level GSLs saplings 27 28 among GSLs but generally the greatest poles 22 24 fewest species were found in amounts in sapling stands were produced at unthinned stands 23 in sapling and 12 in pole m2ma ha 1 or less and pole stands at GSL 23 in stands the only species occurring regularly 14 m2ma ha I1 or less stands GSL unthinned in unthinned stands were oryzopsis asperfoaspermo consistently produced significantly less forage lia danthonia interintennediaintermediaintermedialmedlamedia lathyrus ochroleu all 1 6 same than other GSLs tables the cus and arctostaphylos uva ursi tables 1 6 trend was noted by pase 1958 we identified 89 species in the pinus pon- conclusions AND management derosa understory over all stocking levels and implications tree size classes an additional 19 plants were identified only to genus and 6 others were pinus ponderosa is the most abundant and designated as unknowns pase 1958 who sam- widely distributed tree in the black hills it is pled a greater variety of sites found a total of the major climax tree species in 7 of 12 forested 119 while wrage 1994 tallied 70 hoffman habitat types described by hoffman and alex- and alexander 1987 sampled 10 stands in the ander 1987 and a seraiseral or occasional species pinus ponderosaarctostaphylosponderosaponderosalarctostaphylosArctostaphylos uva ursi habi- in 2 others the climax stands occupy about tat type and found the range of the total num- 600000 ha boldt et al 1983 natural regen- ber of species to be 13 29 uresk and sever eration of pinus ponderosa in the black hills is 326 GREAT BASIN naturalist volume 58 aggressive and prompt with high stand densi- munitiesmunities is limited other exotic species appear ties often resulting boldt and van duesen to be uncommon and restricted to disturbed 1974 hoffman and alexander 1987 theref- sites in open areas such as skid areas on clear ore pinus ponderosa often replaces itself when cuts the presence and distribution of exotic the stand is disturbed only 2 of 7 pinus pon- species should be a part of monitoring activities derosa habitat types have potential seraiseral stages A similar status is afforded threatened en- dominated by populus tremultremuloidesoides quercus dangereddangered and sensitive plant species in the macrocarpamacrocarpa bur oak is a minor seraiseral species pinus ponderosa understory none of the species on another hoffman and alexander 1987 found in this study or wrage s 1994 is listed evidence indicates that eliminating or reduc- as sensitive in the black hills fertig 1993 ing the pinus ponderosa overstory will increase most listed species are found in mesic habi- the number and total productivity of under- tats but this should not preclude a more story species this has 2 apparent advantages intensive survey of pinus understory first and most obvious is that such a reduc- most evidence provided herein represents tion is a viable approach to increasing plant a case for maintaining pinus ponderosa stands species richness of the black hills the habitat at a range of stocking levels while floristic type we studied pinus ponderosalarctostaphyponderosaarctostaphy species richness is greater on clearcutsclearcuts than in los uva ursi is not the most floristically rich unthinned stands the total number of plant we could expect even more dramatic responses species would be greater if both clearcut and to occur in those habitat types that have more unthinned stands were present species such species ege g pinus ponderosaquercusponderosalquercusponderosa Quercus macro as linnaea borealis twinflower shepherdia carpa habitat type hoffman and alexander canadensis buffaloberrybuffaloberry and vaccinium sco 1987 table 4 hence each pinus dominated bariumparium grouseberry are more common under habitat type would likely have a unique under- relatively dense pinus canopiescanopies from an eco- story response to canopy changes eliminating nomic standpoint clearcutsclearcuts and very low pinus overstory in those habitat types where pinus stocking levels promote more growth of seraiseral stages are dominated by aspen would available forage for livestock while intermedi- create an even more diverse community struc- ate stocking levels produce more wood fiber ture peripheral evidence suggests that aspen severson and boldt 1977 wild ungulaungulatesungulatedtes dominated stands contain more species than need open areas for foraging and dense conifer pinus or mixed populus tremultremuloidesoides pinus pon- stands for cover intermediate stocking levels derosa aspen pine stands kranz and linder provide only marginal levels of both sieg and 1973 severson 1996 second a floristically rich and productive historical evidence shows that patterns of understory would support a healthier popula- biotic community development in the black tion of herbherbivoresivores ruminants can select nutri- hills prior to euro american intervention were tious diets from a diverse array of plant species more heterogeneous than those found today provenza 1995 odocoileus virginianusvirgimanusvirgivirginianuslanusmanus white primarily due to the influence of fire graves tailed deer for example have the ability to 1899 progulske 1974 turchen and mclaird select a variety of forages that collectively bal- 1975 see also USDA forest service 1994 ance nutritional needs vangilder et al 1982 sieg and severson 1996 although a few large reducing the number of plant species and the dense pinus stands did exist graves 1899 we resulting decline in selective feeding ability examined only influences of mechanical tree would also increase competitive interactions removal the inclusion of prescribed fire would among herbherbivoresivores have produced different results sieg and sev- several exotic species occur in the pinus erson 1996 litter removal by burning would understory some such as taraxacum officinaofficinalele stimulate a faster understory response due to and tnfohumtrifoliumtlifolium rapensrepens are ubiquitous within nat- exposure of mineral soil and rapid nutrient ural systems others like cirsium vulgare C release A different species response in the arvense and convolvulus arvense are desig- understory could also be expected for exam- nated noxious weeds and must be controlled ple nonsprouting species would be less abun- by law although they may be locally common dant and sprouting species relatively more their distribution within pinus dominated com abundant other species such as epilobium 199811998 PONDEROSA PINE understory 327 angustifolium fireweed characteristically dakota and wyoming a habitat type classification invade burned sites great plains flora asso- research paper RM 276 USDA forest service rocky mountain forest and range experiment sta- ciation 1986 tion fort collins CO A serious approach to ecosystem manage- KAUFMANN MR ET AL 1994 an ecological basis foiforooiool ment including concern for health and main- ecosystem management general technical report tenance of natural systems and economic and RM 246 USDA forest service rocky mountain foi-forest and range experiment station collins CO social needs should consider all habitat types fort KRANZ JJJ J AND RLR L LINDER 1973 value of black hills and all seraiseral stages within those habitat types forest communities to deer and cattle journal of special attention must be given to placing these range management 262639626326 263 265 in optimum spatial and temporal mosaics PASE CPC P 1958 herbage production and composition would include the under immature ponderosa pine stands in the black such mosaics presence of hills journal of range management 1123811 238 243 relatively large dense pinus stands which were PROGULSKEPROCULSKE DRD R 1974 yellow ore yellow hanhairhain yellow present in the black hills prior to this century pine A photographic study of a centuiycentury of forest graves 1899 the historical pattern of fire ecology agricultural experiment station bulletin 616 south dakota state brookingsBrookm gs induced community development coupled with university PROVENZA FDED 1995 postingestivePostingestive feedback as an elemen- patterns of species responses to overstory re- tary determinant of food preferencepieferenceferenee and intake in duction indicates that planning for a higher ruminantslumirumirumlnants journal of range management 48248 2 17 level of landscape diversity would be a logical SEVERSON KEK E AND CEC E BOLDT 1977 options foiroirolfor black hills forest forage both range- approach for ecosystem management in the owners timber or in malsman s journal 4134 13 15 black hills however specific goals under SIEG CHC H AND KEK E SEVERSON 1996 managing habitats current social and economic constraints for white tailed deer in the black hills and bear would likely dictate a landscape that differs lodge mountains south dakota and wyoming a general from one shaped entirely by natural forces review technical report RM 274 USDA forest service rocky mountain forest and range experiment station fort collins CO literature CITED THILENIUS JEJ F 1972 classification of deer habitat in the ponderosa pine forest of the black hills south BENNETT DLD L 1984 grazing potential of major soils with- dakota research paper RM 91 USDA forest ser- in the black hills of south dakota unpublished vicevlee rocky mountain forest and range experiment master s thesis south dakota state university station fort collins CO brookmgsbrookingsBrookm gs TURCHEN LVANDJDLV AND JD MCLAIRD 1975 the black hills BOLDT CEC E RRR R ALEXANDER AND MJM J LARSON 1983 expedition of 1875 dakota wesleyan university interior ponderosa pine in the black hills pages press mitchell SD 126 appp 80 83 in RMR M burns technical compiler silvicul- URESK DWDWANDKEAND KE SEVERSON 1989 understory ovelover ture systems for the major forest types of the united story relationships in pondeiosapondeponderosaiosalosa pine forests black states USDA forest service handbook 445 USDA hills south dakota journal of range management washington DC 4220342 203 208 BOLDT CEC E AND JLJ L VAN DUESEN 1974 silviculture of USDA FOREST SERVICE 1994 the range of natural vanvarivarl ponderosa pine in the black hills the status of our ability for the black hills a fustfirsteustensthirstbirst step appendix A knowledge research paper RM 124 USDA forest pages 1 98 in draft environmental impact state- service rocky mountain forest and range experiment USDA forest service black hills national ment station fort collins CO forest custer SD DUNNETT CWC W 1980 pairwise multiple comparisons in VAN BRUGGEN T 1985 the vascular plants of south the unequal variance case journal of the american dakota and2nd edition iowa state university press statistical association 7537279675372 796 800 ames FERTIG W 1993 black hills national forest sensitive VANGILDER LDL D 0 TORGERSON AND WR porathPORAHI 1982 plant field guide USDA forest serviceSeivice region 2 factors influencing diet selection by white tailed black hills national forest custer SD deer journal of wildlife management 4671146 tii711 718 FFOLLIOT PF AND WP CLARY 1982 understory over WRAGE KJK J 1994 the effects of ponderosa pine pinus story vegetation relationships an annotated bibliog- ponderosa laws on soil moisture precipitation and raphy general technical report INT 136 USDA understory vegetation in the black hills of south forest service intermountain forest and range dakota unpublished master s thesis university of experiment station ogden UT south dakota vermillion GRAVES HENRY S 1899 black hills forest reserve annual report USungeologicalU S geologicalusgeological survey 196719 67 164 received 21 october 1996 GREAT PLAINS FLORA association 1986 flora of the accepted 27 september 1997 great plains university press of kansas lawrence HOFFMAN GRG R AND RRR R ALEXANDER 1987 forest vegetation of the black hills national forest of south great basin naturalist 584 0 1998 appp 328 343

BIRD USE OF RIPARIAN vegetation ALONG THE TRUCKEE RIVER california AND NEVADA

suellen lannilynnilynn1 michael L Mornsonmorrisonl2mornson1212 amy J kuenzilkuenzli jennifer CCC C nealenneale3 benjamin N sacksesacks3 robin hamlm4hamlin4hamline and linnea S hall12

ABSTRACIABSTRACT the truckee river in california and nevada is subject to diverse water regimes and a corresponding variety offlowofdofflowflow rates original riparian vegetation has been altered by these variable flow rates and by a variety of human uses resulting in loss of native riparian vegetation from its historic extent we conducted bird surveys along the truckee rivelriver during spring 1993 to 1 determine relationships between birds and the present vegetation 2 determine the importanceimpotimpoi tance of different vegetation types to sensitive bird species that have declined recently in the western united states due to competition from exotic plant species cowbird molothrus ater parasitism i eductionreduction in nesting habitat or other unidentified reasons and 3 establish a monitoring program and collect baseline data for future comparisons the most frequently detected bird species throughout the study was the brown headed cowbird the greatest number of bird species 98 of 116 was found in the native mixed willow mahxsalixsahx sppapp riparian scrub vegetation type we recommend protecting the remaining native riparian vegetation types for bird habitat along the truckee river

key words bird abundance bird species richness riparian habitat truckee river vegetation type

numbers of neotropical migratory birds are ging gravel removal and grazing klebenow declining throughout north america martin and oakleaf 1984 consequently the truckee and finch 1996 explanations for this decline river riparian corridor is now a thin discon- include reduction and fragmentation of breed- tinuous ribbon of cottoncottonwoodswoods populus sppapp ing wintering and migratory stopover habitat and willows salix sppapp USFWS 1993 ranging stevens et al 1977 finch 1991a riparian cor- up to 250 in wide but averaging approxi- ridors are well known breeding and migratory mately 30 50 in wide where present stopover sites for many neotropical migrants currently there are no baseline data relat- bottorff 1974 stevens et al 1977 wauer 1977 ing bird populations to vegetative communi- szaro and jakle 1985 these corridors are ties along the truckee river our study was important as cover and foraging habitat for designed in cooperation with the US fish and birds migrating through sparsely vegetated wildlife service USFWS to establish a sys- desert areas sprunt 1975 stevens et al 1977 tematic sampling scheme for monitoring bird historically such corridors existed along the numbers and species composition along the truckee river and its tributatributariesries in northeast truckee river and to obtain quantitative base- california and northwest nevada ridgway line data on bird vegetation relationships to 1877 klebenow and oakleaf 1984 satisfy the USFWS operating plan for the at present the native riparian vegetation truckee river our specific objectives were to along the truckee river is greatly reduced from determine 1 bird species composition and its historical extent klebenow and oakleaf relative abundances of birds in the major veg- 1984 USFWS 1993 A number of factors have etation types 2 bird species most likely to be contributed and continue to contribute to the impacted by alterations of the native riparian reduction in riparian vegetation since the late plant communities and 3 vegetative compo- 1800s including varied flow rates from diver- nents successional stage and species composi- sions of water for agricultural use channelization that contribute most to bird abundance tion of parts of the river in the early 1960s log and species richness

school of renewable natural resourcesbesom eh wildlife and fisheries sciences program university of aiarizonaizonaiwona tutsontucson AZ 85721 resenticscnt address department of biological sciences california state university Swsacramentoramento CA 95819 3departentdepal tmentament of environmental science policy and management university of california berkeley CA 94720 4usausU S fish and wildlife serviceSe ivice 4600 keitzkeheitzke lane building C 125 benohenoreno NV 89502

328 199811998 BIRD USE OF RIPARIAN vegetation 329

STUDY AREAS sections based on the approximate elevational border where fremont cottonwood populus we conducted our study along the truckee fremonfremontiifremontiatii changes to higher elevation black river california and nevada approximately cottonwood P trichotrichocarpacarpa USFWS 1993 80 km and the little truckee river 16 km vegetation along the lower truckee river is and independence creek 35353.5 km california characterized by a narrow but extensive strip fig 1 we divided the truckee river into of willow salix sppapp intermixed with occa- lower pyramid lake to sparks nevada and sional clumps of variously aged fremont cot upper floristonflorestonFloriston nevada to lake tahoe tontonwoodswoods agricultural development wide

PYRAMID LAKE

RENO LITTLE TRUCKEE RIVER DEWDAM srampederes

independence CREEK

TRUCKEE

DDNNF A WASHOE WE LAKE

TRUCKEE RIVER

KE TAHOE

10 0 10 20 kilometers

fig 1 map of study site truckee river california and nevada and little truckee river and independence creek california 330 GREAT BASIN naturalist volume 58

spread along most sections of the lower truc- to their occurrence ensuring adequate repre- kee is especially prevalent near the conflu- sentationsentation of the patchy scattered willow and ence of the truckee river with pyramid lake cottonwood vegetation types because the ripar- reservoir cattle grazing also is common near ian vegetation is patchy and thin most survey this confluence hillsidesHillsides bordering the ripar- points sampled 1 vegetation type transects ianlan corridor are dominated by upland shrubs were also distributed along little truckee river primarily shadscaleshadscale atriplex conferticonferttfohaconfertifolialfolial and and independence creek although transects black greasewood sarcobatus vermiculatusverrmculatus on little truckee and independence creek exotic whitetop or peppergrass cafcardanaCardariadarla were established to bisect riparian vegetation draba dominates open disturbed sites aerial photos were not available for these vegetation along the upper truckee river streams and so vegetation was not quantified is also characterized by a narrow strip of wil the beginning of each transect was ran- low cottonwood association black cottonwood domly determined we established fixed points replaces fremont cottonwood between 1800 200 m apart along each transect which ran and 2150 m elevation uplands are dominated parallel to and within 10 m of the stream and by big sagebrush artemisia ttltntrltridentata ripar- within or adjacent to the riparian corridor ianlan vegetation and especially black cotton channelization of the river determined that woods becomes less dense with increasing the transects would be linear ensuring that elevation extensive stands of mixed conifer points were 200 m apart although the num- forest riverbanks reach the nverbanks and dominate the ber of points varied among transects due to higher elevations 1800 vegetation at 2750 m differences in the extent of riparian vegetation along vegetation the little truckee and inde- and accessibility of riverbank all transects hadbad penpendencedence creek resembles the upper truckee at least 8 points we surveyed a total of 250 except the zones along the 2 river riparian points as follows lower truckee 136 upper smaller rivers are dominated by willow alderaider rivers truckee 51 little truckee 45 and indepen- alnus tenuifoliatenuifohd and scrub charac- riparian dence creek 18 terized by willow thickets appendix we surveyed along the lower truckee first because the breeding season began earlier there METHODS relative to the other areas ML morrison we conducted a preliminary study during unpublished field notes 1992 while this may the fall of 1992 to locate appropriate study have confounded our results because we in- areas and determine the latter extent of the cluded migrants only 5 species that we de- breeding season of locally breeding birds tected were not common breedersbleedersbreeders in the area gull observers walked various stretches of the ring billed california gull black chinned truckee river and recorded the presence and hummingbird yellow breasted chat and blue grosbeak frequencies of bird species encountered scientific names in table 2 these during april july 1993 we sampled birds species were all detected in appropriate breed- using the variable circular plot method ralph ing habitat and may have been residents but et al 1993 murray and stauffer 1995 we they were rarely detected and had little impact established evenly spaced points along tran- on our conclusions in addition all of these sects reynolds et al 1980 which were dis- species except ring billed gull have bred in tritributedbuted systematically along the truckee river this area historically ridgway 1877 and there- in a manner that roughly corresponds to the fore were potential breedersbleedersbreeders during our study river stretches used by the USFWS for vege- all points were sampled 3 times with each tation typing USFWS 1993 vegetation types transect surveyed forward twice and backward were identified and quantified by measure- once we completed a round of surveys on ments of percent cover on 1992 aerial pho- each transect before beginning a new round totographs appendix table 1 USFWS 1993 surveys in the lower truckee were conducted vegetation maps were verified by field obser- from mid april to june by 2 observers and in vationsvations vegetation types were homogeneous the other areas from june to early july by 2 and well defined different observers with the exception of one within each river stretch we placed tran- 15 point transect counted by the same observer sects in vegetation types roughly proportional all 3 times each transect was surveyed by 2 199811998 BIRD USE OF RIPARIAN vegetation 331

TABLE 1 percent cover of major vegetation types along the lower upper and overall truckee river and number of birds observed expected and the difference from x2xa2 analysis truckee river california and nevada USFWS 1993

percent cover lumbernumber of birdsabirdisais1 vegetation type lower upper overall observed expected difference open water riverine 12 25 18 b ponds 1 I1 I1 forest sierra mixed conifer 0 0 3 2 636 391 230 black cottoncottonwoodwoodd11 0 7 4 219 521 302 fremont cottonwood willowe 8 3 6 2018 782 1236 shrub alder willow 0 12 6 43 782 739 riparian scrub mixed willow 12 7 10 3011 1303 1708 marsh 3 3 3 161 391 230 gravel bars 10 3 7 56 912 856 sage steppe 16 6 10 1033 1303 270 whitetop peppergrass 14 6 10 381 1303 922 agriculture 23 14 18

basedabased on 7558 observations does not include little truckee river 01or independence creek observations bnotabnotnot quantified fortor these vegetation typesareastypes areas cinecIncincludesludes lodgepole pine jeffrey pine and mixed conifer forest deludesdInc ludes seedling pole sapling and matuiemature stages ofblackosblackofblack cottonwood includes seedling pole sapling and mature fremontpiemont cottonwood with and without whitetop fineFIncincludesludes riparian scrub with and without whitetop

different observers over the season to standard- enough to confidently associate them with ize observer bias verner 1985 all observers vegetation types when the vegetation could be were trained by I1 technician and tested against seen when it could not be seen during a each other to minimize inter observer bias count observers sought out and identified the before performing a count each crew member vegetation after the 5 min count was tested on a practice count where at least we analyzed our data to obtain an index of 90 of all detections were identical between abundance mean number of birdspointcount trainer and trainee species identification and raphael 1987 and frequency of occurrence distance estimations were checked across ob- percentage of points at which a species was servers by informal testing throughout the detected verner 1985 for each species dis- sampling season the paces of each observer cussed because we had small sample sizes of were measured by walking 50 in for 3 replica- individual species in each vegetation type and tions at normal speed distance estimations distances to birds are often difficult to estimate were checked by pacing to stationary objects verner 1985 we included all detections throughout the season ralph et al 1993 regardless of distance from observer in our we counted birds at each point for 5 min abundance analysis blondel et al 1981 sliwa all counts were conducted within the first 4 h and sherry 1992 we also analyzed our data after sunrise and only on days without precipi- to obtain distribution of birds by vegetation tation or significant wind before beginning a type highlighting bird species richness within count the observer waited for 1 min to allow vegetation types and distribution of species possibly disturbed birds to resume their nor- among vegetation types mal behaviors murray and stauffer 1995 all to test the validity of comparing bird de- birds seen or heard at each point were re- tections among vegetation types we examined corded we also recorded the vegetation type the relationship among major vegetation types in which each bird was located appendix and the distribution of detections of birds by detection mode visual song call and dis- distance from the point center using chi square tance from the point to the bird before begin- analysis this analysis tested whether detect- ning any survey each observer was shown ability measured by average detection distance examples of all vegetation types which were of bird species varied among vegetation types distinct and easily identifiable therefore we had they differed comparisons of bird abun- could locate the birds precisely and accurately dance between vegetation types would have 332 GREAT BASIN naturalist volume 58

taboTABUTABLE 2 index of abundancea xY and frequency of occurrenceoccurrenceboccurrence13occurrences13 of birds along the lower truckee upper truckee and little truckee rivers and independence creek nevada and california spring 1993 species of special interest are identified with superscriptsuperscriptssc independence lower TruckTruckeeeed1 upper Trucktruckeeeee little trueTructruckeekeef creeks species scientific nameh Yx s xY s xY s xY s american white pelican 83 57 3 pelecanus erythrorhynchos double crested cormorant 04 01 5 phalacrocorax auritusauntusauretus great blue heron 04 01 2 03 00 1 ardea herodias snowy egret 05 03 2 egretta thula black crowned night heron 03 00 1 nycticorax nycticorax white faced ibis 17 00 1 plegadis chihichihtchahi canada goose 25 33 15 17 00 1 03 00 2 branta canadensis wood duck 06 03 6 aix sconsasponsa mallard 09 05 22 14 20 3 03 00 1 anas platyrhynchosplatyrhynchous northern pintail 03 00 1 anas abutaacuta cinnamon teal 07 03 3 anas cyanoptera gadwall 07 03 2 anas strepera unknown duck 09 08 5 common merganser 08 04 6 04 02 8 12 13 10 03 00 2 mergus merganser turkey vulture 42 76 2 11 07 2 cathartes aura osprey 03 00 1 03 00 1 03 00 2 pandion haliahailahalmehaimehaliaetushalwehalwetitshaliaeetustitsetus red tailed hawk 04 02 7 04 02 3 buteojamaicensisbuteo jamawensis golden eagle 03 00 1 aguila chrysaetos american kestrel 05 02 10 falco sparspansparveriussparvenusspderiusveriusardenusarvenus california quail 06 03 16 14 12 6 callipeplacalhpeplaCalliCalhpepiapepla calicallcailcalifornicacaliformcdcaliforfornicamcd american coot 07 06 3 fulica americanaamencanaemenamencenacana killdeer 06 03 15 03 00 1 charadriusCharadnus vocivociferusvociferousferus american avocet 18 12 1 recurvirostra americanaamencana spotted sandpiper 04 02 9 08 06 20 06 03 21 03 00 2 actipisactitisActitis maculansmamacumaculandmaculariaculandlatiaiarialaria long billed curlew 03 00 1 numenius americamericanusamericanosamencanusanus common snipe 03 00 1 gallinagoGalhnago gallinagogalhn090agoogo ring billed gull 21 26 1 larus delawarensis 199819981 BIRD USE OF RIPARIAN vegetation 333

TABLE 2 continued independence lower Truckeetruckee1Truck eed1 upper TruckTruckeeeee6 little truetruckeeTruckeef creeksgreeks species scientific nameinamehnamebname1 7.7 s x s x s x 8 california gulichgullnhgulinh 07 05 1 larus californicuscalifornicus unknown gull 41 68 12 rock dove 12 13 4 04 02 3 columba livia band tailed pigeon 03 00 1 columba fafasciatafascianasciata mourning dove 07 04 19 07 00 1 03 00 1 zenaida macrouramacroura great horned owl 05 02 1 bubo virginianusvirgimanusvirgivirginianuslanusmanus black chinned hummingbird 07 00 1 archilochusArchilochus alexandri calliope hummingbird 06 04 2 05 02 4 Stellula calliope belted kingfisher 04 02 3 04 02 2 03 00 2 ceryle alcyon red breasted sapsucker 03 00 1 05 03 4 04 02 8 sphyrapicus ruber wilhamsonswilliamsorisWilhamWilliam sorissons sapsucker 05 02 2 03 00 2 sphyrapicus thyroidthyroideuseus downy woodpecker 05 02 3 04 02 3 03 00 1 ficoidesPicpicoidesoides pubescenspubescentpubescens hairy WoodpeckerwoodpeckeinhwoodpeckernhNH 03 00 1 03 00 1 06 05 8 ficoidesPicpicoidesoides villosisvillosusvillosus white headed woodpecker 05 02 1 03 00 1 ficoidesPicpicoidesoides albolarvatus black backed woodpecker 03 00 2 ficoidesPicpicoidesoides arcticusarcticus northern flicker 05 03 3 04 01 9 05 03 12 04 02 4 colantescolaptes auauratusauranusratus olive sided flycatcher 11 03 00 1 04 01 7 03 00 1 03 04 4 Concontopustopus borealis western wood pewee 05 03 8 12 07 23 13 05 31 15 10 36 Concontopustopus sordidulussordidulus willow Flycatcherflycatcher03flycatchercb03CB 03 00 1 empidonax traitrailiilii hammond s flycatcher 03 01 1 empidonax hammondii dusky flycatcher 03 01 2 05 02 1 08 04 32 empidonax oberholseri empidonax species 04 02 3 03 05 3 05 03 10 06 02 28 say s phoebe 03 01 1 sayornisSaysayormsornisorms sayasay a western kingbird 07 05 18 tyrannus vertiverticalisverticalismcalis treeswallownhcbtreeswallownhcb 04 02 1 11 10 6 03 00 2 tachycineta bicolor violet green swallownhcbswallow CB 03 00 1 03 00 1 tachycineta thalasthalassinosina northern rough winged swallow 09 09 22 43 26 5 stelgidopteryx serriserripennissernpenmspennis bank SwallowswallownhNH 08 05 5 ripariariparialRiparia ripariaparlaropana 334 GREAT BASIN naturalist volume 58

TABLETABLL 2 continued independence lower truckeeeed upper TruckTruckeetruckeeetruckee6eee6 little truetruckeeTruckeef creeksgreeks species Truck scientific nameli 1 s xY 8 xY 8 x s cliff swallow 18 33 7 41 56 6 62 79 5 hirundo pvrrhonotapyrrhonotapyrrhon ota bainbarn swallow 09 08 14 17 17 2 hirundo rustica unknown swallow 07 07 4 stellersteilerstelier ssjayjay 14 08 33 08 06 22 04 02 6 cyanocitta stelleristelspellerileri chukclark s nutcracker 07 00 2 Nucinucifragafraga columbiana

blickblack billed magpie 11 09 25 07 00 1 pica pica americanamerleanamencanciowcrow 03 00 1 corvus brachirhunchosbrachyrhynchos common raven 03 01 2 corvus boraxcorax mountain chickadee 12 07 21 09 07 21 12 07 36 parus gambeligamgambelinbeli bushtit 05 03 1 psaltnparuspsaltriparus minimus

red breasted nuthatch 06 03 3 03 04 2 03 00 2 sitta canadensis white breasted nuthatch 05 02 1 03 00 1 03 00 2 sitta carolinensis pygmy nuthatch 00 04 1 sitta pygmaeapygmaean blownbrownbiown creeper 03 04 1 04 01 5 07 03 12 certhia americanaamencanaamencana rockzienrockwienrockroekRoc kWienwren 04 01 5 salpinctes obsoletusobsoletes

bewick s wrenwwi en 08no 07n 7 14ii nlthruomanesthryomanes bewickbewicknbewickebewickiiii house wienwren 11 08 24 17 09 10 16 11 4 07 04 10 troglodytes aedonabdon maishwiennllmarsh wrcnc13n11 03 00 1 cistothorusCistoci&tothoruscistophorusthorus papalustrislustris americanamerlean dipper 03 01 3 05 02 4 cinclus mexicanusmexicanus rubyrubyel ci ownedcrowned kinglet 03 00 1 regulus calendula

blue dayoaycraygray gnatcatcher 03 00 1 polioptila caecaerulearulea mountain bluebird 06 04 2 sialia currucoides townsend s solitaireSolitane 03 00 1 06 03 5 myadestesMyadestes towntownsenditownsendiasendi hermit thrush 03 00 2 tharuscatharscathamscatharusCathams gutguttatusgustatustatus americanamerlean robin 07 05 18 10 07 29 13 07 30 13 09 34 hurcturdusTurclusius miqramigramigratonusinigratoriustuhustunustonus

cedargedalgedar waxwing 17 00 1 bombycillabomhycillaboinbycilla cedrorumcedrorum eulEuieuropeanopean starllngexstarlingexStarlin gEX 12 11 22 04 02 2 sturnus vulgaris solitary vireovirco 04 02 1 03 00 6 vireo solitariussolitanussoltsolisoittariustanus 199819981 BIRD USE OF RIPARIAN vegetation 335

TABLE 2 continued indindependencespende lowilowersr truckeedTrucktrue eed upperupp er truckeeeTruck keeakee6eee ie Truc keeeeekeef i species truetruc littlittlelittie truetruckeef creeks

scientific nameliname x S X s x s X s warbling vireocbvireo 04 03 6 13 08 23 10 06 23 19 10 36 vireo filous changeorange crowned warbler 03 00 1 wehnwennvennivoraverrmvoraVennluoraldoraivora celata nashville warbler 08 05 15 07 00 1 03 04 4 vermivoraVermivora ruftcapillaruficapilld yellow Warblerwarblercbwarbler013gilCB013 05 04 10 09 07 21 17 12 21 13 09 26 dendroica letechiapepetechiatechia yellow rumpedbumped warbler 07 04 10 07 04 11 07 04 14 08 06 22 dendroica coronata

macgillivray s warbler 04 01 2 06 02 6 04 02 3 14 08 6 oporornis tolmietolmieii common Yellowthroatyellowthroatcbyellowthroat015cilcB015 04 01 2 03 00 1 geothlypis tritrltnchastrichasdrichaschas wilson s warbler 05 02 15 08 06 11 07 07 9 07 06 20 Wilwilsoniasoma ausillapupusillasilla yellow breasted chatcbnhchatgb NH 05 03 1 ictericIcictenaicteriateriaterla wrens unknown warbler 04 01 3 western Tanagertanagercbtanager015cilCB015 04 03 3 04 01 4 piranga ludovicianaludoviciana black headed grosbeak 05 02 6 04 01 7 08 08 2 pheucticus melanocephalus blue grosbeak 03 00 1 Guguiracairaca caecaerulearulea lazuli bunting 05 02 7 passerinaPasnassenna amoena green tailed towhee 07 05 14 07 05 8 pipilo chloruruschlorurus spotted towheeCB 07 03 1 05 02 5 pipilo1 macumaculatismaculatuslatus chipping sparrowCB 03 01 2 03 05 3 08 05 6 04 02 8 spizella paspasserinasenna brewer s sparrow 07 04 4 spizella brebrewenmetiwerimerl vesper sarrow 06 03 4 pooecetes gramineusgraminousgramineus black throated sparrow 03 00 1 amphispiza bilineatabilmeatabilineatebtlbilineatameata savannah Sparrowsparrowcbsparrow0808CB 05 02 1 basserrasserfasserfasserculusPasserpasserculusculus sandwichensis fox sparrow 05 02 8 03 00 2 passerella ilincailiacaifiaca song SparrowCB 05 03 9 21 10 33 19 16 21 11 08 22 melospiza melmeimelodiaodid lineolLincollincolnrss sparrow 06 04 2 06 03 3 melospiza fincolniilincolmilincolmi white crowned sparrow 03 00 1 09 07 6 03 00 6 zonotrichia leucophrysleucophrys dark eyed junco 11 06 17 11 08 19 11 07 28 junco hyemalishyemalis red winged blackbird 12 09 23 17 15 5 21 14 5 agelaius phoeniceusphoemceusphoeniceus western meadowlar8MeadowlarkmeadowlarkcbCB 05 03 13 sturnella negneglectsneglectalecta 336 GREAT BASIN naturalist volume 58

TABLE 2 continued independence lower truckeeTruckeed upper TruckTruckeetruckeeetruckee13eee13 little trueTructruckeekeef creeks species scientific nanamelirnell Y s x 8 s s yellow headed blackbird liii111.1 16 6 xanthocephalus xanthocephalus brewer s blackbird iili111.1 090.9og 14 141.4 131.3 13 101.0loio 080.8os 10 eupharuseuphagusEuphagus cyanocephalus brown headed cowbirdEX 15 090.9og 32 090.9og 050.5os 28 060.6og 040.4 16 050.5os 020.2og 22 molothrus ater unknown blackbird 07 06 12 northern oriole 101.0loio 050.5os 28 030.3 000.0oo 1 icterus galbula purple finch 070.7 07 3 carpodacus purpureuspurpurpurpureouseus cassin s finch 070.7 000.0oo 1 090.9og 05 4 11iili1.1 070.7 12 carpodacus cassiniicassicassiniacas&iniiniinil house finch 050.5 050.5os 5 carpodacus inexicanusmexicanusmexicanus pine siskin 030.3 000.0oo 1 090.9og 020.2 2 080.8 040.4 6 carduelis pinus lesser goldfinch 03 000.0oo 1 carduelis pspsaltriaaltria house SparrosparrowwEX 080 8 04 5 passer domesticusdomesticus number of blldspointcountofbirdspointcount inumber1ntiinbeiincumber ofot detections of this speciesspeciestotaltotal counts on this tiantransecttran sectseetveet EX exotic or01 opportunisticoppoioppor tunistictunitungsticsilcsticsile species possibly having a negative impact on native species NH species with sensitive nesting habitat impacted by alterations inin riparian vegetation LBGB species adverselyadveisely affcctedaffected by exotic 01or opportunisticoppol tunistictunitungsticstic species species declining for unknown leasonreason ellig11119llo110IIO points 357 total counts points x 3 u6818 points 204 total counts points X 3 HS145 points 135 total counts points x 3 918 points 54 total counts points x 3 laouIAOUAOU 19831995 not presentpiescntent been invalid in a vegetation type where bird to be most consistently nearer the point cen- calls do not carry well eg dense trees the ter thus explaining some of the bias average distance of detection will be smaller we also conducted chi square analyses to than in vegetation types where bird calls carry determine if there is a difference between long distances eg open grassland for 8 of general bird use of vegetation types and avail- 10 species examined average detection distance ability of these vegetation types to the birds did not vary between vegetation types FP because our survey focused on nonagricul- 010.1aloi the remaining 2 species had 0050.05 FP tural vegetative relationships for this analysis 010.1oloi for these 2 species average detection dis- we excluded individual birds detected in agri- tance was shorter in riparian scrub than in other cultural areas and areas that were not quanti- vegetation types some of our data therefore fied on vegetation maps 36 of all detections are slightly but not usually significantly biased throughout this paper we discuss general toward fewer detections in the riparian scrub trends for all bird species focusing on 21 avian vegetation type however the number of detec- species of special interest tables 2 4 these tions in this vegetation type for these species species include those thought to be decreasing exceeds detections in any other vegetation in abundance due to competition from oppor- type also our survey points often sampled tunistictunistic and exotic species or thought to be multiple vegetation types therefore the dis- impacted by alterations of riparian vegetation tance from the point center to a given vegeta- other species of interest may be increasing tion type varied riparian vegetation tended opportunistic and exotic species eg brown 199819981 BIRD USE OF RIPARIAN vegetation 337 headed cowbird european starling house along the little truckee 4 were detected at sparrow that may adversely impact riparian 10 of the counts and 3 at6ataat & table 2 birds identification of these species of special independence CREEK the most fre- interest was designated by the USFWS based quently detected bird species on indepen- on review of previous works ridgway 1877 dence creek was the mountain chickadee klebenow and oakleaf 1984 and communica- followed by warbling vireo western wood tion with regional biologists USFWS califor- pewee american robin dusky flycatcher nia parks department dark eyed junco unidentified empidonax flycatchers yellow warbler brown headed cow- RESULTS bird song sparrow yellow rumpedbumped warbler and Wilwilsonsorssoiss warbler seven species of special species richness and interest were detected along independence abundance of birds creek 4 were detected at 20 of the counts OVERALL we detected 116 species across and 3 at8ataat table 2 the entire study area the most abundant over all sections of the river 2 individuals species overall was cliff swallow followed by of each of the most frequently detected american white pelican song sparrow turkey species were observed during any single point vulture northern rough winged swallow and count table 2 house wren table 2 mean s of bird species distribution of birds richness per point was 16 4 among transects by vegetation type on the lower truckee 14 4 on the upper truckee 13 2 on the little truckee and 13 RICHNESS AND PERCENT occurrence OF 4 on the single independence creek tran- BIRDS AMONG vegetation TYPES we did not sample sect total bird abundance on each transect each vegetation type equally throughout the truckee river drainage therefore the ranged between 878.7 and 14814.8 birdspointcount following 3 results sections should be consid- with a mean of 11211.2 181.8lsis birdspointcount ered as baseline data to be compared with LOWER TRUCKEE RIVER most fre- the future sampling quently detected bird species along the lower avian highest bird species richness occurred truckee was the brown headed cowbird fol- the in the riparian scrub vegetation type with 17 lowed by northern oriole black billed magpie species detected only in riparian scrub sage- house wren winged blackbird european red brush steppe riparian scrub with whitetop starling northern rough winged swallow and mature fremont cottonwood with and without mallard table 2 eighteen species of special whitetop pole sapling fremont cottonwood detected along lower interest were the truc- with and without whitetop whitetop alone kee 4 at 10 of the counts and the remain- and sierra mixed conifer also had high species ing 14 atata 9 of the counts table 2 at9 richness 40 table 3 of 116 bird species UPPER TRUCKEE RIVER the most fre- observed during our study only the pine quently detected species along the upper siskin white breasted nuthatch white faced song sparrow truckee was the followed by ibis blue gray gnatcatcher brown creeper steller s jay american robin brown headed black throated sparrow cedar waxwing and cowbird warbling vireo western wood hairy woodpecker were never in pewee detected mountain chickadee yellow warbler native riparian vegetation and spotted sandpiper twelve species of spe- species richness was 30 less in riparian cial interest were detected along the upper scrub that contained whitetop however bird truckee 4 at 20 of the counts and the richness in fremont cottonwood was the same remaining 8 at 6 of the counts table 2 with and without whitetop table 3 LITTLE TRUCKEE RIVER along the little riparian scrub vegetation had the highest truckee the most frequently observed bird percentage of detections of all species over all species was the western wood pewee fol- points in our study sagebrush steppe was the lowed by american robin warbling vireo only other vegetation type with 10 of all steller s jay mountain chickadee song spar- birds detected no single successional stage of row spotted sandpiper and yellow warbler cottonwood had 8 of all detections how- seven species of special interest were detected ever 21221.2 of all birds observed were across 338 GREAT BASIN naturalist volume 58

TABLE 3 species richness number and percentage of all birds detected by vegetation type truckee river and vicinity california and nevada spring 1993 a vegetation tipelltypelltype11 number pole sapling fremont cottonwood willow 49 33 matuiematulemature fremontpiemont cottonwood willow 53 66 seedling fremont cottonwood willow with whitetop 9 01 folefoiepole sapling fremont cottonwood willow with whitetop 48 37 matuiematulemature fremont cottonwood willow with whitetop 57 75 riparian scrub 93 287 riparianripal lanian scrub with whitetop 62 96 whitetop 49 43 sage steppe 77 140 maishmarshmalsh 27 16 gravel baibalbar 17 05 seedling black cottonwood 5 01 pole sapling black cottonwood 18 08 mature black cottonwood 28 15 jeffreyjeffi ey pine 39 41 lodgepole pine 40 58 sienaslenasierra mixed conifer 42 57 agriculture 34 21 till1113ased1113liased on 1181211 812 observations iseeselSLL appendix forfoifbiboioor tulitullfuli description ofot vegetation types all stages of fremont cottonwood pure white- quently detected in fremont cottonwood and top stands supplied 434.3 of all detections riparian scrub at low elevations both starlings table 3 and house sparrows were detected primarily the percentage of individual birds detected near buildings and agricultural fields was low in all conifer vegetation types 6 SPECIES POSSIBLY AFFECTED BY EXOTIC OR overall the 3 conifer vegetation types jef- opportunistic SPECIES UNKNOWN REASONS OR frey pine pinusjeffreyipinus jeffreyi lodgepole pine pinus alteration IN RIPARIAN vegetation califor- confortacontortacontorta var murtamurramurmurrayanamurrayandtayanarayanayand and mixed conifer nia gulls common Yellowyellowthroatsthroats spotted contained 15615.6 of all birds detected black towheesTowhees tree swallows willow flycatchers cottonwood which occurs within the conifer marsh wrens chipping sparrows savannah zone contained only 242.4 of bird occurrences sparrows and yellow breasted chats were table 3 rare but most often detected in riparian scrub BIRD SPECIES detections ACROSS vegeta- table 4 tree swallows were also observed tion TYPES thirteen species were detected nesting in mature black cottonwood olive in 10 vegetation types whereas 40 species sided flycatchers thought to be declining were detected in 3 different types brown throughout the west robbinsbobbins et al 1986 headed cowbirds table 4 and american desante and george 1994 were most often robins were detected in all vegetation types observed in sierra mixed conifer and riparian and both were most commonly detected in scrub western Meadowmeadowlarkslarks were fairly fre- riparian scrub vegetation quent across most vegetation types and vio FREQUENCY OF BIRD SPECIES OF SPECIAL let green swallows were only rarely detected INTEREST AMONG vegetation TYPES we con- both species were detected only at lower ele- sidered a bird species to be rare if it was vationsvations song sparrows were common and detected with a frequency of25of 25252.5 during western tanagersTamanagersnagers were rarely detected both 20 of the 750 total point counts were seen across most vegetation types war- EXOTIC OR opportunistic SPECIES brown bling eireosvireos were frequent in riparian scrub headed cowbirds were common and were lodgepole pine and black cottonwood yellow detected in all vegetation types though less warblersWarblers were detected across all riparian frequently at higher elevations table 4 types most frequently in riparian scrub european starlings were frequently detected SPECIES POSSIBLY IMPACTED BY alteration at lower elevations where there were trees IN RIPARIAN vegetation bank swallows were and also in sagebrush steppe similarly the most frequently detected in fremont cotton- introduced house sparrow was most fre wood and sagebrush steppe table 4 hairy 199819981 BIRD USE OF RIPARIAN vegetation 339

TABLE 4 percent detections of species of special interest among vegetation types3tyresatypesa along the truckee river and vicinity california and nevada species cwew cwwbcwwcaw rs rswraw w SS m 9 bebc JP IPlp fmcsmc a ExoticexoticopportunisticexoticopportnnisticOpportunistic european starling 19 31 22 13 4 12 I1 1 0 0 0 0 0 brown headed cowbird 13 12 26 13 8 17 2 1 2 1 1 4 I1 house sparrow 39 10 49 3 0 0 0 0 0 0 0 0 0 competition with exoticsexotica or unidentified reasons olive sided flycatcher 0 0 31 8 0 0 0 0 0 8 8 46 0 willow flycatcher 0 0 100 0 0 0 0 0 0 0 0 0 0 tree SwallowSwallowe0 0 0 42 2 2 0 0 4 46 2 2 0 0 violet green SwalloSwallowwc0 0 33 0 0 0 67 0 0 0 0 0 0 0 marsh wrene 0 0 0 67 0 33 0 0 0 0 0 0 0 warbling vireo 1 14 32 3 1 1 0 0 17 2 18 12 0 yellow warbler 8 17 43 10 2 4 1 0 9 0 3 4 1 common yellowthroat 9 18 0 36 9 18 0 0 0 0 0 9 0 yellow breasted chatcchat0chanc 0 25 13 50 13 0 0 0 0 0 0 0 0 western tanager 15 20 10 15 0 0 5 5 5 5 0 20 0 spotted towhee 0 0 47 24 0 6 0 0 12 6 0 6 0 chipping sparrow 0 0 40 0 20 20 0 0 0 0 0 20 0 savannah sparrow 0 0 67 0 0 33 0 0 0 0 0 0 0 song sparrow 5 2 64 1 2 7 1 0 5 2 2 12 0 western meadowlark 28 26 13 19 2 12 0 0 0 0 0 0 0 impacted by loss of habitat or vegetation alteration california gull 0 0 100 0 0 0 0 0 0 0 0 0 0 hairy woodpecker 0 0 0 0 0 0 0 0 0 0 0 100 0 bank swallow 48 0 24 8 8 32 0 0 0 0 0 0 0

vegetation types and codes found inin appendix bww vegetation type mixed vithwith whitetop alsoaisocalsocaiso affected by loss ofhabitat or alteration of vegetation

woodpeckers were detected only in sierra section s extensive riparian scrub and fremont mixed conifer both species were rare cottonwood stands these vegetation types USE VERSUS availability OF vegetation decreased in area with increasing elevation TYPES overall birds did not use vegetation USFWS 1993 on the upper stretches of the types in proportion to their availability xax22 river higher elevation black cottonwood com- 7254 df 8 FP 00010.001 the discrepancy be- munimunitiesties did not replace lower riparian scrub tween use and availability was highest in ripar- cottonwood communities in terms of bird ian scrub and fremont cottonwood although species richness elevational temperature gra- totaling only 10 cover table 1 riparian dients and arthropod abundances were not ex- scrubmixedscrub mixed willow was used by birds almost amined in this study but may have contributed 40 of the time during our observations bird to levels of species abundance we observed use of monotypic whitetop was significantly black and fremont cottonwood each occupied less than expected given its percent cover the similar absolute areas table 1 hence differ- number ofbirdoxbirdof bird species using these vegetation ences in bird richness were unlikely due to an types supports our findings ofbirdoxbirdof bird preference area effect fremont cottonwood 70 species riparian scrub transects at higher elevations were com- mixed willow 80 species and whitetop 44 posed of coniferous vegetation with a narrow species strip streamsideofstreamsideof riparian vegetation a conifer overstory was often present at streamside in discussion contrast transects at lower elevations were predominantly overall distribution and riparian with a cottonwood overstory and scrub understory abundance of birds thus by virtue of abundance of vegetation types alone lower the lower truckee river harbored the great- elevation areas should be dominated by riparcipar est richness of avifauna of any stream section ian associated bird species while upper eleva we monitored this was due primarily to the tion areas should have fewer riparian associated 340 GREAT BASIN naturalist volume 58

bird species knopf 1985 and finch 1991b and with a much greater frequency than their also reported different bird communities asso- availability therefore a drastic reduction in ciated with different elevations native riparian forest abundance may have more effect species specific considerations on birds than a reduction in any other plant species along the truckee river habitat the brown headed cowbird was widely specialists 40 species found in 533 different distributed but reached its highest numbers at habitat types were observed most frequently lower elevations where agriculture was promi- in riparian scrub even the habitat generalists nent table 1 these birds typically forage in in our study brown headed cowbird and agricultural areas while sometimes flying long american robin the bird species found in distances to find forested nesting habitat the the greatest number of different vegetation brown headedbeaded cowbird was the most fre- types were most frequently observed in ripar- quently encountered bird along the lower ian scrub therefore both dominant riparian truckee and was also found in the greatest vegetation plants cottonwood and willow the yellow number of vegetation types warbler dominant plant species in riparian scrub yellowthroat yel warbling vireo common should be considered in developing manage- song low breasted chat and sparrow were ment plans for protecting the habitat of truc- also detected during study our and are known kee river bird species to be adversely impacted by cowbird nest par- because 98 of 116 bird species were de- asitism friedmann et al 1977 tected in native riparian vegetation 22 exclu- european starlings along common the sively the majority of bird species would be lower truckee were especially numerous near impacted in some way by altering native ripar- agricultural fields and buildings their num- ian plant communities although sagebrush bers decreased rapidly with increasing eleva- steppe also had a high species richness this is tion therefore their potential impact on eav probably due to its proximity to riparian vege- ity nesting species may be of primary concern tation creating an ecotone that attracts only at lower elevations stoner 1939 jackson more sparrows species than the sagebrush steppe vegetation and tate 1974 house were rare gates and were detected primarily around buildings type alone and gysel 1978 willow flycatchers which are declining in impacts of exotic vegetation the west desante and george 1994 rothstein and robinson 1994 were detected at the major exotic plant of interest in our study was whitetop only 1 point on the upper truckee river and or peppergrass whitetop nowhere else dates of the sightings 18 and was used by bird species for foraging S lynn 24 june suggest the probability of breeding personal observation but nesting in this plant activity bent 1942 mccabe and hovel 1991 species was not documented whitetop was but we were unable to confirm this negatively associated with bird species rich- avifauna of little truckee river and inde- ness in riparian scrub however bird richness penpendencedence creek were dominated by species in fremont cottonwood did not differ with the typical of coniferous forests however war- presence of whitetop further research in this bling eireosvireos song sparrows and yellow war area is necessary to determine whether there biers all riparian associated species of concern is a cause effect relationship between white- were also common in both areas throughout top and bird species richness the study in addition the wilson s warbler in summary a varied flow regime overgraz- another riparian associated species was com- ing channelization and other human activities monly observed on independence creek and have altered riparian vegetation along the to a lesser extent on the upper truckee and truckee river and its tributariestributaries klebenow little truckee even in these conifer domi and oakleaf 1984 destruction and removal of natedbated areas small patches of riparian vegeta- native cottoncottonwoodswoods and willows from the ripar- tion apparently are enough to support these ian corridor has likely resulted in a decrease riparian associated bird species in numbers of riparian obligate bird species a historical issue which will be presented in a species richness in vegetation future manuscript also avian exoticsexotica and op- fremont cottonwood and riparian scrub portunisticportuni stic species such as the brown headed willow were used by a wide variety of birds cowbird could potentially reduce sensitive 199819981 BIRD USE OF RIPARIAN vegetation 341 species richness and abundance the effects of BLONDEL J C FERRY AND B FROCHOT 1981 point counts cowbird parasitism and exotic species on sen- with unlimited distance pages 414 420 in CJC ralph and M editors sitive bird species in the truckee area JMJ scott estimating numbers in river of terrestrial birds studies in avian biology 6 warrant in further investigation also warranting BOTTORFF RLR L 1974 cottonwood habitat for birds in further investigation is the effect that the Coloiacoloradodo american birds 2897528 975 979 exotic plant species whitetop may have on bird DESANTE DFD F AND TL GEORGE 1994 population trends distribution unfortunately whitetop is an in the landlandbirdsbirds of western north america studies in avian biology 1517315 173 190 especially hardy is difficult to species that FINCH DMD M 1991a population ecology habitat require- eradicate rosenfels and headley 1944 man- ments and conservation of neotropical migratory agement plans may have to consider it as a birds general technical report RM 205 US permanent aspect of the riparian community department of agriculture forest service rocky range and concentrate on keeping existing patches mountain forest and experiment station fort collins CO 26 appp plant of the from spreading into native ripar- 1991b positive associations among riparian bird ian habitats species correspond to elevational changes in plant managers who are interested in halting communities canadian journal of zoology 69 951 963 declines in bird populations and stimulatinstimulationstimulating9 FRIEDMANN H LEL F KIFF AND SIS I1 ROTHSTEIN 1977 A growth in these populations should consider further contribution to knowledge of the host rela- protecting existing native fremont cottonwoodscottonwoods tions of the parasitic cowbirds smithsonianSmith soman contri- and riparian scrub vegetation as well as per- butions to zoology 235 smithsonianSmithsoman institution haps initiating restoration of these vegetation press washington DC 75 appp types inm degraded areas that we collected GATES JEJ E AND LWL W GYSEL 1978 avian nest dispersion data and fledging success in field forest ecotoneseco tones ecol- will be valuable as a baseline from which to ogy 5987159 871 83 compare future bird surveys along these rivers JACKSON JAJ A AND J TATE JR 1974 an analysis of nest as land uses change or continue in degradation box use by purple martins house sparrows and of native riparian forest researchers starlings in eastern north america wilson bulletin future 8643586 435 449 will be able to use our data to discover and KLEBENOW DAD A AND RJR J OAKLEAEOAKLEAF 1984 historical avi-avi confirm trends among bird species and their faunal changes in the riparian zone of the truckee vegetation requirements along the truckee river nevada pages 203 210 inm RER E warner and river and vicinity KNK N hendrix editors califoiniacalifornia riparian systems university of california pipressess berkeley KNOPF FLEL 1985 significance of riparian vegetation to acknowledgments breeding birds across an altitudinal dinehinecline pages 105 111 in RRR R johnson CDC D ziebell DRD R patten we thank the US fish and wildlife service PFPE ffolliotffolliott and RHR H hamre technical coordinators especially david L harlow of the reno field riparian ecosystems and their management recon- ciling conflicting general office for consulting on logistical arrangements uses technical report RM 120 US department of agriculture forest serviceseiset vicevlee and financially supporting this project we MARTIN TE AND DMD M FINCH 1995 ecology and man- also thank sagehen creek field station uni- agement of neotropical migratory birds a synthesis versity of california for providing housing and review of critical issues oxford university pipressess the landowners along the truckee little truc- new york 489 appp mccabe BAR A AND S HOVEL 1991 the little green bird kee and independence creek for allowing ecology of the willow flycatcher rusty rock press access to their property and several anony- madison wtWI mous reviewers as well as and especially MURRAY NLN L AND DED F STAUFFER 1995 nongame bird steve knick for critiques of this manuscript use of habitat in central appalachian riparian forest journal of wildlife management 597859 78 88 RALPH CJC J GRG R GEUPEL P PYLE TE MARTIN AND DED F literature CITED DESANTE 1993 handbook of field methods for moni-moni toring landbirdslandbirds general technical report PSW AMERICAN ornithologists UNION AOU 1983 check- GTR 144 USU S department of agriculture forest list of north american birds ath6th edition alienallenailen press service pacific southwest research station 41 appp inc lawrence KS 877 appp RAPHAEL MGM G 1987 estimating relative abundance of 1995 fortieth supplement to the american orni forest birds simple versus adjusted counts wilson thologists union checklistcheck list of north american birds bulletin 9912599 125 131 auk 112819112 819 830 REYNOLDS RT JM SCOTT AND RAR A NUSSBAUMNUSSRAUM 1980 A BENT ACA C 1942 life histories of north american flycatch- variable circular plot method for estimating bird ers larks swallows and their allies USU S national numbers condor 8230982 309 313 museum bulletin 179 smithsomansmithsonianSmithsoman institution USU S RIDGWAY R 1877 ornithology pages 303 669 in C king government puntingprinting office washington DC 555 ornithology and paleontology USU S geological explo- PP rations 40th parallel 4 342 GREAT BASIN naturalist volume 58

ROBBINSBOBBINS CSC S D BYSTRAK AND PH GEISSLER 1986 the STONER D 1939 parasitism of the english sparrow on breeding bird survey its first fifteen years 1965 1979 northern cliff swallow wilson bulletin 5122151 221 222 united states fish and wildlife service washington STRONG TR AND CEC E BOCK 1990 bird species distribu- DC tion patterns in riparian habitats in southeastern ari- ROSENFELSrosenlROSENI ELS RSR S AND FBEB HEADLEY 1944 whitetop eradi- zona condor 9286692 866 885 cation bulletin 170 university of nevada agricul-agncul SZARO RCR C AND MDM D JAKLE 1985 avian use of a desert tural department 18 appp riparian island and its adjacent scrub habitat con- ROTHSTEIN SIS I1 AND SKS K ROBINSON 1994 conservation dor 8751187 siisll511 519 and coevolutionary implications of brood parasitism USU S FISH AND WILDLIFE SERVICE USFWS 1993 truc- by cowbirds trends in ecology and evolution 9 kee river riparian vegetation and fluvial geomor- 162 164 phology study appendix C truckee river riparian SLIWA A AND TW SHERRY 1992 surveying wintering corridor cover and land use types USU S department warbler populations in jamaica point counts with of the interior USU S fish and wildlife service sacra-S aeraacra and without broadcast vocalization condor 94 mento CA 924 936 VERNER J 1985 an assessment of counting techniques SPRUNT A IV 1975 habitat management implications of current ornithology 22472 247 302 migrationmigiation pages 81 8686otin proceedings of the sympo- WAUER RHR H 1977 significance of rio grande riparian siumslum on management afforestofforestof forestmorest and range habitats for systems upon the avifauna pages 165 174 in RRR R nongame birds general technical report woiWO 1 johnson and DA jones technical coordinators USU S department of agricultureofagriculture forest service tuc- importance preservation and management of riparian son AZ habitat general technical report RM 43 USU S depart- STEVENS LEL E BTB T BROWN JMJ M SIMPSON AND RRR R ment of agricultureofagriculture forest service tucson AZ JOIINSONJOHNSON 1977 the importance of riparian habitat to migrating birds pages 156 164 in RRR R johnson received 20 november 1995 and DA jones technical coordinators importance accepted 13 november 1997 preservation and management of riparian habitat general technical report RM 43 USU S department of Agriculture forest service tucson AZ

APPENDIX major vegetation types along the truckee river california and nevada USFWS 1993 codeode vegetation type and characteristics fmcsmcme sierra mixed conifer forest an open parklike forest of coniferous everevergreensgreens with crowns often touching several predominant species abies pseudotsuga and cornus are most common on moist sites pinus sppapp and ceanothus sppapp on dry sites the understory typically is sparse consisting of scattered chaparral shrubs and young trees elevation 1500 2100 m ip3 jeffrey pine A tall open forest predominated by jeffrey pine with sparse understory of montane chaparral or sagebrush sppapp elevation 2100 2750 m

1pap lodgepole pine typically a dense forest of slender trees up to 40 ra tall often in pure stands elevation 2100 2750 m bcc black cottonwood A fairly dense mixed riparian forest predominated by black cottonwood with jeffrey pine andor lodgepole pine the shrub and herb layers are well developed elevation usually 1800 m cww great basin cottonwood willow forest open canopied forest predominated by fremont cottonwood and salixsaidosalvesaime laevigatalaevigata primarily east of vista elevation usually 21002 100 in this type was further delineated by the presence of whitetop w added to ending of type code and by successional stage cwbcw2 shrub seedling 3 m tall cwbcw3 pole sapling cwbcw4 mature rs great basin riparian scrub open to dense riparian thickets usually composed of willow open stands may have a dense herbaceous understory elevation all but especially well developed along lower truckee 199811998 BIRD USE OF RIPARIAN vegetation 343

APPENDIX continued code vegetation type and characteristics ss sagebrush steppe A semiclosed steppe predominated by big sagebrush with several perennial bunchbunchgrassesgrasses between shrubs elevation below conifer zone in freshwater marsh predominated by perennial emergent monomonocotscots scirpus and typha dominate sites lack current often permanently flooded freshwater backwater of derby dam oxbow

a aspen dense stands of usually mature aspen located within conifer zone

9g gravel bars w whitetop low dense exotic forb dead stalks persistent and woody often is dominant understory for fremont cottonwood elevation 1800 m

ag agriculture mostly alfalfa creatgleatgreat basin naturalist 584 C 1998 appp 344 351

USE AND SELECTION OF BROOD REARING HABITAT BY SAGE GROUSE IN SOUTH CENTRAL washington

colin M sveumlsveum1 john A Crawfordcrawford22 and W daniel edgededge2

ABSTRACT sage giouseglousecrousegrouse Centrocentrocercuscercus urophasianus brood habitat use was examined during 1992 and 1993 at the yakima titrainingdining centelcenter in yakima and Kitkittitastitas counties washington during the 2 yr we followed 38 broods of which 12 persisted to 1 august afx3f approximately 151.51 5 chicksbroodchicks brood food forb cover was greater at all brood locations than at landomrandom locations hens with broods in big sagebrushbunchgrasssagebrusbbunchgrass habitat artemisiaArtemisiasiusta tridentataagropyrontridentatalagropyron spicaspicatumtum selected for greatelgreater food forbhorb cover total forb cover and lower shrub heights broodsbloods in altered big sagebrushbunch glassgrass habitats selected greatergi eater tall grass cover and vertical cover height broods in grassland showed no preference for any measuredmeasuiedmeasuied vegetation characteristics during the early rearing period post hatching 6 wk each year broods selected sagebiushbunchgrasssagebrushbunchgrass bloodsbroods in 1993 made greater use of grasslands than in 1992 and selected grassland during the late brood rearing period 7 12 wk broods selected for sagebrushbunchgrass during midday but 52 of brood locations in the afternoon weiewelewere in grassland tall glassgrass cover was greater at morning 0500 1000 h and afternoon 1501 2000 h bloodbrood locations than at midday 1001 1500 h and random locations midday brood locations had greater shrub covelcover and height than morning and afternoon locations selection of habitat components was similar to the results of other studies but habitat conditions coupled with a possible lack of alternate brood rearing cover types resulted in low survivalsuisul vivalvivai of chicks

kelikeifkey words broods centrocercusCentro cercus uiourophasianus habitat sage grouse washington

habitat use by sage grouse Centrocentrocercuscercus autenrieth 1981 brood movements to these phasianusurourophasianus for brood rearing was related to habitats are immediate or transitional depend- forb availability in previous studies peterson ing on annual precipitation temperature and 1970 oakleaf 1971 autenrieth 1981 drut proximity from nesting habitat meadows and crawfordgrawford and gregg 1994 drut pyle and upland sagebrush habitats however often have crawford 1994 sage grouse chicks require been grazed excessively causing reduced food protein rich foods including insects and forbs and water availability savage 1969 oakleaf for growth and development from hatching 1971 disturbance in these mesic habitats has through approximately 3 mon savage 1969 increased soil erosion facilitated invasion by dunn and braun 1986 bergerud 1988 drut exotic plants affected vegetative composition pyle and crawford 1994 changes in avail- and lowered water table levels oakleaf 1971 ability of these critical foods may affect sage heimannhofmannhermannhormann 1991 oakleaf 1971 found fewer grouse distribution and habitat selection sage grouse foraging in nevada meadows as wallestad 1971 pyle 1993 furthermore in food supplies diminished because of improper an oregon study drut pyle and crawford grazing practices and soil erosion in oregon 1994 sage grouse productivity was higher drut crawford and gregg 1994 noted that on an area where chicks fed on a diet of 80 broods used larger home ranges where forb forbs and insects than where chicks ate pri- availability is low than where forbs are relatively marily 65 sagebrush artemisia sppapp abundant this may have resulted in reduced sage grouse broods reportedly move from survival these studies reveal the importance nesting habitats to sites where succulent and of forbs and insects in relation to abundance abundant forbs persist ie meadows or upland distribution habitat selection and productivity sagebrush habitats as summer temperatures nevertheless the effect of massive landscape increase and moisture decreases nelson changes on habitat use by broods has rarely 1955 gill 1965 savage 1969 oakleaf 1971 been documented A full understanding of

unitedlunitedlunsted states geological survey biological resources division upper mississippi science center 2630 fanta reed road lacrosse WI 54603 23epaltmcnt of rishcnes and wildlife oregon state university colcorvallisCoi vallis OR 97331380397331 3803

344 199819981 SAGE GROUSE BROOD HABITATHARITAT SELECTION 345 factors affecting habitat use by broods espe- we identified 5 cover types available for cially where populations are severely depleted brooding sage grouse on the YTC 1 big in areas of massive habitat changes remains sagebrush A tridentatabunchgrasstridentatabmchgrsiss which is incomplete dominated by wyoming big sagebrush A t we studied sage grouse in washington wyomingensis rabbitbrushrabbitbrush chrysothamnus where extensive habitat loss approximately sppapp bluebunchbluebunch wheatwheatgrassgrass agropyron spica poa 70 and alteration have reduced sage grouse tum and sandberg s bluegrass sandbergsandbergiiii to 2 separate populations totaling an estimated 2 stiftstiff sagebrush A ngtdasandbergrigidasandberg s blue- 630 birds by 1994 tarhitirhi 1994 our objectives grass 3 altered big sagebrushbunchgrass were to examine selection of cover types by which contains substantially more bare ground sage grouse broods and describe structure and lesser amounts of shrub and herbaceous disturbance 4 and composition of habitat components within cover because of repeated grassland predominantly bunchbunchgrassesbuncligrassesgrasses with cover types ard3rd and ath4th order selection some rabbitbrushrabbitbrush but virtually devoid of sage- respectively see johnson 1980 used by broods brush because of frequent fires and 5 riparian with willow salix sppapp currant ribes STUDY AREA sppapp grasses sedges carex sppapp and baltic rush juncus balbaltitusbalticusticus the study was conducted on the yakima training center YTC a 1058 km2 area in METHODS yakima and kittitasKittitas counties washington which supported approximately 330 sage we captured sage grouse hens using spot- grouse during this study tarhitirhi 1994 eleva- lights and walkinwalk in traps giesen et al 1982 tions on the YTC range from 183 to 1249 in schroeder and braun 1991 and fitted the hot dry summers and cool dry winters typify birds with numbered aluminum leg bands and the study area growing season september necklace attached radio transmitters during august precipitation for 1962 1993 obtained march 1992 and 1993 each year when radio from the US department of commerce cli- telemetry monitoring confirmed nesting com- matmatologicalological database averaged 20420.4 cm grow- pletpletionion fate was determined by inspection of ing season precipitation in 1991 92 and eggshell membranes or in 3 instances 2 in 1992 93 was 16716.7 and 22522.5925985 cm respectively 1992 1 in 1993 observations of hens with above normal precipitation accompanied by young initial brood sizes were estimated from cooler than average temperatures began dur- shells from successful nests which were con- ing winter 1992 93 and continued through sidered initial brood locations we relocated marked hens with broods 1 summer 1993 these weather differences likely radio J timewk visual accounted for greater shrub cover and height and attempted to obtain observations without flushing either hens or broods and more tall grass and forb cover in certain if visual observations were not possible we used cover types in 1993 sveum 1996 recent sign droppings feathers as an because the YTC serves as a military train- ie indication of habitat use each brood location ing ground frequent off road vehicle use has was determined with a global positioning sys- in approximately 5 of the area being resulted tem in universal transverse mercator UTM covered by roads and trails JG stephan coordinates and marked with a flag to facilitate pacific northwest laboratory personal com- relocation for measuring vegetation A brood municationmunication this disturbance has increased was considered successful if J chicks were erosion and facilitated establishment of knap- observed with a radio marked hen after I1 weed centaurea sppapp and cheatcheatgrassgrass brome august the approximate date when brood bromus tectoriumtectorumtectorum cattle and sheep grazing integrity dissolves dalke et al 1960 oakleaf was initiated on the YTC in 1961 livestock 1971 we used a t test to examine the null stocking rates during 1992 and 1993 were 0130.130 13 hypothesis that mean brood sizes were the and 0150.15 animal unit monthshamonthsha fires on the same between years Z tests with a continuity YTC common during summer burned 535.3 and correction zar 1984395 397 were used to 808.0 km2 in 1992 and 1993 respectively LL test the null hypotheses that brood success rates cadwell pacific northwest laboratory per- and cover type use were the same between sonal communication years 346 GREAT BASIN naturalist volume 58

we examined ard3rd order selection of cover were combined and called food forbs for ath4th types and ath4th order selection of particular order analysis these forbs included milkvetchmilkvetch habitat components within cover types habi- astragalus sppapp clover trifolium sppapp hawks tat selection by hens with broods johnson beard crepis sppapp microsterismicrosteris microsterisMicrosteris 1980 availability of cover types ard3rd order gracigracilisfishis and species in the cichorieae milky was determined within a composite minimum juiced composites riparian and stiff sage convex polygon home range odum and kuen- brushbluegrassbrush bluegrass cover types were combined zler 1955 from pre nesting movements of for ard3rd order analysis because of infrequent radio marked hens with a geographic informa- brood use each year and were collectively tion system availability for ath4th order selec- called other 11 tion was identified by measuring randomly we compared cover types used by radio generated UTM coordinates within each cover marked hens with broods observed to the type from the composite home range during availability of each cover type expected with the 1992 93 brood rearing seasons chi square analysis for ard3rd order analysis if a we measured ath4th order characteristics of significant difference between use and avail- hens with broods at brood locations and ran- ability was detected bonferroniBonferroni simultaneous dom sites with 2 perpendicular 10lom m transects confidence intervals were calculated to iden- at each location the direction of the first tran- tify which cover types were used dispropor- sect was determined randomly canopy cover tionationatelytely neu et al 1974 byers et al 1984 of all shrubs along each transect was estimated the null hypotheses for ard3rd order selection following canfield 1941 cover of grasses were 1 broods used cover types during both forbs and litter residual cover and bare early and late brood rearing periods in pro- ground were estimated along transects with portion to their availability and 2 brood loca- ten 010 1 m2ma frames daubenmire 1959 resid- tions by diurnal period were in cover types in ual cover was defined as any dead upright proportion to their availability plant material and consisted primarily of rus- null hypotheses for ath4th order analyses were sian thistle salsola kahkaltkalikait sagebrush knapweed that 1 brood and random measurements within and tumble mustard sisymbrium altissimum cover types did not differ 2 there were no maximum height cm for shrubs and standing differences between early and late brood rear dead vegetation and droop height excluding ing periods and 3 there were no differences flowering parts for grasses were measured among times of day and random locations grass height was classified as short 18 cm fourth order data were treated with analysis or tall 18 cm following wakkinen 1990 and of variance ANOVAAN OVA for unbalanced data and gregg et al 1994 vertical vegetation cover protected least significant difference mean was measured at plot centers with a modified separation tests proc GLM SAS institute inc robel pole robel et al 1970 four readings 1989 vegetation variables with nonnonformalnonnormalnormormalnoimnormormalormainormal 2 each along the 2 perpendicular 10 m tran- distributions were transformed legitlogit transfor- sects were taken 4 m from the pole and 1 m mation for proportional data and log transfor- in height mation for height data however nontrans separate analyses were used according to formed means and standard errors are reported brood age class early post hatching 6 wk herein all statistical tests are 2 tailed and and late 7 12 wk previous research revealed considered significant at a 0100.10olo changes in habitat use and diets of chicks at approximately 6 wk of age martin 1970 RESULTS peterson 1970 drut crawford and gregg 1994 both ard3rd and ath4th order data inm 1993 we captured and fitted 85 sage grouse were apportioned into 3 diurnal periods hens with radio transmitters during march of morning 0500 1000 h midday 1001 1500 both years 45 in 1992 40 in 1993 11 h and evening 1501 2000 h too few broods clutches were hatched in 1992 and 27 in 1993 were monitored in 1992 to analyze diurnal eggs hatched from 22 april to 28 may in 1992 habitat selection primary forbs in the diets of and between 2 may and 19 june in 1993 sage grouse chicks inm oregon drut pyle and more than 80 of clutches hatched in the big crawford 1994 and idaho tennethautenriethAu 1981 sagebrushbunchgrass type initial brood size 199811998 SAGE GROUSE BROOD HABITAT SELECTION 347

TABLE 1 availability and use of cover types by sage grouse hens with broods during early and late brood rearing periods on the yakima training center yakima and kittitasKittitas counties washington 1992 93 brood rearing locations

1992 1993 availability early late early late cover type N nna1 225 n 74 n 6817 n 2410 big sagebrushbunchgrass 46 77b771b 71 68 38 grassland 34 9 29 24 58 altered big sagebrushbunchgrass 8 5 0 9 4 othercotheraother0 12 9 0 0 0

inn burnnurnnumberber of locationsbroodslocations broods b use greater than expected no symbol use inin proportion to availability use less than expected P 0100 10 by bonferronibonfeiromBonBonfeferroniherroniirom confidence intervalintervals cinecIncincludesludes riparian and stiff sagebrushbluegrasagebrusbbluegrassis

TABLE 2 availability and use of cover types by sage grouse broods during 3 diurnal periods yakima training center yakima and kittitasKittitas counties washington 1993

availability MorMorningninga1 midday afternoon cover type N n 1812b n 4915 n 2514 big sagebrushbunchgrass 46 67 650 44 grassland 34 28 25 52 altered big sagebrushbunchgrass 8 5 10 4 otherdothertlgotherd 12 0 0 0

Mo 0500 1000 h midday 1001 1500 h afternoon 1501 h i morningining 2000 lnn number of locationsbroodslocations broods c use greater than expected no symbol use in proportion to availability use lesiless than expected P olo0100 10 by bonfenonibonferromBonfeBonfer nonirom confidence intervals dincludincludes riparian and stiff sagebrushbluegrasssagebrush bluegrass

was greater in 1993 x 717.1 sifs 0420.42 than the mean time between location of a brood in 1992 x 575.7 st 0500.50oso t 1861.86 P and measurements for ath4th order characteris- 0070.07 in 1992 only a single hen was known to tics was 464.6 d recruit young 3 chicks into the august popu- sage grouse selected big sagebrushbunch lation but at least 11 broods x 151.5isls chicks grass during early brood rearing of each year survived to I1 august in 1993 several radio table 1 and used grasslands less than marked hens entered a restricted area during expected during the early rearing period of each summer and the fate of broods that 1992 no cover type selection was detected stayed in the restricted area 1 I1 in 1992 and 4 during the late rearing period in 1992 in 1993 is unknown consequently they are although 71 of locations were in big sage not included in success estimates an addi- brushbunchgrass during the late rearing tional brood was removed in 1993 after con- period of 1993 broods selected grassland and tact was lost with the radio marked hen used big sagebrushbunchgrass in proportion brood success in 1993 was greater than in to availability 1992 10 in 1992 and 50 in 1993 Z 2562.56 during the morning period in 1993 broods P 0ooiool0010.010 1 used all cover types in proportion to their we described ard3rd order selection by broods availability table 2 but at midday they at 29 locations from 5 broods in 1992 and 92 selected big sagebrushbunchgrass we found locations from 19 broods in 1993 no ath4th order no selection during the afternoon period how- data were collected in 1992 because most ever 52 of locationsoflocations were in grassland broods perished shortly after hatching fourth in 1993 brood locations in big sagebrush order data were collected from 72 locations bunchbunchgrassgrass had greater total forb and food from 17 broods and 30 random locations in forb cover and lower shrub heights than did 1993 brood locations in 1992 were not ana- random big sagebrushbunchgrass locations lyzed by diurnal periods for ard3rd order selec- table 3 brood locations in altered big sage tion because too few locations were obtained brushbunchgrass had greater tall grass cover 348 GREAT BASIN naturalist volume 58

tableTABLLTABLF 3 habitat characteristics of sage grouse brood and random locations by cover type on the yakima training centercenten yakima and kittitasKittitas counties washington 1993

Aiteredalterednitered big big sagebrushbunchgrasssage irushbuncagiass sagebrushbunchgrasssagebnishbunchish bunch grass G assasslandaaslandgrasslandland brood random brood random brood random n 43 n 10 nann55 n 10 n 24 n 10 variable S fscs3fs P xsrsxvsxas CST P x S xsFSas P shrub cover 142 202 012 74 123 039 42 32 088 shrub height cm 182 252 010 117 143 070 52 42 075 grassgi ass cover sholtshort 1818cmcm baibal201 202 078 202 192 073 212 233 059 tall 1818cmcm 171 194 076 216 83 004 213 152 024 forb covelcover 252 81 001 154 216 051 193 144 044 food forb cover 81 21 001 42 31 045 41 31 032 residual cover 102 31 001 104 11 041 104 105 031 residual covelcover height cm 103 305 003 21 11 077 205 11 024 vertical cover height cm 152 172 026 174 61 001 91 91 094 bare ground 322 354 054 406 494 022 433 475 043 litter 573 605 067 464 404 043 443 375 019

and taller vertical cover height than random cover in big sagebrushbunchgrasssagebrushbuncligrass apparently altered big sagebrushbunchgrasssagebrushbuncligrass locations provided the best cover for nest success and no differences were detected between brood early brood survival on the YTC and random locations in grassland food forb most late brood rearing locations in 1992 cover was greater at early and late brood loca- were in big sagebrushbunchgrass but in 1993 tions than at random locations table 4 late brood cover type selection switched to midday brood locations had greater shrub grassland hens with chicks made greatest use cover and shrub height than morning and of grasslands during the afternoon period in afternoon brood locations table 5 afternoon montana most early summer brood locations brood locations had less shrub cover and were in sagebrush grassland types but as forbs height than random locations morning and desiccated grouse shifted to black grease- afternoon brood locations had greater tall wood sarcobatus venniculatusvermiculatus and grassland grass cover than midday brood and random cover types in more mesic sites peterson locations during each diurnal period food 1970 wallestad 1971 savage 1969 found that forb cover was greater at brood locations than broods left sagebrush uplands in nevada dur- at random locations vertical cover was greater ing rapid temperature increases which accel- at midday brood locations than afternoon erated forb desiccation in sagebrush habitats brood and random locations broods may remain in sagebrush uplands when free water is available or during years when discussion abundant precipitation increases forb availability oakleaf 1971 dunn and braun 1986 broods exhibited similar patterns of cover fourth order analysis of 1993 data suggests type use during the early rearing period each that hens with broods in big sagebrushbunch year by selecting big sagebrushbunchgrass grass 60 of all locations selectively used which also was the primary nesting habitat sites with more total forbs 25 cover and after hatching before chicks can fly and when more food forbs 8 cover than available ran- mortality is highest patterson 1952 autenrieth domly 8 and 2 respectively during early 1981 broods need food in close proximity to and late rearing periods sites with greater escape cover random big sagebrushbunch amounts of key forbs were selected shrub grass locations had greater shrub cover and height and residual cover were slightly less at height than grassland and altered big sage brood locations than at random sites in big brushbunchgrass short and tall grass cover sagebrushbunchgrass in oregon drut pyle also was abundant in big sagebrushbunch and crawford 1994 found that total forb grass the combination of shrub and grass cover at brood sites which ranged from 11 to 199811998 SAGE GROUSE BROOD HABITAT SELECTION 349

TABLE 4 habitat characteristics of sage grouse brood locations during early and late brood rearing periods and random locations on the yakima training center yakima and kittitasKittitas counties washington 1993 early late random n 53 n 19 n 30 variable aesyes xstsiasesi TSx V x5es0 shrub cover IIhlabiiiabblablabIab 72 122 shrub height cm 142 93 142 grass cover short 18cm18 cm boo201goo 203 211 tall 1818cmcm 192 183 142 forb cover 222 233 153 key forb cover 61aglagia 62a 21b residual cover 103a10 3aaa 06020602aA 204b20420.4 residual cover height cm 10310.3 10310.3 204 vertical cover height cm 142 123 111 bare ground 352a 393ab393 AB 443b443 litter 522 504 463

means within row without letters 01or with the same letter aieare not significantly different P olo0100 10 meanseans within row with different letters are significantly different pap5P 0100 10

14 during early brood rearing and from 19 but 52 of afternoon locations were in grass- to 27 during late brood rearing influenced land which coincided with evening foraging cover type use likewise klebenow 1969 morning and afternoon locations differed from schoenburg 1982 and dunn and braun 1986 midday and random locations by having found more forb cover at sage grouse brood greater tall 18 cm grass and less shrub locations than at random locations relatively cover and height total forb cover and food low availability of forbs in big sagebrush forb cover at brood locations were not signifi- bunchbunchgrassgrass apparently resulted in strong selec- cantly different among diurnal periods day- tion by broods for forbrichforb rich areas within this time brood locations had greater forb and food important brood rearing cover type brood forb cover than random locations dunn and locations in altered big sagebrushbunchgrass braun 1986 found that during the morning had greater tall 18 cm grass cover and verti- broods fed in open homogeneous areas and cal cover than random locations within this during the rest of the day used areas with cover type suggesting that broods seek pro- more horizontal cover and greater variation in tective cover when using this less preferred sagebrush canopy cover to roost and rest sav- cover type no ath4th order vegetation selection age 1969 found that broods fed shortly after was observed for grassland brood locations sunrise loafed in sagebrush during midday compared with random locations probably and moved to feeding areas in the evening because forb cover was sufficiently abundant fourth order location measurements during the throughout and the abundant tall and short morning and afternoon suggested that broods grass cover likely provided adequate conceal- left dense cover to feed but total and food forbs ment for escape were abundant at midday locations as well broods during midday selected big sage late summer mean brood sizes from studies brushbunchgrass and were observed loafing elsewhere ranged from 232.3 to 393.9 chickshenchick shen and dust bathing under large sagebrush mid- keller et al 1941 patterson 1952 nelson day brood locations had greater shrub cover 1955 savage 1969 wallestad and watts 1973 shrub height and vertical cover than morning although brood success increased significantly and afternoon locations midday brood locations on the study area in 1993 the number of also had greater vertical cover height than chicks recruitedhenrecruited hen was lower than in other random locations several studies described studies and may be insufficient to maintain grouse loafing during the midday after morn- population stability brood success was greater ing feeding nelson 1955 gill 1965 savage in 1993 than in 1992 which probably resulted 1969 oakleaf 1971 autenrieth 1981 broods from weather conditions during the 2 yr more used cover types in proportion to their avail- precipitation occurred in 1993 accompanied ability during morning or afternoon periods by cooler temperatures that lasted into the 350 GREAT BASIN naturalist volume 58

tabotablefableTABUTABLL 5 habitat characteristics of sage grouse brood locations by diurnal period and random locations on the yakima trainingti dining center yakima and kittitasKittitas counties washington 1993

diurnal period 1 morning midday afternoon random n 13 n 39 n 20 n 30 vaivalvariablelablelabie YSTest xsast80 5fsox xsas3fso shrubshishl abub covelcover 72ablc72abbc 152c 41b 122ac122 AC shrubshishl abub height cm 92ab 182c 62b 142ac142 AC glassgrass covelcover short isem18cm18 cm 203 191 222 211 tall 18cm18 cm 222a 162b 223a 142b folbforb covelcover 213 222 224 153 key tolbfolbforb covelcover 61agiagla 71a 61aglagia 21bgib residual covelcover 104 103 104 204 residual covelcover height cm 104 104 205 204 veivelverticaltical covelcover height cm ililii111ab11111 aa1aABB 162b 92a lilaliia111a11 JA bare ground 365 362 363 443 littellitter 514 523 524 463

morning omoo0500OTOO 1000IOOO li midday 1001 1500 h afternoon 1501 2000 h iveansimeansmc insuns within lowowtowrow witritliwitli different letters aiealeare significantly different P olo0100 10 meanst ins within lowtowrow without letters 01or with the sainesame letterietter are not significantly different P 0100 10 summer increased precipitation in nevada populations in western states hens did not have resulted in greater forb production delayed the choice of alternative high quality brood plant desiccation and possibly enhanced juve- rearing habitats such as low sagebrush A nile survival oakleaf 1971 peterson 1970 arbusulaarbarbusculausula meadows lakebedslakebeds or broad forb- found greater brood success during wet years rich drainages lack of these critical cover in montana when forb production was 2 3 types at the YTC coupled with existing habitat times that of dry years tennethautenriethAu 1981 noted conditions likely had an adverse effect on that migratory populations of sage grouse had recruitment which may limit populations on high brood success because they were able to the YTC find forbs whereas sedentary populations like those at the YTQYTC had good reproductive suc- acknowledgments cess during moist years when forbs were study funded by pacific abundant but did poorly during dry years an this was northwest laboratory under a contract with the US increase in food and cover on the YTC in 1993 increase in in of defense klabon may have reduced brood movements result- department DOD JA kollasch MA lacroix AW and in lower predator exposure and energetic TM leary ing in hand assisted with data collection we costs of foraging nevertheless by 1 august KD thank cadwell and the late one half of the broods were lost and the LL LE eber- hardt for logistical support and liaisons with remainder declined from a mean of 717.17 1 to 151.5isls1 5 DOD MA gregg provided valuable comments chicks approximately an 80 loss on the manuscript this is manuscript 11112 of we concluded that nesting success 45 the oregon agricultural experiment station was rather typical for sage grouse but brood 1 august 14 and especially recruitment to literature CITED the number of chicks recruited 24 from 45 successful hens during the 2 yr were far below autenrieth RER E 1981 sage grouse management in average we suggest that brood rearing habi- idaho idaho department of fish and game wildlife tat was a strong limiting factor for this small bulletin 9 238 appp BERGERUD A T 1988 population ecology of north ameri- population our results indicated that big sage AT can grouse pages 578 685 in AXA T bergerud and brushbunchgrass and grasslands are impor- MWM W gratson editors adaptive strategies and pop- tant cover types throughout brood rearing ulation ecology of northern grouse university of within brood rearing habitat there was selec- minnesota press minneapolis for with forb cover especially BYERS CRC R RKR K steinhorst AND PR KRAUSMAN 1984 tion sites greater clarification of a technique for analysis of utilizautilize forbs used as food and shrub or grass cover for tion availability data journal of wildlife manage- concealment unlike many other sage grouse ment 48105048 1050 1053 199819981 SAGE GROUSE BROOD HABITATHARITAT SELECTION 351

CANFIELD R 1941 application of the line interception OAKLEAF RJR J 1971 the relationship of sage grouse to method in sampling of langeiangerange vegetation journal of upland meadows in nevada unpublished mastelmaster s forestry 3938639 386 394 thesis university of nevada reno 64 appp DALKE PD DBD B PYRAH DCD C STANTON JEJ E CRAWFORD ODUM EPE P AND EJE J KUENZLER 1955 measurement of AND E schlatterer 1960 seasonal movements territory and home range size in birds auk 72 and breeding behavior of sage grouse in idaho 128 137 transactions of the north american wildlife confer- PATTERSON RLR L 1952 the sage grouse of wyoming ence 2539625 396 407 sage books inc denver CO 341 appp daubenmire RFR F 1959 A canopy coverage method of PETERSON JG 1970 the food habits and summer distri- vegetation analysis northwest science 3322433 224 227 bution of juvenile sage grouse in central montana DUNN PO AND CEC E BRAUN 1986 summer habitat use journal of wildlifeofwildlife management 3414734 147 155 by female and juvenile sage grouse journal of wild- PYLE WH 1993 response of brood rearing habitat of life management 5022850 228 235 sage grouse to prescribed burning in oregon DRUT MSM S JAJ A CRAWFORD AND MAM A GREGG 1994 brood unpublished master s thesis oregon state univer- habitat use by sage grouse in oregon great basin sity corvallis 47 appp naturalist 5417054 170 176 ROBEL RJR J JNJ N BRIGCSBRIGGS ADA D DAYTON AND LCL C HULBERT DRUT MSM S WH PYLE AND JA CRAWFORD 1994 tech- 1970 relationship between visual obstruction mea- nical note diets and food selection of sage grouse suresurementsments and weight of grassland vegetation jour- chicks in oregon journal of range management nal of range management 2329523 295 297 479047 90 93 SAS INSTITUTE INC 1989 SASSTAT user s guide version glesenGIESEN KMK M TJ schoenberg AND CEC E BRAUN 1982 6 volume 2 ath4th edition SAS institute inc carygary NC methods for trapping sage grouse in Coloiacoloradodo wild- 795 appp life society bulletin 1022410 224 231 SAVAGE DED E 1969 the relationship of sage grouse to GILL RBR B 1965 distribution and abundance of a popula- upland meadows unpublished master s thesis uni- tion of sage grouse in north park colorado unpub- versity of nevada reno 101 appp lished master s thesis colorado state university schoenburgSCHOEN BURC TJ 1982 sage grouse movements and habi- fortcolhnsfort collins 147 appp tat selection in north park colorado unpublished GREGG MAM A JAJ A CRAWFORD MMSS DRUT AND AKA K mastermasterss thesis Coloiacolliadocoloiadocoloradodo state university fort DELONG 1994 vegetational cover and predation of collins 86 appp grouse sage in oregon journal of wildlife manage- SCHROEDER MAM A AND CEC E BRAUN 1991 walkinwalk in traps ment 5816258 162 166 for capturing greater prairie chickens on leks jour- HOFMANN LAL A 1991 the western sage grouse centro nal of field ornithology 6237862 378 385 cercus urophasianus phaiosphacos on the yakima training SVEUM CMC M 1996 habitat selection by sage grouse hens center in central washington unpublished master s during the breeding season in south central wish-wash- thesis central washington university Ellensellensburgellensbergburg ington unpublished mastelmaster s thesis oregon state 177 appp university corvallis 86 appp JOHNSON DHD H 1980 the comparison of usage and avail- TIRHI MJM J 1994 management of sage grouse centro ability measurements for evaluating resource prefer- cercus urophasianus in washington washington de- ence ecology 6165gigs61 65 71 partpartmentment of fish and wildlife olympia 109 appp KELLER RJR J HRH R SHEFERDSHEPERD RNR N RANDALL 1941 survey WAKKINEN WL 1990 nest site characteristics and spring park of 1941 north jackson county moffit county summer movements of migratory sage grouse in including comparative data of previous seasons sage southeastern idaho unpublished master s thesis grouse survey colorado game and fish depart- university of idaho moscow 57 appp ment denver 31 appp WALLESTAD ROR 0 1971 summer movements and habitat klebenowKLERENOW DAD A 1969 sage grouse nesting and brood use by sage grouse broods in central montana habitat in idaho journal of wildlife management journal of wildlife management 3512935 129 136 3364933 649 662 WALLESTAD ROR 0 AND CRC R WATTS 1973 factors affect- MARTIN NSN S 1970 sagebrush control related to habitat ing annual sage grouse productivity in central mon- and sage grouse occurrence journal of wildlife tana federal aid to wildlife restoration job progress management 3431334 313 320 report W 120 11 3 montana fish and game depart- NELSON 0OCC 1955 A field study of the sage grouse in ment helena 23 appp southeastern oregon with special leferencereference to repro- ZAR JHJ H 1984 Biostatistical analysis and2nd edition prentice duction and survival unpublished master s thesis hall inc englewood cliffs NJ 718 appp oregon state university Corvalcorvallishs 112 appp NEUCWCRNEU CW CR byersandjmBYERS AND JM PEEK 1974 A technique received 5 february 1997 for analysis of utilization availability data journal of accepted 24 november 1997 wildlife management 3854138 541 545 gleatgreat basin naturalist 584 0 1998 appp 352 362 SOIL vegetation RELATIONS OF recovering SUBALPINE RANGE OF THE WASATCH PLATEAU

james 0 Klemmedsonklemmedson11 and arthur R Tiedemanntiedemann22

absrraclabsnlaur on degraded subalpine langeiangerange of the wasatch plateau we examined the hypothesis that lecoveryrecovery of veg- etationetation as manifested by its composition and biorbiomassbiornassbiornessnass yield was related to soil phosphorus P and sulfur S status we sampled 6 topographic locations to determine the relationship among composition and yield of grasses and forbs litter covelcovercoven and soil characteristics including lockroekrock cover organic carbon co total N ntN available nitrogen nynav total phosphorus Ppt organic01 game P poP inorganic P piP total potassium K total S stS and element ratios we also evaluated aspect effects an alternativealtel native hypothesis was that productive potential was a function of depth of soil leremainingmaming after the period of destructive grazingglazing differences among locations were significant for all vegetal attributes and for all soil characteristics except total K and coC aspect was significant only for forb yield and ptP regression coefficients for yield and percentage composition of grasses were always opposite in sign to those for forbs yield and composition of grasses and forbs as groups weiewelewere oppositely and strongly related to soil element ratios ofofcaofcpcoptCP NPQ coptcop and costcos but were not i elatedrelated to soil Ppt 01or stS there was no clear support for acceptance of the hypothesis that soil P andor S were malormajor factors in recoversrecovery of this subalpine langeiangerange after destructive grazing differences in regression coefficients and lower r val aesues among species within grass and forb groups than for the groups themselves to soil nablesnabiesvavariables is a reflection of species individuality this indicated a need to examine soilvegetationsoil vegetation relationships at the species level percentage com- positions of gigrassesasses and forbs were oppositely related to the depth of A B horizon lending support to acceptance of the alternativealtet native hypothesis

key words summer range plant composition and cover herbage yield litter soil C N P S and K

after 35 years of destructive grazing by cat- our observations suggest that improvement tle and sheep in the late 1800s the subalpine in soil and vegetal conditions reached a plateau range of the wasatch plateau east of ephraim about 1930 1940 based on ellisorsellisonsellisonsrEllisorssons 1954 utah was in poor condition reynolds 1911 records and has remained essentially the same sampson and weyl 1918 sampson 1919 from the time of ellisorsellisonsElliellisonsrsolssorssons studies intermoun- depletion of vegetation reached such severe tain research station provo utah unpublished proportions that most of the soil A horizon was data johnson 1964 these observations led us lost by erosion and mudrockmud rock floods were a to ask why after 30 40 yr of rapid improvement common occurrence in the canyons and val- under reduced grazing pressure should sec- leys below reynolds 1911 croft 1967 in ondary succession apparently stabilize at a mid many places only subsoilssub soils remained when seral stage and remain so until the present grazing regulation was begun with the 1903 there are perhaps several possible explana- establishment of the manti national forest tions for the apparent stable state lewontin reynolds 1911 sampson and weyl 1918 elli- 1969 laycock 1991 that exists on the wasatch son 1949 transient livestock herds were abol- summer range two explanations derive chiefly ished and livestock numbers greatly reduced from degradation of the ecosystem and massive but most of the summer range was so badly erosion that occurred over the long period of deteriorated that these management changes livestock overgrazing 1 loss of most of the were insufficient to halt continuing soil loss soil A horizon and hence most of the soil ellison 1954 although condition of the range organic matter and nutrient capital and alter- improved over the next 4 decades most of the ation of nutrient cycling processes nikiforoff summer range was still unstable in 1950 and 1959 anderson 1988 2 loss of extinction accelerated erosion was continuing but at much prone perennial grasses mack and thompson reduced rates ellison 1954 meeuwig 1960 1982 oconnor 1991 which are the key climax

school of renewable natural resources 325 biological sciences east building univeisityuniversity ofanzonaof arinaarlariAnzonana tucson AZ 85721 2paclfic2pacific noithwestnorthwestnortirwest research station 1401 gekeler lane LaGilagrandeande OR 97850 correspondingcone&pondmg author

352 199811998 SOIL VEGETAL RELATIONS OF recovering SUBALPINE RANGE 353 dominantsdominants in similar subalpine landscapes ous shales with minor interinterbedsbeds of sandstone throughout the west and were thought by oil shaleshaie conglomerate gypsum and volcanic sampson 1919 to have dominated the pris- ash weber 1964 schreiber 1988 soils of the tine vegetation here plateau are mostly fine mixed argieargle cry although both explanations for the appar- oborollsoborolls but lithic pachie and vertic cry ent static condition of soil plant systems on oborollsoborolls also are present they are shallow to the wasatch summer range have merit we moderately deep subsoilssub soils are silty clays or focus on the former in this paper more specif- clay loamscoams thickness of the A horizon aver- ically we hypothesized that losses of soil phos- ages just 4 cm that of the B horizon averages phorus P andor sulfur S following the 52 cm range 30 74 cm based on typical period of destructive grazing and erosion have profile descriptions HK swenson natural diminished levels of one or both nutrients to resources conservation service boise ID the extent that accumulation of organic C and personal communication these relative hori- N to pre degradation levels has been impeded zon thicknesses indicate that much of the orig- walker and adams 1958 cole and heil 1981 inal A horizon was lost by wind and water ero- this in turn has limited soil development dur- sion following the period of unrestricted grazing range recovery an alternative hypothesis ing prior to 1903 was that neither P nor S was limiting relative vegetation of the wasatch plateau is chiefly to other elements but that soil loss was so herbaceous but patches of engelmann spruce extensive that productive potential is now picea engelmanniiengelmanniaengelmannii and subalpine fir abies largely governed by the amount of remaining lasiocarpalasiocampa occupy steep northerly exposures soil ie A and B horizon under either hy- of east west ridges and dot the plateau land- popothesisthesis regaining climax conditions of the scape because remnants of pristine vegeta- former ecosystem would seem to require soil tion do not exist ellison 1949 1954 opinions formation over a very long time to reestablish differ regarding its exact character ellison the original steady state soil profiles character- 1954 described the original herbaceous com- istic of the pre 1870 climax soil plant nutrient munity as mixed upland herb dominated by system olsen 1958 jenny 1980 tall forbs while sampson 1919 considered wheatwheatgrassesgrasses agropyron sppapp the primary cli- STUDY AREA max dominantsdominants

the study area is centrally located on the METHODS wasatch plateau about 17 km east of manti utah the area extends south 7 km from near the original strategy for testing the hypothe- the alpine station along skyline drive road ses was a comparative analysis of paired eroded 139 to snow lake the long narrow plateau and uneroded soil plant systems at various is oriented approximately north and south with locations however after an exhaustive search riblike ridges extending east and west the it was apparent that grazing use of this sum- plateau top is gently rolling but gradient steep mer range had been so complete during the ens up to 65 on slopes of east west drain- period of devastation that uneroded sites were ages average annual precipitation is about nonexistent even on plateaus isolated by steep 840 mm 23 of this falls as snow between terrain that we believed would limit access to november and april precipitation averages livestock 173 mm during summer months june through instead we selected 6 topographically sep- september but varies considerably mean arated locations mostly small knolls and sam- annual temperature is about OOC ellison 1954 pled soil and plant attributes on up to 4 aspects during the growing season may through all 6 locations had a similar grazing history october average maximum temperature is until the 1930s since then elk knoll EK and 21c average minimum is 5cac ellison 1954 alpine cattle pasture CP have been pro- soil parent materials are of the flagstaff tected from livestock ideally this would give formation stanley and collinson 1979 that us an array of site conditions that would per- outcrop over about 7200 km2 in central utah mit differentiation among locations and aspects schreiber 1988 the dominant lithology is based on soil parent material and vegetation freshwater lacustrine limestone and calcaracalcare properties and permit determination of key 354 GREAT BASIN naturalist volume 58 variables influencing herbage composition and digestion bowman 1988 inorganic P pipj was production on the summer range determined with the same color development on each location we attempted to restrict on samples ignited at 550c5500c for 2 h olsen and aspect sampling points to a single parent mater- sommers 1982 while organic P PO was ial stratum hence elevation among aspects determined by difference available nutrients was near constant however this was probably Xxyaxy were determined as follows P by ascor- futile because individual parent material strata bic acid color development following 050.5os M were usually very thin 05os050.5 m klemmedson sodium bicarbonate extraction olsen and and tiedemannTiedemalmmarm 1994 there was little confi- sommers 1982 N by steam distillation of 2 N dence in sampling the same parent material KCI extracts keeney and nelson 1982 and S among all aspects of a location all locations with 11 water extracts followed by ion chro- were within 7 km of each other northeast matomatographygraphy dick and tabatabai 1979 SE SW and NW aspects were sampled on to facilitate comparison among sites and elk knoll EK elevation 3116 m and a knoll aspects we summarized soil horizon data and adjacent to snow lake SL elevation 3133 m expressed the data for the 0 to 15 cm soil two aspects were sampled on trail ridge TR layer and for the entire solum the data were elevation 3216 m skyline drive SD eleva- analyzed by 2 ANOVAs 1 for EK and SL knolls tion 3200 m and alpine cattle pasture CP with data for all 4 aspects the and2nd with data elevation 3066 m a single aspect was sam- from all 6 locations where the number of pled on south knoll SK elevation 3109 m aspects sampled was unequal in the latter slope gradient was 10 30 among aspects on ANOVA the interaction term was calculated SL gradients were 5 on other locations using data only for EK and SL locations back- we sampled at selected aspects from ran- ward stepwise multiple regression analysis domly located soil pits and vegetal litter cover was used to relate herbage yield and composi- plots soil pits were dug and profiles described tion to soil surface and 0 to 15isemcm soil layer by standard terminology single samples of properties known volume were taken from each horizon 3 5 above the C or R horizon for laboratory RESULTS AND discussion analysis herbage and litter were sampled in 6 location and aspect randomly located 05 m2ma plots near each pit differences basal cover of litter bare ground and rock vegetation AND SOIL COVER analysis of and foliar cover by species were visually esti- variance for all locations showed significant mated mass of grasses forbs and litter was differences among locations for 6 attributes of determined by harvesting each component vegetation and cover at the FP 5 0050.05 level separately followed by oven drying 70c700c table 1 of these forb yield was the only and weighing attribute that also differed significantly among for chemical analyses we air dried soils aspects table 1 it was highest on NW aspects sievedsievek them to remove the 2 mm fraction and lowest on NE and SE aspects table 2 and then ground them to pass a iso150 rmlm sieve the significant location response of forb com- samples were analyzed for total C by dry composition by foliar cover must be qualified bustion nelson and sommers 1982 in a because of the significant location X aspect LECOLEGO high frequency induction furnace interaction table 1 forb composition re- lecolegoLECOLEGO corp st joseph MI organic C Ccp sponded differently to aspect at EK and SL of soils was determined by difference after locations especially on SE and SW aspects determining carbonate by a gasometric table 2 this response may at least in part method dreimanis 1962 total N ntN was be influenced by parent material at the SL determined by semi micro kjeldahl bremner location in a companion study klemmedson and mulvaney 1982 and total S stS by dry and tiedemannTiedemarm 1998 parent material was combustion in the LECO high frequency highly associated with vegetal properties induction furnace tiedemannTiedemaimmarm and anderson based on values for the 6 vegetation and 1971 total soil P ptP was determined using cover attributes discussed above the locations ascorbic acid color development olsen and appear to form 2 distinct groupings fig 1 sommers 1982 following hydrofluoric acid the EK CP and SK locations were similar for 199811998 sollsoilnegetalSOIL VEGETAL RELATIONS OF recovering SUBALPINE RANGE 355

TABLE 1 probability values from analysis of variance for all locations and for elk knoll EK and snow lake SL locations alone

probability valuevalues s all locations ekanakanEK andasldslSL variables location aspect lxaalxa location aspect vegetation AND SURFACE COVER total yield gmegm2 0033 NS NS NS NS forb yield gmegm2 0001 0038 NS 0004 NS grass composition N 0036 NS NS 0090 NS forb composition 0034 NS 0050 0090 NS bare ground 0006 NS NS NS NS rock cover 0029 NS NS 0004 NS

SURFACE SOIL 0 15 CM organic C kgm2 0071 NS NS 0053 NS available N gmegm2 0001 0099 NS NS NS total P gmegm2 0009 0010 NS 0002 0049 inorganic P gmegm2 0010 0069 NS 0013 NS organic P of total 0013 NS 0010 0047 NS total K kgm2 NS NS NS 0019 NS COPcopt 0004 NS NS 0005 NS cost 0007 NS NS 0001 NS NPntptnept 0001 NS NS 0001 NS ntpftpntpo 0002 NS NS 0138 NS

TABLE 2 influence of aspect on forb yield and interaction of location and aspect on forb composition by foliar cover aspect NE SE SW NW LSD forb yield gmegm2 75 76 98 128 24 forb composition 17 elk knoll 67 84 87 90 snow lake 61 53 28 72

P 005oos0 05

composition of grasses and forbs less so for than the EK and CP locations and vegetal cover ofbareof bare ground and rocks and differed composition was about equally divided between in herbage yield fig 1 yield of forbs and grasses and forbs these locations SL TR total herbage for the EK and CP locations was and SD all had large amounts of bare ground similar and much greater than that for the SK and exposed rock fig 1 location herbage yield at all 3 locations was the 6 locations break out into the same dominated by forbs 80 of yield and com- groupings based on species composition of position these locations had little exposed the 25 species comprising at least 3 of the rock 5 and moderate amounts of bare composition table 3 only 2 grasses agropy- ground 13 26 ron trachycaulumtrachycaulum and stipa lettermanilettermannlettermani and I1 the SL TR and SD locations form the and2nd forb achillea millefoliummillefolium occurred on all 6 group they were similar to each other for locations composition of these 3 species was most attributes especially yield of forbs and similar among the SL TR and SD locations composition of grasses and forbs fig 1 they and from 202.0io10 to 66 fold higher than that for had significantly lower total herbage yield the EK CPCV and SK locations the 2 groups of 356 GREAT BASIN naturalist volume 58

250 uz3uza total M forbs 300 1 J lotalotoiotototal hP3 Ea inorganicino racrgcanlcanac P

20020q L tn 240 N E 1 E i 1 150ISO 1 180 I1 0 it j- CL 1 6 X T 100 t 120 J 0ID 0 1 1 E 1 50 a 1 t ra 60 1 X M Ail INH an N ivl 0n km ivl 1n 601 0 100 a grass M forbs 60 A CL 80 J iY 45 c 60 0 0 w i L 30 0 40 T 0 CL E 1 1 2.2 I I 1 C 80 20 I 15 N n 0 0n jl 0

60 I1 1 bare ground f rock 16 3 cgcorg kgm2 W 2 JL navnovnownopNQV gmgm2gme 45 C M 12 N 0 j E X 30 W0 8 X i 1 00 i 15 2L I1 4 JL 1 t rfaafa 3 h 1 1 f 0n ti EK SL TR SD CP SK 0 N location EK SL TR SD CP SK location fig 1 effect of location on yield offorbsof forbs and total vegetation composition by foliar cover of grasses and forbs fig 2 effect of location on amounts of total and inorganic and cover ofbareofbare ground and rocks P percentage organic PE and amounts of organic C and available N in soils locations also differed in distribution of several forbs six species asterfoliaceusaster foliafoliaceusfoliaceouspollaceus v canbyi among the 6 locations table 1 differences erigeron speciosusspeciosus geranium fremonfremontiifremontiatii ligus- were significant at the P ooi0010.01 level for nynav ticum porteriporponteriteri fenstemonbenPenFenpenstemonpentstemonstemon rydbergrydbergiiii and poten- pt piP and the cypcoptgypgop coscostcys and npfnepfntpt ratios at tilla gracilis were found only on the EK CP the FP 0050.05 level for percentage popy and at P SKS K at least 2 of the 3 locations and locations 0100.100 10 for coCQ in the case of popy content as a per- while 2 species artemisia ludoludovicianaviciana v inm centage of ptP there was a significant L X A compta and cymopterus lemmoniinii occurred cometa lemmo interaction table I1 popy was markedly higher only on the SL TR and SD sites table 3 at the EK than the SL location for all aspects in a companion study on the SL location klemmedson and tiedemannTiedemarm 1998 C lem except SW joniimonii was highly associated with rocky sites results of the analysis of variance for EK with shallow soils of low nutrient content and SL locations alone table 1 were similar SOIL properties nine properties of the to that for all locations with 2 exceptions 0 to 15isemcm soil layer differed significantly these 2 locations did not differ significantly in 199819981 sollsoilnegetalSOIL VEGETAL RELATIONS OF recovering SUBALPINE RANGE 357

TABLE 3 species composition at 6 knoll locatilocationsonsa1

knoll locationlocationllocationslocation1111

vegetal component EK SL TR SD CP SK

GRASSES AND GRASSLIKE agropyron trachycaulumtrachy caulum 76 105 131 89 36 34 alopecurus pratensisprapratenszstensis 34 carex microptera 36 stipa columbiana 67 stipa letterietterletietlettermanilettermamlettermanntermanimam 38 282 305 366 52 60 others 32 10 06 27 30 08 total 182 464 442 482 118 136

FORBS AND SHRUBS achillea millefoliummillefoliumliumilum sspasp lanulanulosalosa 84 126 150 113 58 56 artemisia ludovicianaludoviciana v incompta 188 47 197 asterfoliaceusaster fohaceus v canbyi 13613.613 6 astragalus miser v oblongifoliusoblongifohus 303.03 0 castilleja sulphureasulphured 46 52 cymopterus leimomilemmoniilemmomilemmoniinil 52 98 106log10 6 erigeronengeron speciosusspeciosus 53 49 34 erigeronengeron ursinus 828.28 2 geraniumfremontiigeranium frefremontufremontefrhbremontu 13513.513 5 22422.422 4 lesquerella utahensis 333 3 ligusticum porten 52 82 Penpenstemonpentstemonstemon rydbergii 109log10 9 44 36 potentilla glanduloseglandulosa 343.43 4 potentilla gracilisgraczitslisils 31 10810.810 8 solidago parryiparrys 55255.255 2 swertia perenisperennisperenmsperennisfennis 303.03 0 taraxacum officina le 46 65 valeriana occidentoccidentalisoccidentahsoccidentalistalisails 42 36 vicia americana v americana 64 46 vicia nuttalhiliiiliill v nuttalnuttalliinuttallialii 585.85 8 others 188 60 52 69 126 98 total 818 586 558 518 883 864 ababaseded on foliar cover species with 3 composition not listed bseeabseesee text for names and descriptionsdeseidesel iptionsoptions of locations

nynav but they did differ in total K moreover tionaldional differences among locations fig 1 there was a significant L X A interaction for Ppt table 3 and give clues to the seraiseral plateau although ftpt was higher at SL than at EK fig these systems have been in for the past 50 yr 2 the absolute and proportional differences simple linear regression relating vegetal were a function of aspect attributes with soil properties especially those for soil properties the 6 locations did not shown to be significant in table 1 indicates break out into the 2 groups observed for vege- that grasses and forbs as groups responded tation and cover attributes those groupings differently but consistently to these variables were apparent only for I1 of the 9 soil proper- table 4 in fact for each independent vari- ties gopCQPcopt ratio found to differ among loca- able shown in this table for regressions with tions fig 3 analysis of data for variables of herbage yield and composition as dependent the entire solum added very little information variables the regression coefficients for grasses to that shown for the 0 to 15isemcm soil layer and were always opposite in sign to that for forbs hence are not shown here certainly this was not the case for all independent variables we sampled but for the soil vegetation relations large majority the trend was very noticeable the marked differences described above in in a companion study at the SL location vegetal attributes and soil properties among klemmedson and tiedemann 1998 the dom- locations caused us to pursue further soil veg inant grass stipa letlettermanilettermanntermani and forb cymoncymop etationelation relations that might explain legetavegeta terus lemmolemmoniinii were oppositely related for 358 GREAT BASIN naturalist volume 58

200 30 cafcpf 0 cyle r 07100.7171 gost 0 gefrgear r 0520.520 52 V cost 0 iso150 ntptnept X 10 20

loo100 0 0

E 10 IV I1 0 W 50 0 71 0 07 1 a 0 6 0189biag E M i 2.2 0 e 0 u 60 80 100 120 EEKK S L T R D K SL TR SSD CCPP SSK V 0 agatrgat ratio location CLQ- FE 50 0 0 agtragar r 0330330.33 fig 3 effect of location on elemental of soil ratios 0 40 9 sile r 061ogi0.610 61

30 every variable sampled but for groups of 20 0 grasses and forbs comprising many species 0 0 table 3 it is remarkable that these groups 0 10 would respond min an opposite manner so con- 00 sistently to various growth soilsitesoil site parame- 0 0 ters although not always with high r values 10 20 30 40 50 60 table 4 that correlation coefficients associ- gorbatcorgptgorqat rioratio ated with specific independent variables in fig 4 Relationsrelationshiphipbip of composition by foliar cover of table 4 were not high as a rule is a reflection cymopterus leimonalemmonalemmoniilemmoniinil and geraniumftemontiigeranium fremonfremontnfremontepremontn to cstcoscys ratio of the individuality of species within grass and and composition of Agropyron trachycaulumtrachy caulum and stipa let forb groups in response to location and aspect teanite ani to capcopcgpgopCopt ratio figure 4 illustrates this individuality ofaof2of 2 grasses and 2 forbs in their response to coptcop and costcos variables that were both highly corre- which may have significantly influenced vege- lated with grass and forb composition table 4 tal yield and composition contrasting responses of grass and forb groups table 5 summarizes our attempt to predict also have been observed by huenneke et al herbage yield and composition with 2 vanablevariable 1990 in short term fertilization experiments equations using backward multiple regression on serpentine soils within vegetal groups hav- overall percentage rock cover was the most ing common properties species responded to efficient predictor of yield and composition of experimental treatments in an individualistic both grass and forbs based on standardized manner also chapin and shaver 1985 ob- regression coefficients percentage litter cover served this kind of species community behav- and bare ground were about equal as predic- iorlor in response to manipulation of environment tors of total yield popg was as efficient as soil in tundra surface features only in the grass yield equa- of those variables one would ordinarily tion when a ard3rd variable was allowed in the associate with soil fertility only co and the 2 equation 3 variable equations not shown p0papp element ratio variables were correlated with filled that role in 4 of 5 cases based on varia- yield and composition table 4 the high r tion explained in the 3 variable grass yield values for percentage litter cover and rock equation cocg became the ard3rd and most efficient cover are at first puzzling however when one variable that variables portraying soil surface considers the extent of correlation usually features would appear as the most efficient observed among soil plantpiant litter properties predictors of vegetal yield and composition in both positively and negatively high r values multiple regression is not surprising in view of for soil surface variables table 4 are not sur- results from simple regression table 4 and prising moreover these soil surface variables susuggestsests a high degree of intercorrelation with may be expressing the influence of soil physi- soil properties more commonly associated with cal properties that we did not sample but productivity 199819981 sollsoilnegetalSOIL VEGETAL RELATIONS OF recovering SUBALPINE RANGE 359

TABLE 4 contrasting regression relations between yield and composition of grasses and forbs and several independent variables regression coefficient correlation coefficient dependent independent variables variables grass forbs grass forbs yield gmegm2 organic C gmegm2 7176 25158 043 068 r 1 total N gmegm2 0090 0234 030 040 total P gmegm2 0146 0416 027 040 total S gmegm2 0431 0799 025 024 copt 1273 2895 063 073 ntpftpntpo 6627 14116 064 067 rock cover 3356 9188 058 081 litter cover 0051 4123 013 076

Compositioncompositionscomposition1a1 organic C gmegm2 5993 5940 054 054 total N gmegm2 0079 0076 049 049 total P gmegm2 0087 0092 028 029 organic P gmegm2 0435 0449 042 043 copt 0881 0890 0740 74 074 cost 0820 0830 0690ogg69 070 rock cover 2686 2754 079 0800.800 80

significant at P 0050oos05 significant at P 001 basedabased on foliar cover

TABLE 5 statistics from backward multiple regression predicting herbage yield and composition based on foliar cover with 2 vanablevariable equations herbage standardized probability of component variables coefficient significant F fi square

YIELD total regression 0002 072 litter cover 02380.2380 238 0001 bare ground 02160.2160 216 0067 grass regression 0026 052 rock cover 049300.493493 0024 organic P 045400.454454 0085 forbs regression 0001 076 litter cover 04080 408 0001 rock cover 05530 553 0021 composition grass regression 0007 063 litter cover 00940.0940 094 0016 rock cover 07300 730 0015 forbs regression 0005 064 litter cover 00920 092 0013 rock cover 07440.7440 744 0012

we demonstrated strong association between marked differences among the 6 study loca- grass versus forb dominated vegetation and tions based on these soil plant relations several soil nutrient and surface soil properties although vegetation was strongly associated on the whole it appears that forbs responded with P and to a lesser extent S it seems pre- positively to variables generally associated mature to accept the primary hypothesis regard- with better growing conditions and the oppo- ing the importance of P and S the opposite site for grasses moreover results demonstrate relationship of grasses and forbs to depth of A 360 GREAT BASIN naturalist volume 58

0 grassgrossgrans r 0500.50 9 forbs r 0510510.51 lisheddished here all 6 sites were unprotected dur- 100 ing the period prior to 1905 and they appear so to have suffered more or less equally losing 50 90 of the A horizon klemmedson and 60 tiedemannTiedemarm 1994 grazing over the past 80 yr 0 differed among the 6 locations EK and CP 40 0 have been reasonably well protected since the 20 0 8 0 1930s but the impact of differential grazing 0 1905 0 since though marked in the case of EK and CP locations appears to be small com- 50 0 alfra9fr r 0440.44 pared to the differences among sites coupled 0 stleastle r 038 with loss ad4d the earlier of so much of their productive capacity through erosion CF 30 conclusions CL 20 E 0 0 0 10 0 0 although we demonstrated strong relation- 0 ships among several soil and vegetal proper- 0 ties cause and effect relationships were not ogo forthcoming from this information compari- 30 0 cyle r 0600.60 0 asforasfoc r 0560360.36 sons with undisturbed areas would perhaps have provided such linkages nonetheless it

20 is apparent that yield and composition of vegetation were closely tied to soil properties 0 of 0 the physical properties rock cover was the 10 best predictor of yield and composition of grasses and forbs litter cover was the best vegetal attribute for predicting forb yield and 0 4 20 ad4d 6 80 composition of the soil chemical characteris- thickness of01 8 horizon cm tics the CQPcopcopt ratio was the best predictor of forb and grass yield and composition fig 5 effect of thickness ofofabocabA B horizon on composi- the opposite response of grasses and forbs tion by foliar cover of grasses forbs agropyron trachy as groups as indicated by regression coeffi- caucaulumcaulurnlurn stipa lettermaniletlettermanntermani cymopterus lemmonulemmonslemmoniilemmlemmo onnonuniinil and aster cients to all measured attributes suggests that foliafoliaceusfoliaceousceus v canbyi these 2 groups occupy different seraiseral positions in successional development of this area their roles in successional dynamics are not clearly defined and will require more careful study of and B horizon fig 5 supported the alternate their responses to soil development contrast- hypothesis that plant attributes are governed ing responses of individual species within by depth of soil remaining after the period of plant groups figs 4 5 suggest the need to destructive grazing it is also apparent from focus on species level responses in a compan- the differential response of individual species ion study we are attempting to do this by within each group fig 5 that these relation- examining foliar cover response to fertilization ships are not straightforward with 5 combinations of N P K and S for about we believe soil and plant attributes that 100 species over a 5syryr period distinguish the 6 locations were primarily a depth of remaining soil over 90 B hori- reflection of the parent materials and the soil zon has not been emphasized in earlier stud- that remained mainly B horizon after many ies of this area but may be an important deter- years of destructive grazing and severe ero- minant in the composition of grasses and forbs sion differences among soils which in this as plant groups depth of B horizon varied case were chiefly a function of differences in widely among sites and locations and may be a composition of parent material were mani- primary reason that location was such an fested in soil vegetation relationships estabbestab important factor in the analysis 199811998 SOIL VEGETAL RELATIONS OF recovering SUBALPINE RANGE 361

acknowledgments reduced grazing and complete protection unpublished master s thesis brigham young university provo this research was supported by grants UT KEENEY DRD R AND DWD W NELSON 1982 nitrogen inorlnorinor- 2 85 CRSR 2717 and 9138300615691 38300 6156 range ganic forms pages 643 698 in ALA L page editor research grant program USDACSRSUSDA CSRS the methods of soil analysis part 2 and2nd edition agron- authors gratefully acknowledge the shrub omy monographs 9 american society of agronomy improvement and revegetation project inter- and soil science society ofamericaofamerica madison WI mountain research station provo utah for klemmedson JOJ 0 AND ARA R TIEDEMANN 1994 soil and vegetation development in an abandoned sheep cor- providing facilities for this research field assis- ral on degraded subalpine rangeland great basin tance of gary jorgensen range technician naturalist 5430154 301 312 intermountain research station and dr 1998 lithosequenceLithosequence of soils and associated vege- clyde blauer professor snow college both of tation on subalpine range of the wasatch plateau analysis by research note PNW RN 524 USU S department of ephraim utah and laboratory agriculture forest service pacific northwest justine mcneil university of arizona we research station portland OR 16 appp thank dr paul doescher oregon state uni- LAYCOCK WA 1991 stable states and thresholds of range versityversity and dr richard everett pacific north- condition on north american rangelands journal of west research station for reviews of the man- range management 4442744 427 433 LEWONTIN RC 1969 the meaning of stability brookhaven uscript we also thank peer selected lewontinrc reviewers symposia in biology 221322 13 24 by the great rasinbasin naturalist for their helpful MACK RN AND JN THOMPSON 1982 evolution in suggestions steppe with few large holvedhooved mammals american naturalist 119757119 757 773 literature CITED MEEUWIG ROR 0 1960 watersheds A and B a study of surface runoff and erosion in the subalpmesubalpine zone of central utah journal of forestry 5855658 556 560 ANDERSON DWD W 1988 the effect of parent material and NELSON DW AND LE SOMMERS 1982 total carbon soil development on nutrient cycling in temperate carbon and organic matter pages 539 580 in ecosystems biogeochemistry 5715 71 79 organic in ALA L page editor methods of soil analysis part 2 BOWMAN RAR A 1988 A rapid method to determine total 2ndand edition agronomy monographs 9 phosphorus in soils soil science society of america american in society of agronomy and soil society of journal 52130152 1301 1304 science america madison WI BREMNER JMJ M AND CSS MULVANEY 1982 nitrogen 1959 reappraisal of the soil science total pages 595 624 in ALA L page editor methods of nikiforoff CC 129 186 196 soil analysis part 2 2ndand edition agronomy mono- 129186 OCONNOR 1991 perennialerenmal graph 9 american society of agronomy and soil TG local extinction in grass- science society ofamericaof america madison WI lands a life history approach american naturalist 137753137 753 773 CHAPIN FS ililiiIII111 AND GRG R SHAVER 1985 individualistic OLSEN S 1958 of and soil changes on growth response of tundra plant species to environ- JSJ rates succession mental manipulations in the field ecology 66 southern lake michigan sand dunes botanical in gazette 564 576 ilg119125119 125 170 OLSEN S R AND L E SOMMERS 1982 phosphorus COLE CVV AND RDR D HEIL 1981 phosphorus effects on SR LE terrestrial nitrogen cycling ecological bulletin stock- pages 404 430 in ALA L page editor methods of soil holm 3336333 363 374 analysis part 2 2ndand edition agronomy monographs CROFT ARA R 1967 rainstorm debris floods arizona agri- 9 american society of agronomy and soil science cultural experiment station tucson 36 appp society of americaofamerica madison wlWI DICK WA AND MAM A TABATABAITABATABAL 1979 ion chromato- REYNOLDS RXRRVR 1911 grazing and floods a study of graphic determination of sulfate and nitrate in soils conditions in the manti national forestfoiest utah US soil science society of america journal 4389943 899 904 forest service bulletin 91 16 appp DREIMANIS A 1962 quantitative gasometric determina- SAMPSON AWA W 1919 plant succession in relation to range tion of calcite and dolomite by using chittick appa- management USU S department of agriculture bul- ratus journal of sedimentary petrology 3252032 520 529 letin 791 76 appp ELLISON L 1949 establishment of vegetation on depleted SAMPSON AW AND LH WEYL 1918 range preserva- subalpmesubalpine range as influenced by microclimate eco- tion and its relation to eicsionerosionmicsion control on western logical monographs 199719 97 121 grazing lands USU S department of agriculture bul- plateau letin 675 35 1 1954 subalpine vegetation of the wasatch appp utah ecological monographs 9489248924 89 124 SCHREIBER JFJ F JR 1988 final report on the flagstaff HUENNEKE LFL F SPS P HAMBURG R KOIDE HAH A MOONEY limestone paleocene early eocene in the manti AND PM VITOUSEK 1990 effects of soil resources lasal national forest east of manti ephraim san on plant invasion and community structure in cali- pete county utah unpublished report department fornian serpentine grassland ecology 7147871 478 491 ofgeosciencesofgeosciencesGeoceosciences university of arizona tucson JENNY H 1980 the soil resource origin and behavior STANLEY KO AND JWJ W COLLINSON 1979 depositional springer verlag newnewyoikyork 377 appp history of paleocene lower eocene flagstaff lime- JOHNSON HBH 1964 changes in vegetation of two re- stone and coeval rocks central utah american asso- strictstricteded areas of the wasatch plateau as i elatedrelated to ciation of petroleum geologists bulletin 6331163 311 323 362 GREAT BASIN naturalist volume 58

TIEDEMANNTILDEMANN ARA R AND T D ANDERSON 1971 rapid analy- WEBER JN 1964 carbon oxygen isotopic composition of sis of total sulphur in soils and plant materials plant flagstaff carbonate rocks and its bearing on the his- and soil 3519735 197 200 tory of paleocene eocene lake flagstaff of central WALKERWALKLR TW AND AERA FR ADAMS 1958 studies on soil utah geochimica et cosmochimicacosmocbimica acta 28 organic matteimattelmatter 1 influence of phosphophosphorusiuslus content 1219 1242 of palentparent materials on accumulation of carbon nitrogen sulfur and organic phosphorus in grassland soils received 27 october 1997 soil science 8530785 307 318 accepted 17 april 1998 great basin naturalist 584 0 1998 appp 363 374

understory PATTERNS IN CUT WESTERN JUNIPER JUNIPERUS occidentalsoccidentaleoccidentalisOCCIDENTOCCIDENTALSallsALIS SPP occidentalsoccidentaleoccidentalisOCCIDENTOCCIDENTALSallsALIS HOOK WOODLANDS

jon D bateslabatesl2Batesbates1212 richard F millerl3Millermiller1313 and tony Svejcarsvejcarasvejcar44

ABSTRACT western juniper juniperusJumpetusperus occidentoccidentalisoccidentalistalis sppapp occidentoccidentalisoccidentalistalis has rapidly expanded into shrub steppe com- munimunitiesties in the intermountain northwest during the past 120 yr cutting juniper is a management tool used to lestorerestore shrub steppe communities response of the understory after cutting is strongly influenced by plant species composition existing prior to treatment this study assessed distribution patterns of understory plants over 2 growing seasons aftelafterarrelanterarrer tree cutting in a western juniper woodland cover density and diversity of understory species were compared among 3 locations interinterspacesspaces duff zones previously under tree canopiescanopies and debris zones beneath cut trees plant cover and density increased in all zones following tree cutting understory vegetation in cut woodlands exhibited strong zonal dis- tributiontribution cover and density of poa sandbergii and Sitsitanionsitamonsidamonanion hystrix and canopy cover of annual forbs were greatest in duff zones FP 005 density and cover of other perennial grasses and total densities of perennial forbs and annual forbs were greatest in interinterspacesspaces P 0050.050 05 debris zones tended to have the lowest overall understory cover and plant density values under juniper debris many species common to interinterspacesspaces were reduced in density although plants that survived or established beneath debris grew larger than their counterparts in interinterspacesspaces species that increasedinciincl eased in density and covelcover under debris were plants characteristic of duff zones and whose seeds are typically wind dispersed

key words western juniper understory patterns diversity juniper debris species composition zonal succession

pinyon juniper woodlands in the western et al 1987 bates 1996 intercepting precipi- united states have rapidly expanded into tation larsen 1993 and causing physical or shrub grasslands since the late 1800s tausch allelopathicallelopathic interference by litter layers jame- et al 1981 west 1984 miller and wigand son 1966 peterson and buss 1974 padien 1994 western juniper juniperus occidentoccidentalisoccidentalistalis and laethalajtha 1992 attributed understory spatial sppapp occidentoccidentalisoccidentalistalis hook has invaded extensive patterns in pinyon juniper woodlands to dif- areas of sagebrush grasslands and other plant ferencesferences in nutrient availability shade protec- communities in the pacific northwest burk- tion seed dispersal and seed germination hardt and tisdale 1969 miller and rose 1995 management prescriptions employed to re- the transition from shrub steppe communities duce western juniper dominance in rangelands to woodlands has resulted in reduced under- have been successful in increasing understory story productivity and diversity johnson 1962 productivity and cover evans and young 1984 jameson 1967 burkhardt and tisdale 1976 vaitkus and eddleman 1987 rose and eddle- tausch et al 1981 tausch and tueller 1990 man 1994 bates 1996 however the influence bates 1996 understory distribution patterns of spatial distribution on plant succession in canopy and interspace zones become more following juniper elimination particularly distinctly developed during woodland develop- with removal methods that leave substantial ment johnson 1962 pieper 1990 vaitkus and amounts of juniper debris on site are poorly eddleman 1991 understory patterns proba- documented bly reflect a mosaic of canopy and interspace evaluating understory distribution patterns micromicroenvironmentsenvironments Junijunipersjuniperuspers influence the may prove useful in generating hypotheses on microenvironmentmicroenvironment under tree canopiescanopies by mod- species zonal interactions and in predicting ifying temperatures and light levels pieper successional responses and pathways follow- 1990 accumulating soil nutrients doescher ing juniper control this study was designed

lrangelandangeland resources department oregon state university eastern olegonoregon agricultural research centelcenter bumsburns OR 97720 2plesent2present address eastern olegonoregon agricultural researchRe seaichsealch center olegonoregon state university hc71 hwybwy 205 bumsburns OR 97720 the eastern olegonoregon agricultural researchRe seaich center isis operated jointly by the USDAARSUSDA ARS and the oregon agricultural experimentexpelexper imentament station of oregon state university aliallthoiAllhauthor3authorthoithor to whom coicol respondencecorrespondence should be addressed 4usdaUSDA agricultural researchReseaichsearch serviceselvideSe ivice hwybwy 205 bumsburnsbugs OR 97720

363 364 GREAT BASIN naturalist volume 58

1 to assess understory distribution patterns cally cool and wet summers are warm and dry within an intact western juniper woodland prior the majority of annual precipitation falls be- to tree cutting and 2 to evaluate the effects of tween november and june mean water year tree cutting on understory zonal patterns 1 I1 october 30 september precipitation at weather stations located 27 km southwest ele- METHODS vation 1300 m and 30 km northwest 1250 m of the site average 28228.2 and 24924.9 cm respec- study site tively study was conducted on steens moun- the experimental design and tain in southeastern oregon 9.595gs km southeast 95 measurements of diamond IISWW 42wn4255n elevation at the site is 1525 m and aspect is west facing in june 1991 we established eight 08osha ha with a 22 slope the site is dominated by an replicated plots along the contour of the ridge 80 yr old western juniper woodland juniper slope plots were selected for similarities in canopy cover averaged 23 and tree density overstoryunderstoryoverstory understory cover and density soil averaged 228 trees haglhagiha 1 the following indi- type and aspect after being measured for cated a fully developed juniper stand limited baseline vegetation characteristics basal cover terminal and lateral leader growth on juniper and density juniper trees were felled with trees lack of juniper seedling recruitment chainchainsawssaws in august 1991 on half of each plot and most artemisia dentatatritridentatatridentate sppapp vastyanavaseyanavaseyana 04040.4 ha cut trees remained on the plots we nutt mountain big sagebrush were dead began subsequent measurements of under- understory perennial plant basal cover aver- story characteristics and soil moisture content aged 252.5 in interinterspacesspaces and 292.9 under tree in april 1992 and concluded in september canocanopiespies based on existing shrubunderstoryshrub understory 1993 in this paper we report data from the vegetation soils and aspect we judged the cut plots only in order to highlight zonal dif- original community prior to woodland domi- ferencesferences nance to have been artemisia dentatatritridentatatridentate sppapp understory measurements were canopy and vaseyanastipavaseyanalstipa thurberianathurberiana type basal cover density and diversity sampling prior to treatment the dominant understory was spatially separated into 3 zones duff plant was poa sandbergsandbergiiii vasey sandberg s juniper debris and interspace duff zones are bluegrass comprising nearly 75 of the total defined as those areas formerly beneath tree understory perennial plant basal cover other canopiescanopies with a surface layer of old juniper species characteristic of the site included stipa needle litter debris zones are former inter- thurberianathurberiana thurber s needlegrassneedlegrass sitanionSitanion space zones that are covered by felled juniper hystrix bottlebottlebrushbrush squirreltail agropyron trees interspace zones are open areas that are spicatumspicatum bluebunchbluebunch wheatwheatgrassgrass astragalus not influenced by old or newly felled juniper filipes basalt milkvetchmilkvetch microsterisMicrosteris gracilis tree litter microsterismicrosteris and alyssum alyssioides pale we estimated understory plant density alyssum 1991 1993 and canopy cover 1993 only for soils were classified as clayey skeletal each zone in each replicate in the 4 cardinal montmorillonite frigid lithic agrixerolls they directions for duff and interspace zones only are shallow 40 50 cm deep and are underlain around 12 trees per replication using a 30530.5 X by a thick welded ash layer of rhyolite and 61glemgiemcm frame 48 subsamples per zone per rhyodacite composition which limits root replication trees were randomly selected penetration each year for the debris zone we estimated domestic livestock grazing has occurred on density and cover by randomly samplingsubsamplingsub 4 this site since the late 1800s the ridge on locations under each of the 12 cut trees in which the study site is located was used as a each replication we subsamplessubsampledsubsampled along the sheep wintering area through the 1930s since outer 13 of the duff zone interspace zones the 1940s the site has been grazed by cattle in were located approximately 3 m from the the early spring april livestock were outer edge of the duff zone or at the midpoint excluded from the site during our study between duff zones climate in southeastern oregon is semiarid using the cover class technique described by and continental winter and spring are typl daubenmire 1959 we estimated canopy cover 199811998 understory PATTERNS 365 in this study 7 cover classes were designated abundant species hill s modified evenness trace 0 1 I1 1 5 II11 5 25 III111 25 50 ratio was used to compare relative abundances IV 50 75 V 75 95 and VI 95 100 of species among zones ludwig and reynolds midpoints of cover classes were used for sta- 1988 tististicaltical analysis understory basal cover of perennial grasses RESULTS and forbs was measured along five 305 m line climate conditions and soil water intercepts positioned parallel to the slope in all 8 cut plots in 1991 baseline year prior to water year 30 september 1 october pre- cutting 1992 and 1993 transects were per- cipitation values were 20 below average in manentmanentlyly marked in 1991 using rebar stakes 1991 and 1992 at weather stations 27 and 30 Groundgroundcovercover provided by juniper debris and km from the site study site precipitation in old juniper litter in duff locations was also the 1992 water year totaled 21321.321913 3 cm half of estimated along the transects which was received in june and july 1992 the gravimetric soil water content was sampled 1993 growing season was cooler and eastern in interspace and debris zones at 2 depths oregon received record amounts of moisture 0 20 and 20 40 cm we collected biweekly precipitation totals at nearby weather stations samples on 12 dates during the 1992 growing were 140 and 149 of longtermlong term averages season april september and on 13 dates respectively water year precipitation on the during the 1993 growing season in each plot study site totaled 41841.841 8 cm 5 randomly located subsamples were collected in 1992 soil moisture at both depths was for each zonal depth during each measure- significantly greater P 0050.05oos0 05 in debris zones ment period soils were weighed oven dried than in interinterspacesspaces from late june through at 106c for 48 h and reweighed to determine september fig 1 in 1993 soil water at 0 20 percent water content cm depth was significantly greater under debris than in the interspace fig 1aaa except statistical analysis in for 2 periods mid may and july there were understory data were compared among no differences in soil water content at 20 40 zones over time using ANOVA techniques for cm between the 2 zones in 1993 fig ab1b a randomized block design main effects were there were significant time by zone by year and zone understory measurements depth interactions for soil water in 1992 and were also analyzed each year to help explain 1993 P 0050.050oos05 these interactions indicated year by zone interactions subsamples of under- that soil water decreased as the growing sea- story density and canopy cover were averaged son progressed and was greater at both depths by zone per replicate for statistical analysis in the debris zone than in the interspace n 8 for each year soil water content was depth by zone P 0050.05oos0 05 interactions indi- analyzed each year using a repeated measures cate that soil water was greater at 20 40 cm ANOVA for a randomized block design main than 0 20 cm in the soil profile effects were zone and soil depth understory density and cover all statistical analyses were performed using the statistical analysis system SAS institute pretreatment vegetation PATTERNS 1988 data were tested for normality using perennial plant basal cover in interspace and the SAS univariate procedure data not nor- duff locations was very low prior to the cutting mally distributed were log transformed to sta- treatment fig 2aaa total plant basal cover did bilize variance when interactions were signif- not differ between zones however for several icant means were separated using Dunduncanscarsears species we detected significant differences in new multiple range test the alpha level was cover between zones basal cover of poa sand set at P 0050.05oos for statistical significance bergli and Sitsitanionsitamonsidamonanionanlon hystrix was greater in duff diversity indices were determined for each zones than in interinterspacesspaces P 0050.05oos0 05 fig 2aaa zone using density measurements in 1992 and basal cover of stipa thurbenanathurberianathurthurberbenanaiana was highest in 1993 hill s 1973 NI and n2na diversity indices interinterspacesspaces P 0050.050 05 densities of poa and were used as indicators of plant diversity the Sitsitanionsitamonsidamonanionanlon were greater in the duff zone than in n2na index is a measure of very abundant interinterspacesspaces P 0050.050 05 fig 3aaa density of stipa species and the n1na index is a measure of agropyron spicaspicatumtum and alyssum allyssioides 366 GREAT BASIN naturalist volume 58

35 A 0 20 cm 0 1 interspacenterspace debris 30

25 0 ayyy 0 20

35 B 20 40 cm

30 all

an 0 25

U0 20 interspace debris

15 1 A M J J A S 0 A M J J A S 1992 1993 during fig 1 volumetric soil water content in interspace and debris zones from A 0 20 cm and B 20 40 cm depths the 1992 and 1993 growinggi owing seasons data are in means s asterisks denote significant differences between zones P 0050 05

was greater in interinterspacesspaces than in the duff posttreatmentPOSTTREATMENT vegetation PATTERNS zone FP 005oos0.05 figs 3aaa 4aaa density values cover and density of understory species dif- for annual forbs in 1991 are probably low and fered significantly among debris interspace incomplete because sampling did not take and duff zones P 0050.05oos after cutting figs place until july well past the peak for annual 2 4 cover of perennial plant species and forb growth density of most other understory species 199811998 understory PATTERNS 367

uu505.0 454.5 A 1991 Preprecuttingcutting I1 I1 interspace 404.0 duff & 353.5 b 0 303.0 a b 0 252.5 5 a i g 202 0 1 0 151.5 CO a 13L 101.0loio b da 050.5 i1 F 1 F 1 m r B 11 FII FA oo 1 1 00 1 1 1 I1 I1I I 000.0 niI I1 I I posa stthotth pssp sihybihy PGT alfiasfi PFT total 505.06jj0 11111111111111 454.5 BR i 9921992 debris 404.0 1 1 interspace 0 o 353.5 duff 303.0 D a b 00 z250 ab ia5 202.0 b W fl ca0 1.54 C a m 15 a b a 101.0iolo a iiilii b a a 050.5 1 rffll 1 nj MOM B Jra M mltmfmtm mromsomM oo li 00 1 000.0 i i posa suth pssp sihybihy PGT alfiasfi PFT total

505.0 i ah f 003002qq3 lllllllllllltfj1h fohricbohric15 454.5 C 1993jwjlwj debrisUGUI r 404.0 1 1 interspace 1 H &S 353.5 duff 303.0 0 252.5 08 c EM iai5co 202.0 b g i a 1 151.5 EM m h a a 1 IN i- l m a HM 101.0loio c bb HA fllBall a a ab hah1 050.5 H r b UM 1 000.0noo n posa stthotth pssp sihybihy PGT alfiasfi PFT total

fig 2 understory basal cover in A 1991 precuttingprecutting data B 1992 and C 1993 for the most common perennial plants different letters denote significant zonal differences P 0050.050oos05 between species or plant groups species abbre- perennial viationsviations posa poa sandbergsandbergiisandbergnsandberghii suthsurh stipa thurberthurberianathurbenanathurbehanabenanaianalana agspagap agropyron spicaspicatumtum smy Sitsitanionsttomonstromonanionanlon hihystrixstrix PGT perennuinul grass total brtearte bromus tectoriumtectorumtectorum alfiasfi astragalusfilipesastragalus filipes PFT perenperennialmalmai forb total 368 GREAT BASIN naturalist volume 58

16 A 19911991precuttingPreprecuttingpnecutting b interspace 14 a CP 12 duff E 10 2 8 ca faf1 C 6 aU 3 b C a b a iroicoM b FLa 2 a 1 0 stthotth agspagap sihybihy PGT posa batebade alfiasfi PFT

16 b 13 debris 14 1992 CP 12 a a inter E b 10 duttduff 8 6 b a a a 3 a a a b b C b b bb a M a a b FL 2 a a aa a

0 suth agspagap sihybihy PGT posa batebade alfiasfi PFT

16 b b Cc19931993 debris 14 inter C 12 E a 10 a duff Z 1018 b b 7U.7 a 0a 00 a b 4 a b CE b b b jcaaca 3 b a a a b a a b a 0 2 a

0 stthotth agspagap sihybihy PGT posa batebade alfiasfi PFT

fig 3 understory density in A 1991 precuttingpre cutting data B 1992 and C 1993 for the most common perennial plants and annual grasses different letters denote significant zonal differences P 0050.050 05 between species or plant groups species and plant group abbreviations are defined in figure 2 199819981 understory PATTERNS 369

100 1 I1 A 1991 Preprecuttingcutting I interspace 8u80au CM duff E 60 a

28 a 1 bD 5 C 24 u 20 C 16 W0 12 0Q 8 b 4 n vmF 1 0 R I1 I1 I1 I1 I1 I1 alaalai cepidepi eppa lase figrmigr rute OAF AFT

100 B 1992 bi EBIH debris 1 I1 inter 80 E 60 duttduffpuff 28 duff a 0 C 24 1 20 a 1 16 1 C a ai AS 12 b 0CL b b b b 8 a 1 1 aaaa araana a a 1 4 I 1 10 m L rb 1 0 1 1 1 J J 1I1 T T i alai cepidepi eppa lase figrmigr rute OAF AFT b 100 C 1993 1 debris inter ft bhi 80 hb N PMMM D ff 1 1 u I I duff a 1 E bu60 n wj S 28 c c mm I11 H w w m i C 24 a0 20 C I1 1 b C 16 a b b t0taW a b a 12 b 1 8 b jida aaa a 1 a i 4 a j M m I1 a TO 0n J m alai cepidepi eppa lase figrmigr rute OAF AFT

fig 4 annual forb density inm A 1991 preprecuttingpiecuttmgcutting data B 1992 and C 1993 different letters denote significant zonal differences FP 0050.050 05 between species and plant groups species abbreviations malmaialai alyssum alyssioides cepidepi descurainiadescuraima pinpinnatepinnatanata eppa epilobium paniculatumpamculatum lase lactuca sesefsernoianolanolorriola figrmigr microstensmicrosterisMicrosterisstens gracilis rute ranunculus testiculatus OAF other annual forbs ege g cirsium sppapp ciliagilia sppapp AFT annual forb total 370 GREAT BASIN naturalist volume 58

32 1 1 EM debrisde bnsans F 1 interspace EM duff 30 a a 28 r a T 18 ft J 8 2 16 1 1

u 1 1 14 S 00 12 a abyb CL a a b 7 a 0c 1 0 10 1 0 b 1 1 8 a a Cc b r 1 6 a b c a a C 4 ana a i a 1 b b b a 0 b 1 1 n8 I b a a a a b a 2 b a J ri ab 1 B 1 a H B mm B ji d 1 11 M70 i 0 ik i i Ai stthotth agspagap sihybihy PGT posa alfiasfi croc PFT brtearte aldeaide figrmigr lase cepidepi AF AFT total

fig 5 Postposttreatmentestmenteatmenttreatment canopy cover in 1993 for interspace duff and debris zones different letters denote significant zonal differencesdiffeidiffee encesances P 0050.05oos0 05 among species and plant groups new species abbreviation croc crepis occidentoccidentalisoccidentalistalis otherothel species and plant group abbreviations are defined in figures 2 and 4

increasedinci eased significantly between 1991 and 1993 debris zones the change in total perennial in all zones P ooi0010.010 01 species that increased grass cover under debris was a result of inin density P 0050.050 05 included sitanionsttamonsteamonStSitanionanlontamon agropy- creased cover of sitanionSitanion ron and most perennial and annual forbs ex- Groundgroundcovercover also increased by the addition cept alyssum of jumperjuniper debris Groundgroundcovercover provided by basal cover of sitanionsttamonsteamonStSitanionanlontamon astragalus lipesfilipesft juniper debris averaged 18 in cut plots lomatium donnelli donnell s lomatiumlomatmm and together with cover provided by interspace perennial forbs as a group exhibited signifi- vegetation 18 see fig 5 and by old juniperjumper cant year by zone interactions P 0050.050oos05 due litter in duff zones 22 total groundcoverground cover to an increase in basal cover in duff and debris after cutting was about 58 in 1993 we zones between 1991 and 1993 for most species observed that juniper debris was effective in there was a lack of year by zone interaction trapping sediment washed downslope from indicating that plant cover and density domi adjacent uncut woodlands after heavy rain nance patterns remained consistent among storms in july 1992 zones between 1991 and 1993 basal cover PLANT DIVERSITYDIVE RSITY diversity and species figs 2bcnbc density figs 3bcnbc and 4bcnbc evenness tended to be greater in the duff zone and canopy cover fig 5 of poa sitanionsitamonsidamonSiSitaniontamon and in 1991 and 1992 than in the interspace or annual forbs remained greatest in duff com- debris 1992 only zones table 1 in 1993 pared to other zones P 0050oos05 density and diversity and evenness indices in both duff cover of other perennial grasses primarily and debris zones were significantly greater than agropyron and stipa and density of perennial in the interspace P 0050.05oos higher diversity forbs and alyssum were greater inm interinterspacesspaces and evenness ratios in duff and debris zones than in either duff or debris zones P 0050.050 05 compared to the interspace were due to higher canopy covelcover and density of sifsitanionSitsztamonanionanlon and species richness of annual forbs and lower canopy cover of annual forbs tended to be densities of alyssum and microsterisMicro steris gracilis greater in debris zones than in interinterspacesspaces P the high densities of alyssum reduced even- 00500.05oos05 between 1992 and 1993 basal cover ness and diversity in interspace zones plant of perennial grasses increased fourfold in diversity increased between 1991 and 1993 in 199819981 understory PATTERNS 371

TABLE 1 plant diversity hill s NI and n2na and evenness on warm sunny days soil temperatures 5 ratios in interspace duff and debris zones evenness cm depth under tree canopiescanopies were 4 ac dominance of several 7c ratios in this table show a co species lower than in spaces during the growing with an array of less abundant species in interspacesinter season 1993 data not shown buffering of yearlocationYear Location hill s ni hill s n2na evenness temperature extremes beneath tree canopiescanopies 1911 may favor plants that initiate growth relatively interspace 313.13 1 alag 2424aaaa 05aa05 aa early in the spring such as poa and sitanionsitamonsidamonSitanionanlon 26 aa 05os aa duff 3636aaaa 26aa262.6 050.505aa cooler temperatures under tree canopiescanopies re- haworth and mcpher- 1992 duce evapotranspiration interspace 51ab51 ab 353.535abab 060.6og06aaaa son 1995 and therefore may decrease under duff 707.0tobbbb 49bb49 bb 06aab060.6og aab story water stress debrishdebris3 52ab525.2 ab 36aa36 aa 06aab060.6og aab we did not measure photosynthetic active PAR this study were 1993 radiation in but there interspace 595.9sg59abab 4040abab 0505aaaa indications that light levels for plant growth duff 858.58 5 bc 6266.22 bc 07bb07 bb were reduced under canopiescanopies in duff zones in debris 878 7 bc 63bb63 bb 07bb070.7 bb 1991 in 1991 we observed that perennial difieientidifferentindifferent lowercase letters indicate significant zonal differences inin a yeaiyealyear grasses tended to exhibit etiolatedetiolapetiolatedted growth P 005oos0 05 under tree cano diffrientdiffeient uppercase letters denote signiticintsignificant yeaiyeatyear differences within a zone patterns in duff zones canopiespies P 005oos0 05 several studies have reported that as light lev- debris values inin 1992 and 1993 are compared to 1991 interspace value because beneath canopy with before cutting in 1991 debris zones were interspaceintel space aieasadeasareas els decrease the tree pinyon juniperjumper woodland development perennial grass cover decreases schott and pieper 1985 pieper 1990 therefore lower light lev- all 3 zones P 0050.05 species evenness in- els in duff zones than in interinterspacesspaces may be creased between 1991 and 1993 in debris and responsible for lower cover and density of duff zones perennial grasses the combination of reduced there were similarities and differences in light levels and lower temperatures also reduces terms of the most abundant species in each seed germination of annual plants baskin and annual zone abundant species in the duff and debris baskin 1985 which may explain why zones zones in 1993 were alyssum microsterisMicrosteris des plant densities were lower in duff ver- curainia pinnatapinnate pinnate tansytansymustardmustard lac- sus the interspace tuca serriola prickly lettuce bromus tectoriumtectorumtectorum Postposttreatmenttreatment vegetation patterns cheatcheatgrassgrass astragalus poa and sitanionSitanion abundant species in the interspace were agro- there were significant increases in under- pyron stipa poa alyssum and microsterisMicrosteris story cover density and diversity after juniperjumper cutting these increases occurred mainly dur- discussion ing the 1993 growing season which was characterized by more favorable growing condi- patterns pretreatment vegetation tions higher spring moisture plant response differences in understory composition be- to the cutting was limited in 1992 due to dry tween duff and interspace zones indicate that conditions occurring from late winter through juniper influences the development of zonal the spring however the ability of the under- micromicrositessites conditions inm duff zones were story in the cut treatment to respond in the more favorable for poa and sitanionSitanion than asso- high precipitation year was made possible by ciated species whereas conditions in the eliminating juniper competition for soil moismolsmois- interspace favored stipa and alyssum figs ture and N bates 1996 we believe the under- 2 5 these plant zonal patterns do not appear story response in the cut treatment resulted to be related to soil moisture content as there from elimination ofjuniperofjuniper competition because were no differences detected in soil moisture in adjacent uncut woodland the understory availability between duff and interspace zones showed little response to increased precipita- bates 1996 duff and interspace understory tion in 1993 bates 1996 patterns may be related to other factors influ- zonal understory plant composition did not encing plant establishment and growth such change in duff or interspace zones after cut- as temperature light and nutrient availability ting species with greater cover figs 2 5 or 372 GREAT BASIN naturalist volume 58 density figs 3 4 in duff zones such as poa seedling establishment by physically impeding and sitanionSitanion remained dominant in the duff seed soil contact reducing soil temperatures zone while stipa agropyron and alyssum and restricting plant growth via alleallelopathylopathy remained dominant in the interinterspacesspaces these jameson 1966 peterson and buss 1974 results suggest that early postcuttingpostcutting under- although allelopathicallelopathic effects should not be story composition particularly for zonal domi discounted they seem unlikely given growth nants is predictable based on pretreatment patterns of plants established in duff areas understory floristics predicting understory plants that were established or became estab- dynamics after cutting however provides only lished in the duff zone tended to grow larger a qualitative estimate predicting a quantita- than their counterparts in the interspace par- tive response is more difficult due to a num- ticulticularlyarly annual grasses and forbs ber of uncertainties particularly postcuttingpostcutting there was a propensity for greater estab- weather conditions uncertainty is also intro- lishmentlishment of sitanionSitanion and several annual forbs duced by lack of knowledge about the quan- lactuca cirsium sppapp thistlthistleswhistleses in duff zones tity and composition of soil seed bank reserves than in interinterspacesspaces this may be a product of koniak and everett 1982 and the level of seed dispersal and catchment mechanisms understory seed production following release seeds of sitanionSitanion and these annual forb species from juniper competition for instance the are typically wind dispersed we hypothesized increase in plant diversity and species rich- that old juniper needle litter in duff zones is ness table 1 in our study appears to have more effective in trapping wind dispersed seeds largely resulted from the emergence of plants than the relatively bare soil surfaces character- from soil seed banks and belowgroundbelowground bulbs izing the interinterspacesspaces and tubers understory succession following juniper debris had negative and positive pinyon juniper removal by fire is also guided effects on understory plants prior to cutting by initial site floristics although prediction of in 1991 debris zones were interinterspacesspaces after postposttreatmenttreatment response is again limited to trees were cut and debris zones created com- qualitative estimates everett and ward 1984 position of the understory shifted developing after cutting plant cover increased more greater similarity to duff zones than to inter- than did density especially for perennial plants space zones in both years following juniper because existing perennial plants grew larger cutting cover and density of sitanionSitanion and the in size between 1991 and 1993 total perennial following wind disseminated annual forbs grass basal cover increased by nearly 200 increased under juniper debris lactuca epi- but perennial grass densities increased by only lobium paniculatum willowweedwillow weed and cir- 65 in duff zones and 43 in interinterspacesspaces the sium sppapp these plants tended to establish lower density response of perennial grasses along the outer edges and less heavily shaded was due to the lack of seed production except or open patches of the debris zone these re- for sitanionSitanion little seed production in the sults indicate that environmentalmicromicroenvironmental changes perennial component was observed in 1991 or can alter species composition and successional in 1992 in 1993 we observed that perennial pathways grasses allocated a large portion of their the increase in sitanionSitanion density under growth to reproduction the higher perennial debris suggests that debris also may be benefi- seed crop in 1993 may alter plant composition cial for establishment of other grass seedlings on the site in subsequent years once needles fall from the debris and suffi- litter layers in duff zones continued to in- cient perennial grass seed sources are avail- fluence species establishment and growth fol- able juniper debris may serve as important lowing cutting plant density and cover were micrositesmicrosites for other perennial grass seedlings greatest in the outer portion of the duff zone total annual forb density and cover and decreasing with proximity to the tree stump density of perennial grasses except sitanionSitanion where litter depth was greatest bates 1996 and boaboopoapoo and perennial forbs were still signifi- in other pinyon juniper woodlands plant den- cantly lower under debris than in interspacesinterspaces sity and cover decreased as the tree bole was P 005oos0.05 in 1992 and 1993 negative effects approached and litter layers thickened everett of juniper debris on perennial grasses except and sharrow 1985 dye et al 1995 juniper sitanionSitanion and rodroapodpoa and forbs were particularly litter may interfere with seed germination and evident in 1992 when compared to interspace 199819981 understory PATTERNS 373 values in 1991 perennial grasses such as stipa not be radically altered by juniper debris since and forbs eg astragalus were killed under debris coverage averages only 18 across the heavy debris accumulations as evidenced by site their reduced densities the negative impact of juniper debris on acknowledgments annual forbs particularly alyssum and micros teristefistensteffs may have resulted from lowered seed we thank fred otley and family for use of germination andor plant establishment dimin- their property during the study thanks to ished light levels and lower soil temperatures kara paintner and jeff rose for statistical help due to shading by juniper debris may reduce and assistance in the field thanks are due seed germination and establishment of annual to lee eddleman dave ganskopp wendy forbs thus resulting in decreased forb density maichlerwaichlerWa ichler and 3 anonymous reviewers for their however annuals and perennials that did estab- comments and suggestions this manuscript paper lish in debris were generally larger and stayed was submitted as technical 11161 for active longer into the season than their coun- the oregon agricultural experiment station terpartsterparts in the interinterspacesspaces bates 1996 the larger individual sizes of plants and literature CITED their prolonged growing season under debris BASKIN JMJ M AND CCC C BASKIN 1985 the annual dormancy may have been a product of improved plant cycle in buried weed seeds a continuum bioscience water relations higher soil moisture levels 3549235 492 498 fig 1 under debris are hypothesized to have BATES JDJ D 1996 understory vegetation response and cycling following cutting of western resulted from a combination of reduced evap- nitrogen juniper unpublished dissertation oregon state university orative loss and lower moisture demand by Corvalcorvallishs 230 appp plants reduced temperatures and increased BURKHARDT JW AND EW TISDALE 1969 nature and boundary layers provided by pinyon juniper successional status of western juniper vegetation in debris lower vapor pressure gradients thereby idaho journal of range management 2226422 264 270 gifford 1976 causes of juniper invasion in southwest reducing transpirationtranspirationalal demand and idaho ecology 5747257 472 484 shaw 1973 the lower density of plants under daubenmire RER F 1959 A canopy coverage method of debris also may have resulted in more avail- vegetational analysis northwest science 334333 43 64 able resources per plant than in the inter- DOESCHER PS LEL E EDDLEMAN AND MSM S VAITKUS 1987 evaluation of soil nutrients ph and organic space thus contributing to larger plant size matter in rangelands dominated by western juniper results from this study support findings northwest science 619761 97 102 from other pinyon juniper systems that under- DYE KL DN UECKERT AND SG WHISENANT 1995 story plant composition is influenced by zonal redberry juniper herbaceous understory interac- of range management 4810048 loo100 107 location has been little discussion abouabout tions journal there EVANS RAR A AND JAJ A YOUNG 1984 plant succession in the effects on understory zonal patterns of following control of western juniper juniperusJumperus occi juniper removal by cutting fire or other means dendentacentacentedentalsdentalisdentahstalistailshs with pipicloramcloram weed science 336333 63 68 our study determined that while there were EVERETT RLR L AND SHS H SHARROW 1985 understory re- smgleleafsingleleaf and significant increases in plant cover and density sponse to tree harvesting of singlesingieleaf pinyon utah juniper northwest science 4510545 105 112 removal was little resulting from juniper there EVERETT RLR L AND KOK 0 WARD 1984 early plant succes- change in relative species understory zonal sionslon in pinyon juniper controlled burns northwest dominates composition in duff or interspace science 585758 57 68 zones predicting qualitative species composi- GIFFORD GFG F AND CBC B SHAW 1973 soil moisture patterns on two chained pinyon lumperjumperjuniper sites in utah journal tion in duff and interspace zones at least in of range management 2643626 436 440 the first 2 yr postcuttingpostcutting is possible from pre- HAWORTH K AND GRG R mcpherson 1995 effects of treatment community floristics understory re- quercus eonyiiconyneonyn trees on precipitation distribution and sponse to the deposition of juniper debris is microclimate in a semisemiaridarid savanna journal of andaridarld environments 3115331 153 170 less predictable changes to the microenviron HILL MOM 0 1973 diversity and evenness a unifying nota- ment caused by juniper debris rapidly shifted tion and its consequences ecology 5442754 427 432 understory plant composition how juniper JAMESON DAD A 1966 pinyon juniperjumper litter reduces growth debris affects vegetation dynamics over a longer of blue grama journal of range management 19 214 217 period of is currently being monitored on time 1967 the relationship of tree overstory and herba- this site we hypothesize that overall plant ceous understory vegetation journal of range man- community composition and development will agement 2024720 247 249 374 GREAT BASIN naturalist volume 58 jolJOIJOHNSONINSON FNTN 1962 one seeded juniper invasion of north- SASAS INSTITUTE 1988 user s guide release 603 edition ern arizonaanzona grasslands ecological monographs 32 SAS institute carygary NC 187 207 SCHOTTCHOTT MRM R AND RDR D PIEPER 1985 influence of canopy KONIAK S AND RLR L evereheverenEVERLHEVERETT 1982 seed servesreservesle in soils characteristics of one seed juniper on understory of successional stages in pinyon juniperjumper woodlands grasses journal of range management 3832838 328 333311 americanamerleanamencan midland naturalist ioslos108295108 295 303 tauschAUSCH RJR J AND PT TUELLER 1990 foliage biomass LARSENLARSLN RER E 1993 interceptionintel ceptioncaption and water holding capac- and cover relationships between tree and shrub ity of westeinwestern juniper unpublished dissertation dominated communities in pinyon juniper wood- oregon state university corvallisCoigoigol vallis lands great basin naturalist 5012150 121 134 ludwicLUDWIG JAJ A AND JEJ F REYNOLDS 1988 statistical ecology tauschAUSCH RJR J NEN E WEST AND AAA A NABI 1981 tree age john wiley & sons new york and dominance patterns in great basin pinyon MILLERmilmii lekLLK RER F AND JRJ R ROSEROSL 1995 western juniper expan- juniper woodlands journal of range management sionslon in eastern oiegonbiegonoregon great basin naturalist 55 3425934 259 264 37 45 VAITKUSAITKUS MSM S AND LEL E EDDLEMAN 1987 composition millenMILLERMILLLR RER F AND PE WIGAND 1994 holocene changes and productivity of a western juniper understory in semiarid pinsonpinyon juniper woodlands responses to and its response to canopy removal pages 456 460 climate firefnan e and human activities in the USU S great in RLR L everett editor proceedings of the pinyon basin bioscience 4446544 465 474 juniperjumper conference general technical report INT PADIENPADILN DJD J AND K LATHALAJTHA 1992 plant spatial pattern 215 USDA forest service intermountain forest and nutrient distribution in pinyon junipeijuniper wood- and range research station ogden UT lands along an elevational gradientgi adient in northern new 1991 tree size and understory phytomassphy tomass pro- mexico intelinternationalnational journal of plant science 153 duction in a western juniper woodland great basin 425 433 naturalist 5123651 236 243 PIEPERpli PLRper RD 1990 overstory understory lelationsrelations in WESTEST NEN E 1984 successional patterns and productivity pinyon juniperuniper woodlands in new mexico journal of pinyon juniper ecosystems pages 1301 1332 in of range management 4341343 413 415 developing strategies for range management west PLILRSONPETERSON GB AND WR BUSS 1974 determination of view press boulder CO the plesencepiepresencesence location and allelopathicallelopathic effects of sub- stances produced by Jumjuniperusjumperusperus scopulorum proceed- received 11 june 1997 ings of the south dakota academy of science 53 accepted 21 january 1998 298 rosirosfROSE JR AND LE EDDLEMANEDDLCMAN 1994 Pondepondeiosaponderosaiosalosa pine and understory glowthgrowth following westeinwestern juniper removali northwest science 687968 79 85 great basin naturalist 584 0 1998 appp 375 379

comparative demography OF THE HIGH ALTITUDE LIZARD sceloporus GRAMMICUS phrynosomatidae ON THE iztaccihuatl VOLCANO PUEBLA MMEXICOXICO

julio A lemos espinallespinellespinal1 royce E Ballinger 2 and geoffrey R smithasmith3srnith3

ABSTRACT population density reproduction and survivorship were compared between 2 populations of sceloporus grammicusgrammicus occurring at different altitudes 3700 m and 4400 m on the eastern slopes of iztaccihuatl volcano puebla mexico lizards in both populations matured at the same age 14 15 mon and size 39 42 mm SLV population density was slightly greater at high altitude 131 163 per ha than at low altitude 52 83 per ha survivorship and rorp were higher at the low altitude area but in general there were no significant demographic variations between altitudes that have been reported in lizard population at higher latitudes studies of lower elevation populations might reveal some differences because previous studies indicate that litter size increases at lower altitudes although they do not differ between our 3700 m and 4400 m populations

key words lizard life history demography reproduction altitude variation sceloporus grammicusgram micus population density replacement rate survivorship

life histories and demographic traits of southern texas USA to the state of oaxaca lizards can vary along elevational gradients mexico conant and collins 1991 flores vil- ballinger 1979 grant and dunham 1990 lela and gerez 1994 this species has been smith and ballinger 1994a 1994b altitudinal poorly studied and little has been published variations in life history characteristics often on its biology except for studies on its repro- mimic variation observed across broader geo- duction guillette and casas andreu 1980 graphic ranges that can be attributed to differ- 1981 ortega and barbaultBarbarbauldbault 1984 general pop- ences in environmental conditions eg adolph ulation biology lemos espinal and amaya and porter 1993 1996 in addition some stud- elias 1986 growth lemos espinal and ies have shown that altitudinal variation can ballinger 1995a and thermal biology lemos have at least a partial genetic basis smith et espinal and ballinger 1995b al 1994 ballinger et al 1996 just as studies on geographic variation have shown ferguson MATERIALS AND METHODS and talent 1993 niewiarowski and roosen burg 1993 most of these studies have focused the 2 populations we studied are located in on lizard populations in north temperate lati- the campo experimental forestal san juan tudes understanding how life histories and tetla ion1910nlon 98ww9836w on the eastern slope demography vary in response to latitude and of iztaccihuatl volcano puebla mexico at altitude combinations may be useful in identi- 3700 and 4400 in on iztaccihuatl volcano S fying variables responsible for such changes grammicusgrammicus can be found up to 4600 in eleva- in this paper we present data on demo- tion at this latitude tree line is 4000 in the graphic variation of 2 populations of scelo- low elevation site hereafter designated laguna porus grammicusgrammicus wiegmann 1828 at different of approximately 4 ha is located in a pinus elevations 3700 in and 4400 in to examine hartweghartwegiiii forest surrounding a natural lake whether variations in demography occur at lizards were seen primarily on logs and subtropical latitudes sceloporus grammicusgrammicus is stumps but were occasionally found under a small viviparous lizard that occurs from tree bark or in cracks in tree trunks this site

1proyectoloyecto fauna silvestre CENID COMEFSARH avenida progreso 5 viveros de Coyocoyoacanacan mexico DFD F 04110 school of biological sciences university of nebraska lincoln lincoln NE 68588 conespondencecorrespondence author 3departruentsdepartment of biology williawilliam jevelljevelijewell college liberty MO 64068

375 376 GREAT BASIN naturalist volume 58

laguna paredosparedon 6

y 53005.300 0194x R 085 5 y 81088.108 0247x R 088 5 4 aenmen W a 4 N 3 a a tm W U ow a a 3 2 imme J 2 MW a

0 1 30 40 30 404 0 50 60 50 60 snout vent length mm snout vent length mm

fig 1 relationship of litter size to body size in pregnant sceloporus grammicusgrammicus from low laguna and high pare don altitude populations on the iztaccihuatl volcano puebla mexico was studied from november 1984 to june recorded for each female all means are given 1988 and from september 1990 to january ls unless specified otherwise 1992 the high elevation site hereafter desig- using jolly s 1965 stochastic method which nated Paredon of approximately I1 ha is a vol- is relatively insensitive to differences in chance canic rock formation surrounded by grassland of capture or survival among animals carothers composed primarily of festuca tolucensis 1973 we calculated population density for lizards at this site live under rocks and in rock each month although young and old lizards crevices we studied this site from november may have differed in capture frequency and 1985 to june 1988 and from september 1990 survivorship bias in population estimates was to january 1992 from may 1991 to april 1992 probably small smith 1981 average minimum temperatures for these 2 lizards were aged according to size at first sites were very similar laguna 202.090go 06c capture since lemos espinal and ballinger mean isid1 paredosparedon 222.2 06c how- 1995a found that lizards from both study ever average maximum temperatures for sites show the same growth rates we used the laguna were higher 13113113.1 09c than for same size categories for both populations size paredosparedon st57575.7 05cosc lemos espinal and class I1 females 39 mm SVL males 42 mm ballinger 1995a SVL individuals in their ist yr size class 2 both populations were censusedcenscensusesused monthly females 39 45 mm SVL males 42 49 mm for each captured lizard we measured snout SVL individuals in their and2nd yr and size class vent length SVL to the nearest mm using a 3 females 45 mm SVL males 49 mm clear plastic ruler and body mass BM to the SVL 3 yr or older for life table analyses we nearest ooiool0010.01 g using a pesola spring scale estimated age by recapture of animals marked we also recorded sex tail condition broken as hatchlingshatch lings or by using von bertalannyBertabertalanffylanny 1957 regenerated or unbroken time of capture growth analyses lemos espinal and ballinger and micromicrohabitathabitat of capture site each lizard 1995a survivorship was estimated for each was permanently marked by toe clipping to age class as the proportion of marked animals examine reproduction we collected females in recaptured the following year adjacent areas more than 500 m from the 2 study sites n 67 for laguna n 54 for RESULTS Paredon during may 1991 and dissected them to examine reproductive tracts specimens litter size increased with female body size currently in JAEsJALs personal collection size and in both populations fig 1 r 0850.85 n 67 number of yolked follicles or embryos were P 000010.0001 for laguna and r 0890.89 n 54 199819981 demography OF sceloporus CRAMMICUSGRAMMICUS 377

P 000010.0001 for Paredon females from laguna leoieo1e0 4 had significantly larger litter sizes than did females from redonPaparedosparedon after controlling for dif- ferencesferences in body size with ANCOVA 3643643.64 25 0100.10olo n 54541 vs 3313.31 0130.13 n 67167 f1117F Q E N 85 laguna 4924.92 FP 0030.03 the interaction term was not 0 86 laguna ae 1 significant was no indication our 0 5e a 87 laguna there in Z 86 paredosparedon study or in that guilletteofofguillette and casas andreu w 87 paredosparedon 1980 that females have more than I1 litter U per year lizards at both study sites were born at 19 20 mm SVL females attained sizes of oe0aeooeo approximately 39 mm SVL by 14 mon of age 0 1 2 3 4 5 lemos espinal and ballinger 1995a the age years smallest reproductive female was 39 mm SVL at both laguna and Paredon but an SVL of fig 2 survivorship alxajlx1j curves for sceloporus frammigrammi high Pa typical cus from low laguna and redon altitude popu- approximately 40 42 mm was the minmin- lations on the lztaccihuatliztaccihuatl volcano puebla mexico imum size of reproductive females fig 1 these data indicate that females at both study sites mature at an age of 14 15 mon ie in their and2nd fall low elevation site had slightly larger litters annual survivorship alxlxL was calculated for than did individuals from Paredon the high 1985 86 and 1986 87 at laguna and for elevation site this difference may help 1986 87 at Paparedosparedonredon in general survivorship explain the difference in rorg between these tended to be greater at laguna than at pare populations laguna s rorg suggests a growing don fig 2 the number of individuals per ha population whereas Parparedonparedonsparesonedonss suggests a was greater at paredosparedon than at laguna for all decreasing population it is interesting to note years of study table 1 that litter sizes of S grammicusgrammicus from lower contribution of the different age classes to elevation populations 2000 3200 inm are even age specific fertility was similar at both study larger mean 525.2 than from our laguna site sites age classes 2 and 3 contributed the most guillette and casas andreu 1980 32 and 29 at laguna and 30 and 31 at the lack of major differences between these redonPaparedosparedon see table 2 average generation time 2 populations of S gramgrammicusmicus is in contrast to was 3323.32 yr for laguna and 3373.37 yr for pare several other studies of elevational variation in don replacement rates varied between years life history and demographic traits such as as did average population density table 2 growth grant and dunham 1990 smith and lower rory values in 1987 may have resulted ballinger 1994a and survivorship smith and because both study sites were sampled only 6 ballinger 1994b while it is tempting to attrib- mon in 1988 until june 1988 thus some ute differences between the present study and lizards that survived from 1987 to 1988 may other studies to geography ie differences in not have been registered latitude or elevation present study took place at higher elevations than other studies such a discussion conclusion is premature our results taken along with those of guillette and casas andreu in general the 2 populations of S frammigrammi 1980 do suggest there may be additional ele- cus studied here do not differ greatly in their vatvationalional differences among populations of S biology survivorship estimates appear to be grammicusgrammicus if a broader range of elevations slightly higher in the laguna population but were studied our results also suggest that the difference is quite small growth rates and further studies comparing populations at dif- body temperatures also do not differ between ferent elevations from a variety of latitudes these populations lemos espinal and ballinger would be useful in elucidating potential causes 1995a 1995b one of the few population dif- of life history and demographic variation in ferencesferences is litter size females from laguna the lizards and other ectotherms 378 GREAT BASIN naturalist volume 58

TABLE 1 average population density for 2 populations of sceloporus gramgrammicusgramrmcusmicus from the iztaccihuatllztaccihuatl volcano puebla mexico for 5 yr densities are given as individuals per hectare population 1985 1986 1987 1988 1991 laguna 81 79 83 52 77 paredosparedon 155 163 131 135

TABLE 2 age specific fertility rates ixmxm and RS for low laguna and high Paredon altitude populations of sceloporus gramgrammicusgramrmcusmicus from the lztaccihuatliztaccihuatl volcano puebla mexico absolute longevity is unknown but syroid5 yr old animals have been i ecordedrecorded life table was arbitrarily stopped at the end of the ath6th yr laguna paredosparedon age 1985 1986 1987 mean 1986 1987 mean

0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 2 0423 0439 0273 0378 02070.2070 207 01810.1810 181 02440.2440 244 3 0375 0398 0243 0338 03340 334 017000.170170 02520.2520 252 4 0239 0259 0154 0217 02230 223 0980 98 0160 5 0152 0169 0099 0140 01480.1480 148 00560 056 01020.1020 102 6 0095 0107 0062 0089 00960.0960 096ogg 00300 030 00630 063

Rro 1284 1372 08310.8310 831 11591.1591 159 11081.1081 ioslos108 05350.5350 535 082108810 821

acknowledgmentsACKNOWLE DGM E NTS CONANT R AND JTJ T COLLINS 1991 reptiles and amphib- ians eastern and central north america ard3rd edition co field assistance and use of facilities we houghton mifflin boston for FERGUSON GWGWANDAND LG TALENT 1993 life history traits thank G praxedis martinez and workers from of the lizard sceloporus undulatesundulatus from two popula- the campo experimental forestal san juan tions raised in a common laboratory environment tetla financial support to E was pro- oecologia 938893 88 94 JL FLORES VILLELA P GEREZ 1994 biodiversidad y vided by the nacional de investiga 0 AND institute conservationconservacionconservacion en mexico vertebradosvertebrados vegetationvegetacion y cioneschiones forestallsforestalesForestales y agropecuarias and the uso del suelo DRD R universidad nacional Autautonomaautonomyonoma consejo nacional de cienciaciancia y tecnologia de mexico mexico E is especially grateful to ing C gonza- GRANT BW AND AEA E DUNHAM 1990 elevational covari- JL ation in environmental constraints in life histories of les and S sanojasaboja sarabia for advice in in vicente the desert lizard sceloporus mernamimerriamimerrimerrlami ecology 71 and support 1765 1776 GUILLETTE LJL J AND G CASAS ANDREU 1980 fall repro- literature CITED ductive activity in the high altitude mexican lizard sceloporus grammicusgrammicus micromicrolepidotuslepidotus journal of herpetology 1414314 143 147 ADOLPH SCS C AND WP PORTER 1993 temperature activ- 1981 seasonal variation in fat body weights of and lizard life histories american naturalist 142 ity the mexican high elevation lizard sceloporus gram 273 295 micus micromicrolepidotuslepidotus journal of herpetology 15 growth seasonality and lizard life histories 1996 366 371 age and size at maturity oikosbikos 7726777 267 278 size JOLLY GMG M 1965 explicit estimates from capture recap BALLINGER RER E 1979 intraspecific variation in demogra- ture data with both death and immigration stochastic phy and life history of the lizard sceloporus jarjarrovirovi model blomebiometnkabiometrikaBiome trika 522255292552 225 247 along an gradient southeastern altitudinal in arizona LEMOS ESPINAL JAJ A AND J AMAYA ELIAS 1986 Aspectaspectsaspectosos ecology 6090160 901goi 909 generates sobrebobre laa ecologia poblacionpoblacionalal de la lagar BALLINGER RER E GRG R SMITH AND JWJ W nletfeldtNIETFELDT 1996 tija sceloporus grammicusgrammicus micromicrolepidotuslepidotus sauriasauna elevational variation in age at maturity in sceloporus iguanidae en la vertientevertiente oriental del volcan iztacizzac jarrovi an experimental evaluation southwestern cihuatl puebla cienciaciancia forestal 5912759 127 151 naturalistnatuiahst4141179179 182 LEMOS ESPINAL JAJ A AND RER E BALLINGER 1995a ecology BERTALANI FY L VON 1957 quantitative laws in metabolism of growth of the high altitude lizard sceloporus and growthglowth quarterly review of biology 3221732 217 231 grammicusgrammicus on the eastern slope of iztaccihuatl vol- carothersadCAROTHERCAROTHERSSADAD 1973 the effects of unequal catcheatchcatchabilitycatchabihtyhatchabilityability cano puebla mexico transactions of the nebraska on jolly seber estimates biometrics 297029 70 100 academy of sciences 227722 77 85 199811998 demography OF sceloporus CRAMMICUSGRAMMICUS 379

1995b comparative thermal ecology of the high SMITH GRG R AND RER E BALLINGER 1994a temporal and altitude lizard sceloporus grammicusgrammicus on the eastern spatial variation in individual growth in the spiny slope of the lztaccihuatliztaccihuatl volcano puebla mexico lizard sceloporusjarrovisceloporus jarrovi copeiacopela 199410071994 1007 1013 canadian journal of zoology 73818473218473 2184 2191 1994b survivorship in a high elevation popula- niewiarowski PH AND WM roosenburgroodenburgROOSENBURG 1993 reci- tion of sceloporus jarrovi during a period of drought procal transplanttiansplant reveals sources of variation in copela 199410401994 1040 1042 growth rates of the lizard sceloporus undulatesundulatus SMITH GRG R RER E BALLINGER AND JWJ W NIETFELDT 1994 ecology 74199274 1992 2002 elevational variation of growth rates in neonate ORTEGAOBTEGA A AND R BARBAULT 1984 reproductive cycles sceloporus jarrovi an experimental evaluation in the mesquite lizard sceloporus grammicusgrammicus journal functional ecology 82158 215 218 of herpetology 1816818 168 175 SMITH DCD C 1981 competitive interactions of the striped received 11 june 1997 plateau lizard sceloporus virvirgalusvirgatusgatus and the tree lizard accepted 20 january 1998 Urourosaurussaurus ornatus ecology 6267962 679 687 great basin naturalist 584 C 1998 appp 380 385

NEW BISEXUAL FORM OF cavernocyprisCAVERNO CYPRIS subterraneasubterraneanSUBTERRANEA WOLF 1920 CRUSTACEA OSTRACODA FROM IDAHO

okan kulkoyluoglu1kfilk6y1fioglul and gary L Vinyard 1

ABSTRACT males of cavemocypriscavemocypns subterrane a were found for the first time in an idaho spring the bisexual form is described based on soft body parts and valves the genus cavernocyprisCavernocavemocypnscypris now includes 2 bisexual and I1 parthenogenetic species

key words cypridopsinaecypndopsinae Cavernocavernocypriscdvernocypnscypris subterranesubterraneasubterraneana crenobiont bisexualformbisexual formporm ecology idaho

A recent revision of the genus cavemocypris fication is based both on soft body parts and hartmann ostracoda cypridopsinae by mar valves ecological and physical data collected monier et al 1989 suggested that this genus from the collection site are shown in table 1 contains 3 species cavemocypris subterranesubterraneasubterraneana all materials have been retained in the depart- wolf 1920 C commacoreana mckenzie 1972 and ment of biology university of nevada reno C wardi marconiermarmonierMarmonier meisch and danielopol except for 7 specimens deposited at the musee 1989 of these only C coreana and its sub- national dhistoire naturelleNaturelle luxembourg species C coreana elonelongataelongategata in south korea by dr claude meisch have been reported to exist in bisexual populations mckenzie 1972 marconiermarmonierMarmonier et al 1989 description parthenogenetic populations of C subterranesubterraneasubterraneana are known from europe and central asia and in general modern ostraostracodeostracodscods are described parthenogenetic C wardi has been reported based on their soft body parts and valves from the western united states marconiermarmonierMarmonier et detailed information about the terminology al 1989 forester 1991 ward et al 1994 an and more description of the parts can be found undescribed species cavemocypris n sp has in moore 1961 and van Morkmorkhovenworkhovenhoven 1962 also been reported from arizona danielopol MALE shell viewed dorsally is elongate et al 1994 and the width less than half the length there this study presents the first report of the is no double folded inner list on the posterior bisexual form of cavemocypriscavernocyprisCaverno cypris subterranesubterraneasubterraneana margin of the left valve LV and LV range and provides the first description of males of 0670.67 0730.73 mm is slightly longer than the right the species valve RV range 0630.63 0730.73 mm height H is less than half the length L range 0260.26 0340.34 MATERIALS AND METHODS mm and approximately equal to the width W range 0270.27 0330.33 mm in lateral view figs IA we collected 25 individuals 8 males and 17 IB valves are elongate and LV overlaps RV females on 4 august 1993 from head spring both anteriorly and posteriorly the posterior near brush creek in malad city snake valley end is slightly narrower than the anterior but bannock county tilsTIIS r38e sec 7 idaho both are rounded in dorsal view the fused specimens were collected using a hand dip zone of the inner lamella is wider at the ends net with a mesh size approximately I1 mm2mma and broader anteroventrally valves are whitish preserved in 10 formalin and subsequently opaque and smooth in some european speci- stored in 70 ethanol after isolating ostraostracodsostracodecods mens the valves may have a dorsa median from the samples we dissected specimens and band with pits some individuals may lack mounted them in lactophenollactophenol species identididenti this band C meisch luxembourg personal

1 department of biology 314 program in ecology evolution andconservationand Conservation biology university of Nevada reno reno nv89557 0015

380 1998 NEW BISEXUAL CAVERNOcavernocyfrjscavernocyprisCYPRIS SUBTERRANEA 381

TABLE 1 ecological and physical data collected from the communication in our specimens decalcifica- study site tion has destroyed this pitted area if it was name head spring malad city present and location snake valley bannock county the ist antenna fig IQ1cac or antennula al sec idaho t11s r38e 7 has 7 segments joints with long natatory latitude 42484348048434204843 N longitude n20609w1120609 W setae as in females numbers of setae on elevation 1842m1842 m each segment of al are 311 234 544 water temperature 96c unlike females there is no row of delicate conductivity 528 lscmuscmusamusem setae on the ist segment of al of male speci- ph 773 mens the rome organ r is poorly developed the and2nd antenna aaa2 is 4 segmented fig ID swimming setae of a2aa on the and2nd segment are reduced to 6 small setae 5 of which

s1sa 100um s2sa

50ym

50um 50um

1 7

z2za 77 jaj6

25um

fig 1 cavemocypriscavernocypns subterranesubterraneasubterraneana A left valve of male inner view B right valve of female inner view C ist antenna antennula al D last 3 joints of 2ndand antenna aaa2 and2nd joint bears 5 short and 1 longer setae E mandibula FV mandibular palp G rakelikeracelikerakelike organ with 7 teeth 382 GREAT BASIN naturalist volume 58

2aaa C

50um

2aaa 2 G B

50um 2aaa

50um F

is 50um

E 50um

fig 2 Cavernocavernocypriscypris subterranesubterraneasubterraneana A maxillarmaxillaemaxil lae of male formed as right prehensile palp and left 13 prehensile palp C maxillarmaxillaemaxil lae of female in normal shape D maxillulamaxillula and maxillularmaxil lular palp of male E zenker organ with 11 whorls F hemipenis lateral shield Is is spatula shaped G furcafureaburcabureaaurea of female

barely extend to the proximal margin of the are also 2 other setae klki and kak2 the position penultimate joint or podomorepodomerepodomore ard3rd segment ofkloaklof kak1 is between the ist and and2nd teeth and kak2 the ist dorsally located natatory seta extends is between the and2nd and ard3rd halfway up the penultimate joint as in females the mandibular palp edpmdp is 4 segmented 3 all sensory clubs aesthetasc Y YIy y2yya y3ya are with a respiratory plate fig IF the aX P present the aesthetasc Y is I1 segmented and and 7 setae are present on the ist and2nd and its length equal to 33 of the dorsal edge of ard3rd segments respectively the ist segment the and2nd segment and 37 of the ventral edge also carries I1 long seta and sl seta but the s2sa of the and2nd segment the t and z setae are seta is reduced or absent the P3 seta on the reduced on the ard3rd segment claws gl and and2nd segment and the 7 seta on the ard3rd seg- g2ga are about equal in length 23 longer than ment are feathery while the a seta is small and g3ga and 512 longer than the and2nd segment smooth there are also 2 small and 2 long claws are serrate length of the CMGM claw on setae on the and2nd segment which reach almost the ath4th segment is about equal to g3ga and 38 to the tips of the claws the ard3rd segment has 2 longer than the claw gm that is about the groups of setae 8 total in the istI1 st dorsal group same size as or slightly longer than y3ya are 2 small and 2 medium setae whereas the the mandibula fig IE ends with 6 teeth and2nd ventral group has 4 setae all similar in and 3 small setae I1 hairy and 2 smooth there size the ath4th segment has 2 claws and 2 setae 199819981 NEW BISEXUAL cavernocyprisCAVERNO CYPRIS SUBTERRANEA 383

25ym

50m50um

fig 3 Cavernocavernocypriscavemocypriscaoerno cypris subterranesubterraneasubterraneana male A ist thoracopod tlTI B and2nd thoracopod taUt2

the maxilla max is formed as left and right seta or 2 small setae and 1 beaklike claw prehensile palspalps with 10 12 medium sized fig ab3b no serrate setae occur apical setae placed on the opposite end of the A flagellum like furca is absent in males but endopodite right prehensile palp fig 2aaa is present in females A small hook shaped seta slightly larger than left fig ab2b the subter- is located on the proximal part of the furca minal segment of the left palp has 2 small the rake shaped organ fig IG is T shaped clawlikeclawclaylikelike setae J1 on the right palp two feathery and has 7 teeth branchial filaments are seen on the exopodite the zenker organ has I111I1 spinous whorls plate a diagnostic character of the species fig 2eae and 1 specimen of 3 shows 3 zenker two small setae aa2a are also seen on the oppo- organs rather than 2 the hemipenis fig af2f site side is of cypridopsine type and subtriangular in the maxillulamaxillula daxlmaximaxl fig ad2d bears 3 mas- shape the base of the lateral shield Is of the ticatoryticatory processes and 1 maxillularmaxillular palp the peniferum is straight the medial shield ath4th joint the terminal segment of the palp is rounded rectangular in outline with 5 6 setae I1 of which FEMALE shape and surface structure of is always small the length of the and2nd segment the valves are as in males fig IB mean of the palp is twice its width the outer ard3rd length and width of the shell of 12 females were masticatory process of the maxilmaxillulalula has 2 smaller than that of males L 0670.67 0690.69ogg mm spinelike setae and is scarcely toothed H 0200.20 0300.30 mm W 0200.20 0260.26 mm A row the ist thoracopod tlTI walking leg has a of delicate setae is found on the ist segment of long faintly toothed distal claw fig 3aaa its al the ard3rd segment has 2 setae J1 is reduced length nearly equal that of all 4 segments the in males claws gl g2ga and g3ga on a2aa are distal seta of the penultimate segment is long subequal the maxilla is shaped normally and and well developed in addition to the distal not formed into palspalps fig 2qaq2cac flagellum like claw the ath4th segment bears I1 small seta furca is present fig 2gag other appendages the and2nd thoracopod tat2 with 4 segments are similar to those reported by marconiermarmonierMarmonier et is ended with I1 pincer organ I1 well developed al 1989 384 GREAT BASIN naturalist volume 58

TABLE 2 comparison of males of the species C coreana and C subterranesubterraneasubterraneana character C coreana C subterranesubterraneasubterraneana size 0660.66ogg uhnnunmm elonelongataelongategata 0700.70 mm 072mm coreana al 6 segment 7 segment g3ga claw short seta normal t and z setae present reducedabsentreduced absent aesthetascae&thetasc Y long 2 segment I1 segment maxilla with 4 filaments 2 filaments zenker organ 6 8 whorls elongateelongataelongata 11 whorls 8 9 or 12 coreana lateral shield duck head shaped spatula shaped habitat cave waters cold springs rivers caves mountain lakes

200um20qmm

BCD 50m501tm

a 2aaa

fig 4 cavemocypriscavernocyprisCaverno cypris coreana elonelongataelongategata male A right valve B right prehensile palp C left prehensile palp D right hemipenis inner view from marconiermarmonierMar monier et al 1989 with permission of dr claude meisch

discussion terrandaterraneaterranea 2 al is 7 segmented in C subterhubter ranea but 6 segmented in C coreana 3 the determination of sexual dimorphism in g3ga claw on the antenna is reduced to a tiny cavernocyprisCaverno cypris subterranesubterraneasubterraneana is based on the short seta in C coreana but g3ga is normal and occurrence of reproductive organs and furca clawlikeclawclaylikelike in C subterranesubterraneasubterraneana the t and z setae size of the shell shape of the maxilla and dif- are reduced in C subterranesubterraneasubterraneana while C core- ferencesferences in the length of claws gl g2ga g3ga ana has 2 t and 2 z setae 4 C subterranesubterraneasubterraneana the description of females of C subterrane a has 2 branchial filaments on the exopodite by marconiermarmonierMarmonier et al 1989 also included plate of the maxilla but C coreana has 4 descriptions of 2 additional species of these branchial filaments fig ab4b 4qaq4cac 5 the lat- C coreana is bisexual the males of these 2 eral shield of the hemipenis fig ad4d is duck species table 2 can be compared as follows head shaped in C coreana but spatulate 1 C coreana fig 4aaa is smaller than C sub shaped in C subterranesubterraneasubterraneana 6 zenker organ 11 199811998 NEW BISEXUAL cavernocyprisCAVERNO CYPRIS subterraneasubterraneanSUBTERRANEA 385 whorled in C subterrane a but 8 9 whorls in acknowledgments C coreana coreana and 6 8 in C coreana elongataelongateelongata which can also bear 12 whorls mar we thank dr claude meisch musee monier et al 1989 although 1 of our speci- national dhistoirecthistoire naturelleNaturelle luxembourg mens had 3 zenker organs this is considered for his help comments and encouragement as aberrant and is not used for comparison of these we prepared this study thanks are also given 2 species however this characteristic may be to dr dinger gulenguien and his colleagues at the important if the condition is found to be com- university of istanbul turkey for their common morphological and anatomical anomalies ments and assistance obtaining the camera of some freshwater ostraostracodsostracodecods C meisch per- lucida to dr burkhard scharf department of sonal communication were reported from inland water research germany for per- sweden and poland after the Cherchernobylnobyl acci- sonal communication and to robert schroeter dent B scharf germany personal communi- university of nevada reno USA for his cation but the presence of 3 zenker organs review of an early draft of this manuscript was not observed further studies are needed to assess causal factors for this anomaly habi- references tats of these 2 species are different C coreana is known only from the cold limestone cave danielopol DLD L P MARMONIER ajA J BOULTON AND G BONADUCE 1994 world subterranean ostracod bio- waters while C subterranesubterraneasubterraneana is creno in korea geography dispersal or vicanancevicariancevicavieavicariancenance hydrobiologia blont and occupies a wider range of environ- 287119287 ilg119 129 ments including cold spring waters river and FORESTER RMR M 1991 ostracode assemblages from springs alluvial bed sediments caves and the littoral in the western united states implications for paleo hydrology the entomological society of zone of mountain lakes marmonierMarmonier et al 1989 memoirs of marconier canada iss155181155 181 201 based on these differences we propose MARMONIER P C MEISCH AND DLD L danielopol 1989 that the bisexual form of C subterranesubterraneasubterraneana is con- A review of the genus Cavernocavernocypriscavernocypnscypris hartmann generic with the bisexual forms of C coreana ostracoda cypndopsmaecypridopsinae systematics ecology and but not conspecific they constitute 2 distinct biogeography bulletin societe naturelleNaturelle de luxembourg 8922189 221 278 species MCKENZIE KGK G 1972 results of the speleological survey in south korea 1966 XXII subterranean ostracoda conclusions from south korea bulletin national science museum tokyo 1515515 155 166 twentyfivetwenty five 8 males and 17 females indi- MOORE RCR C 1961 treatise on invertebrate paleontology part Q arthropoda 3 crustacea ostracoda univer- Caverno subterhubter viduals of the species cavernocypriscypris sity of kansas press lawrence 442 appp ranea were examined the bisexual form of C VAN MORKHOVEN fecFPCMFPC M 1962 post palaeozoic ostra- subterrane a was found in a cold water spring coda volume I1 their morphology taxonomy and in idaho and is described for the first time economic use elsevier publishing company new york description of the males is based on valves 204 appp WARDJVJAWARD JV JA STANFORD AND JNJ N VOELZ 1994 spatial soft body parts and comparison with the other distribution patterns of crustacea in the floodfloodplainplain known bisexual form C coreana differences aquifer of an alluvial riverriven hydrobiologia 28711287 11 17 of morphological characters structures and ecological parameters indicate that males of C received 30 september 1997 accepted 6 january 1998 subterranesubterraneasubterraneana are different from males of C coreana but the same as parthenogenetic females ofofcofaC subterranesubterraneasubterraneana found in europe cleatgleatgreat basin naturalist 584 C 1998 appp 386 389

AN undescribedundescribedastragalcsastragalus leguminosae FROM SOUTHERN UTAH A NEW subsection OF THE GENUS AND validation OF THE combination sphaeralceajaneaesphaeralcea JANEAE WELSH WELSH

stanley L welshlwelsh1welshel

ABSFRACI one new species astragalus conconcordiuscordius welsh sp nov is described from washington and iron counties utah and section argophylhargophylli subsection concordi welsh subsect nov is pi oposedopposedproposed A complete bibliographic citation is supplied to validate the nomenclatural combination sphaeralceajaneae welsh welsh memoirs great basin naturalist 9 4231987423.1987423 1987

key words taxonomy astragalus new species nomenclature

while I1 was preparing keys to the species barneby 1964 then indicates examples of of astragalus for the flora north america pro- potential species pairs between those with ject my attention was drawn again to some basifixed and those with malpighian pubes- peculiar plants from the pine valley and kolobdolob cence possibly this is the situation between A portions of washington county and adjacent piupiutensistensis and the new proposal the species is iron county utah because of the peculiar leaf nevertheless anomalous in any of the previ- pubescence contrasting sharply with that of ously proposed subsections of section argo the pod the plants will not key to any species philliphylli known for or nevada either of the utah in section argophylli A gray previous treatments by barneby 1964 1989 or welsh et al 1987 1993 the plants superfi- subsection concordi welsh cially resemble A piupiutensistensis barneby & mab subsection nov argo berley A marianus rydberg of section similis sectionssectionesectione argophylli subsectionesubsectionssubsectione and most have philliphylli subsection argophylli argophylli in leguminilegumleguminaini pubespubescentcenti sed allter similarity been identified as such the main difdifferetdifferedferet et similis subsectionesubsectionssubsectione missourienses in from involves the long hairy pods apart habit pilis dolabriformdolabriformisis sed in legumleguminileguminaini pubescentipubes centi however the plants in question are appressed differt dolabri- strigose with definitely malpighian or tyteTYPETOTE SPECIES astragalus conconcordiuscordius welsh known from form pubescence a feature not spsanovspnovnov subsection argophylli but typical of subsection subsection concordi is clearly allied to sec- missourienses only A amphioxusamphioxys of subsection argophylli subsection argophylli with tion missourienses occurs within the range of which it shares caudex features shaggy long the plants in question and that plant has hairy pods and general habit but differs in merely strigose pods barneby 1964697 states the malpighian pubescence of the herbage it shares the feature of herbage pubescence with the subsect missourienses is neatly circumscribed members of subsection missourienses but not and defined by the pieplesencepresenceseneesence of dolabriform hairs but it would be hazardoushazaidous to assume that it is a the pod pubescence truly natural monophyletic group on the contrary this proposed new species has long passed it seems possible that the species have arisen inde- under A piupiutensistensis barneby & mabberleymobberleyMabberley pendently eitherelther singly or in pairs flomfrom already although placed in a different subsection of argophylli with basifixed vesture 01or from existing argophylli because of contrasting pubescence piecuisoisprecursors of these at some i emoteremote period in the past types it appears to be most closely allied to A

1 provo I MLL bean lifilitelifealifi science musminandMusmuseumminandinciincl deaudepudepartmenttmentament ofit botany brigham young university ut84602UT 84602

386 199811998 NEW SOUTHERN UTAH AsTMastmgalusastragalcsastragalusASTRAGALcalusGALUSCS 387 piupiutensistensis the long hairy pods of A con els is disjunct by many kilometers from the cordius are not shared by other species of sub- present proposal with the nearest approach in section missourienses but are known in some the black mountain vicinity in northeastern species in subsection newberryaniNewberryani in that iron county relationship of subsection con- subsection the most similar species so far as cordi to subsection newberryaniNewberryani appears to be pod pubescence is concerned is the strictly tenuous acaulescent not acaulescentsubacaulescentsub A walshiiwelshii astragalus conconcordiuscordius welsh sp nov barneby which has only incipiently malpighian fig 1 hairs on the herbage and differs in other regards welsheswelches milkmilkvetchvetch an endemic of similis astragaliAstraastragalogalo piupiutensitensi sectionssectionesectione argo south central utah mainly on igneous grav philliphylli subsectionssubsectionesubsectione argophylli in aspectemaspectem

g HO TY F

fig 1 photograph of type specimen of astragalus conconcordiuscordius welsh 388 GREAT BASIN naturalist volume 58 generalesgenergeneralemalem sed pubescentispubes centis dolabriform is nee 11 basifixis foliolisfbliolisfbfomoliolisliolis laepesaepe rotunrotundatisdatis vel apiculaapiculatistis et calycecacyce tantum strigulosis differt z p-0 perennial acaulescentsubacaulescentsub 9 15 cm tall from am IRON a branching caudex pubescence malpighian stems 0 6 cm long the interinternodesnodes mostly C Z GARFIELD cedar city by stipulesstipuledpules these 3.535 9 mm long concealed sti 35 0 all distinct leaves 3 9 mm long leaflets 11 17 3.535 13 mm long 121.2lgig 5 mm broad obo- 35 min 0 vate to oblanceolate or elliptic rounded to 1 KANE apiculate or acute appressed strigose on both WASHING N peduncled 1 argey sides Pedpedunclesuncles 10 cm long racelesracemes 2 to st arges 8 flowered the flowers ascending at anthesis the axis 050.5os 5 cm long in fruit bractsbraats 252.5 454.5 ARIZONA minmm long pedipedicelscels 151.5 252.5 mm long bractebracete oles 0 calyx 10510.5losios 12 mm long the tube astragalus coconconcordiuscordius welsh 858.5 959.5gs mm long cylindric strigulose the 2 map of southwestern utah showing distribution teeth 2 3 minmm long subulate flowers 21 25 fig 0off astragalus conconcordiuscordius welsh in washington and iron minmm long pink purple or whitish to lilac counties tinged pods spreading ascending sessile or nearly so the body 15 40 mm long 9 13 mm thick ovoid to lancelanee acuminate obcom- wash t39s r11wRIIW s34 28 may 1986 LC pressed almost straight to incurved densely higgins 16741 do horse ranch mt t38s shaggy hirsute unilocular ovules ca 30 r12w s24 9 july 1987 K thorne & S clark TYPE USA utah iron co flat top mt 5368 do kolobdolob plateau t38s r11w s2&34 ca 6 mi NE new harmony t37s r12w s31 2 july 1988 GI baird 3021 do north creek 6200 ft elevation 24 may 1976 S welsh K t40s r11w s343s34 3 may 1988 KH thorne & taylor and E peabody 13160a holotype BRY MA franklin 6014 do kolobdolob vicinity t40s 4 isotopesisotypesisotypes distributed previously as A mari- r11w s12 29 april 1989 SL welsh & SL anus redbrydb clark 24162 do hop valley t39s r12w OTHER collections PARATYPES ALL BRY s1223s12 23 may 1989 LC higgins 18372 USA utah iron co spring creek SE flowering occurs mainly during april and Kanarkanarravilleraville 16 may 1985 D atwood 11003 early may hence most specimens are in fruit do W grants ranch t37s r14w s27 30 the species occurs with ponderosa pine man- may 1986 RB warrick 1672 do upper zanita oak aspen mixed mountain brush grants ranch t37s r14w s36 3 june 1986 pinyon juniper and less commonly with fre- RB warrick 1762 washington co along mont poplar willow and ash or rarely with santa clara river t41s r17w s17 22 april creosote bush at 1200 1340 2600 in mainly 1961 A terril sn do E slope pine valley on sandstone or soils derived from sandstone mts t39s r13w sig 8 june 1981 D anomalous in any of the currently known atwood 7901 do 5 mi SW enterprise reser- subsections ofargophylli A conconcordiuscordius is most voir t38s r19w sl 10 june 1981 D similar vegetatively except for its malpighian atwood 9362a do S kolobdolob reservoir t39s hairs with A piupiutensistensis from which it differs r1rar11w1waw s27 8 june 1983 LC higgins &ahbah& AH also in several less tangible features ie barnum 13606 do E slope pine valley mts leaflets are commonly rounded to apiculate t39s r13w 16 may 1984 D atwood 9652 not obtuse to emarginate or acuminate calyx do pine spring wash t39s r11w s34 23 is merely strigulose not pilosulouspilosulous and at april 1984 B franklin & G baird 462 do least some pods are much longer kolobdolob terrace t39s r11w s34 7 june 1948 distribution of the species fig 2 centers SL welsh L higgins & K thorne 22941 in the harmony mountains iron county utah do pine valley mts main canyon t38s and pine valley mountains and kolobdolob plateau r14w s33 2 june 1986 RB warrick 1715 regions of washington county the area occu- do pine valley mts t38s r13w s9sa 17 may pied by most known collections is an oval 1986 RB warrick 1379 do pine spring approximately 40 km long and 20 km wide 199819981 NEW SOUTHERN UTAH astragalus 389 trending along a northwest southeast axis welsh welsh published without citation of only 2 collections are known to be remote bibliographic reference of the basionymbasionym the from the main body of the species one along full citation should read sphaeralcea janeaejaneace the santa clara river ca 10 kmkin west of the welsh welsh memoirs great basin natural- town by that name and the other from the ist 9 423 1987 basibasionymonym sphaeralcea lepto bull valley mountains southwest of enter- phylla varjaneaevarvan janeaejaneace welsh great basin natural- prise reservoir the plute milkmilkvetchvetch is mainly ist 40 27 1980 this information merely val- a plant of the southeastern great basin with idates the earlier intentional combination only a slight overlap of distribution in the pine valley mountains literature CITED of the numerous specimens initially con- BARNEBY R C 1964 atlas of american astragalus sidered to be A piupiutensistensis only one is from the RC north memoirs of the new york botanical garden 13 pine valley mountains the remainder are 1 1188 from other utah and nevada localities thus 1989 fabalesbabales pages 1 279 in A cronquist AHA H the 2 species are evidently disjunct though holmgrenHohngren NHN H Hohnholmgrengren JLJ L reveal and PK contiguous as are other closely related species holmgren intermountain flora 30130 1 279 WELSH S L NDN D ATWOOD S GOODRICH AND LCL C HIG- the genus SL elsewhere in GINS 1987 A utah flora great basin naturalist memoirs 919 1 894 A matter unrelated to the new species of 1993 A utah flora and2nd edition revised monte L astragalus was called to my attention by dr bean life science museum provo UT 986 appp gandhi gray card index bib- KN herbarium received 21 november 1997 liographerlio concerning the lack of proper for- accepted 17 february 1998 mat in what turned out to be an incomplete citation a lapsuskapsus calamus of sphaeralceajaneae gleatgreat basin naturalist 584 0 1998 appp 390 392

AGE AND GROWTH OF JUNE SUCKER chasmistes LIORUS FROM OTOLITHS

markcmarkmaracC bellbenlbenibelki

key words june sucker chasmistes liorusdiorus age growth life history otolithsotolithus utah lake

june sucker chasmistes horushofus is endemic sion of wildlife resources standard length to utah lake utah county utah miller and total length mass and sex were recorded for smith 1981 this species is federally listed as each individual lapillicapilli were removed cleaned endangered and the wild population may and embedded in epoxy resin Seriserifixserifimfix struersstruess number 500 individuals based on mark corporation westlake OH for sectioning recapture estimates C keleher utah division otolithsotolithus were sectioned in the frontal plane of wildlife resources personal communica- using a lapidary grinder struessStrustruersers model DAP tion the remaining C horus population 7 struersstruess corporation westlake OH and appears to suffer from lack of recruitment to mounted on a glass slide thin sections were the adult population apparently in part due to observed with a wild dissecting microscope at predation on juveniles by an abundant popula- 24x to determine number of annuli age and tion of introduced white bass morone chrysops images were captured via a video camera USU S fish and wildlife service 1995 present mounted on the microscope and a TARGA recovery efforts include artificial propagation board in the computer to measure annual adult C horus are captured as they proceed increments on otolithsotolithus annual increments for up the provo river to spawn galetesgametes are the first 10 presumptive annuli were mea- stripped and combined and offspring are sured with the use of MOCHA image analysis raised in captivity until they reach a size large software jandel scientific inc estimated enough to avoid predation by white bass these length at age was calculated using the follow- juveniles then are returned to utah lake ing formula modified fraserleefraser lee method except for those retained as brood stock to campana 1990 effectively evaluate survival of captive reared individuals and to estimate age at recruitment L lo10L L lorLR ROORR roR to the breeding population one must under- stand natural patterns of C horus growth in where L is estimated total length at age x lgL utah lake although adult size is well docu- is length at capture rxarxR is otolith radius at age mented no data are available on C horus x and rpRC is otolith radius at capture loly10 is esti- growth patterns in this paper I1 report growth mated length at swim up 11 mm snyder and pattern size at age and age at death of indi- muth 1988 and rorp is estimated otolith radius viduals estimated from presumptive annuli on at swim up 0090090.09 mm measured from otolithsotolithus otolithsotolithus lapillilacapillipilli from 10 C horus from the wild an age growth curve for ages 1 10 was gener- population this study provides previously ated by averaging back calculated sizes at age unavailable data on age and growth to serve as for each sex fig 1 a baseline for comparing C horus growth pat- estimated ages of individuals ranged from terns 10 to 41 yr two individuals had 10 presump- ten C horus 7 in june 1992 and 3 in june tive annuli at death in 1992 all others had 1994 died from unknown causes possibly as a more than 25 annuli at death table 1 chas result of stress associated with spawning activ- bistesmistes horushofus exhibited a 3 stage growth pat- ities and were recovered by the utah divi tern back calculated lengths at age indicated

liepartldepartidcpaitmcntent of zoology brighambnghambangham young university provo UT 84602

390 199811998 NOTES 391

TABLE 1 presumptive age sex total length year of death and estimated total length at presumptive ages 1 10 for 10 chasmistes horushonus total lengths at ages 10 yr were not calculated estimated total length at annulus presumptive age sex TL death 1 2 3 4 5 6 7 8 9 10

10 M 502 1992 87 256 355 421 461 477 492 498 501 502 10 F 526 1992 101 277 388 437 468 487 503 520 522 526 26 M 518 1992 91 259 343 380 393 414 430 449 469 470 31 F 580 1992 197 294 353 397 425 457 476 497 517 538 34 F 539 1992 149 273 302 332 351 366 390 421 434 457 35 F 568 1992 70 176 232 262 299 330 348 368 382 403 41 F 555 1992 141 204 264 320 365 395 420 438 451 455 30 F 558 1994 78 203 296 325 366 383 393 411 421 430 36 F 592 1994 97 205 251 291 327 348 373 394 405 424 37 M 502 1994 96 172 214 255 286 322 350 369 380 400

xT 544 111 232 300 342 374 398 418 437 449 460 s 317 39 45 59 64 63 60 57 54 53 49

500 although 10 is a relatively small sample size malemaie this sample may represent 2 of the remain- E female population well above the sam- E 400 ing proportion growth C pled in most studies on age and offish 300 because C liorusdiorus is endangered and individu- 4 als cannot be obtained easily validation of a 200- ages derived from otolithsotolithus has not been done validation of age estimates using otolith annuli 0 loo100 has been done for other cyprincyprinidsids leatherside chub gila copeicopel johnson et al 1995 utah 0 chub gliacilacliagila atrariaatratwariaaria unpublished data and C 0 1 2 3 4 5 6 7 8 9 10 liorusdiorus appears to exhibit similar patterns of age annulus formation ages derived from sectioned otolithsotolithus of xyrauchen tetanustexanus have been validated for younger age classes and fig 1 mean isyalsylslsy total length at age 1 10 for male n 3 and female n 7 C horushornsborus in utah lake utah co ages appeared reliable for older age classes utah USA length at age calculated from presumed mccarthy and minckley 1987 however annual increments on otolithsotolithus length at ages 10 was until a validation study of ages derived from calculated not sectioned otolithsotolithus of C liorusdiorus is possible ages in this study should be considered preliminary beamish and mcfarlane 1983 rapid growth for the first 3 5 yr and individu- in most fishes growth rate decreases after als averaged 69 of mean total length at death sexual maturity almaimaimalm 1959 chasmistes liorusdiorus by the ath5th presumptive annulus following examined in this study show a decreased rate this rapid growth individuals exhibited inter- of growth after about 5 annuli and all individ- mediate growth rates until about age 8 10 uals in this study were reproductively mature eighty five percent of mean total length at assuming that decreased growth rate indicates death was achieved by the loth annulus table probable maturation C liorusdiorus may mature as 1 growth after age 10 was further reduced early as age 5 but at least by age 10 in 1980 growth trajectories did not appear to differ the smallest reproductive individuals were between sexes within the first 10 yr and esti- 440 and 490 mm total length for males and mated length at age 10 did not differ between females respectively shirley 1983 if growth sexes FP 010.1oiol wilcoxon rank sum test SAS patterns of these fish are similar to that docu- 1990 fig 1 based on condition of monadsgonads mented in this study then the smallest indi- and presence in the spawning aggregation in viduals likely would have been 6 10 yr old provo river all individuals were reproduc- US fish and wildlife service 1995 tively mature at time of death reported total length of reproducing females 392 GREAT BASIN naturalist volume 58 was larger than males in both 1980 and 1991 transactions of the american fisheries society 112 until age 10 growth patterns do not differ 735 743 CAMPANA S 1990 reliable growth calcacalcu all SEE how are back between sexes in this study however lations based on otolithsotolithus canadian journal of fish- females were larger than males suggesting erieserles and aquatic sciences 47221947291947 22199219 2227 that differences in total length between sexes JOHNSON JBJ B MCM C BELK AND DKD K SHIOZAWA 1995 age result from increased growth of females rela- growth and reproduction of leatherside chub gila great tive to males after sexual maturation copeicopet basin naturalist 5518355 183 187 MCCARTHY MSM S AND WL MINCKLEY 1987 age estima- C horus age and growth patterns appear tion for razorback sucker pisces catastomidaecatostomidae from similar to those of other large bodied western lake mohave arizona and nevada journal of the suckers eg chasmistes cajuscujus scoppettone arizonaanzona nevada academy of science 2187918721 87 97 1988 scoppettone and vinyard 1991 xirauxyrau MILLER RRR R AND GRG R SMITH 1981 distribution and evolution of chasmisteschasrmstes pisces catostomidae in chen tetanustexanustexanus mccarthy and minckley 1987 western north america paper 696 occasional papers delayed maturity and long adult life may be of the museum of zoology university of michigan adaptations to uncertain recruitment caused ann arbor 46 appp by environmental fluctuations scoppettone SAS 1990 SAS user s guide statistics version 6 edition garscarygary and vinyard 1991 characteristics have SAS institute inc cars NC these scoppettone GGG G 1988 growth and longevity of the allowed populations of C horusborus and other sim- eulcuieui ui and longevity of other catostomids and ilar species to persist even though recruit- cyprincyprinidscypnnidsids in western north america transactions of ment has been extremely limited because of the american fisheries society lit117301117 301 307 scoppettone VINYARD recent human disturbance is my hope that GGG G AND G 1991 life history it and management of four endangered lacustrine recovery efforts can improve before recruitment suckers pages 359 377 in WL minckley and JEJ E the aging adult population becomes extinct deacon editors battle against extinction native fish management in the american west university of press I1 thank C keleher and the utah division arizona tucson of wildlife resources and US fish and SHIRLEY DLD L 1983 spawning ecology and larval development of the june sucker proceedings of the bon- wildlife service for help in obtaining C neville chapter american fisheries society 1983 horus otolithsotolithus A lambert prepared and sec- 18 36 tioned the otolithsotolithus D shiozawa C keleher SNYDER DED E AND RTR T MUTH 1988 description and G scoppetone and H tyus provided reviews identification of june utah and mountain sucker larvae and early juveniles report 88888 8 utah state and valuable suggestions for improvement of division of wildlife resources 107 appp this note US FISH AND WILDLIFE SERVICE 1995 june sucker chasmisteschasrmstes horus recovery plan salt lake city literature CITED UT 55 appp received 6 october 1997 ALM G 1959 connection between maturity size and age accepted 17 february 1998 in fishes institute of freshwater research drott ningringholmningholmholm 40540 5 145 BEAMISH RJR J AND GAG A mcfarlane 1983 the forgotten requirement for age validation in fisheries biology great basin naturalist 584 0 1998 appp 393 395

RAVENS COWBIRDS AND STARLINGS AT SPRINGS AND STOCK TANKS MOJAVE NATIONAL PRESERVE california

richard L knightlknight1knightly richard J campicamp and heather AL knight2knighty

key words common raven brown headed cowbird european starling water mojave desert

we investigated use of natural and artificial scrub joshua tree yucca brevifoliabrevifolia woodland water sources by common ravens corvus and pinyon juniper pinus monomonophyllaphylla juni- boraxcorax brown headed cowbirds molothrus perus sppapp woodland munz and keck 1959 ater and european starlings stamusstumus vulgaris between 2 june and 5 july 1993 we sam- in the mojave national preserve california pled 60 sites consisting of 20 springs 20 stock our study was motivated by earlier observa- tanks and 20 points located away from springs tions of these 3 species in which they were and stock tanks controls springs stock tanks seen at stock tanks but not detected at springs and control points were chosen from US geol- ravens cowbirds and starlings have been ogical survey maps control points were posi- viewed as detrimental to wildlife for a variety tioned 300 m from a road and 1 I1 km from of reasons including depredations on endan- stock tanks or springs all points were 3 km gered species eg desert tortoise gopherusGopherus from human habitation stock tanks and springs agassizagassiziiii by ravens boarman 1993 nest para- all contained open flowing water stock tanks sitism of native songsongbirdsbirds by cowbirds trail were metal or concrete structures and aver- and baptista 1993 and competitive displace- aged 3 m in diameter controls springs and ment of cavity nesting birds by starlings stock tanks were located so as to represent as weitzel 1988 kerpez and smith 1990 broad an area as possible within the mojave the mojave national preserve located in national preserve maximize the distance be- san bernardino county california comprises tween sites and reflect dominant regional approximately 681000 ha it falls within an area vegetation bounded on the north by US interstate high- because our study examined whether bird way 15 the east by the colorado river the species were associated with water but not south by US interstate highway 40 and the diurnal activity patterns we visited sites west by the convergence of these 2 highways throughout daylight hours there were no sta- the study area consists of mountain ranges tististicaltical differences in time of day when the 3 interspersed with basins varying in elevation site categories springs stock tanks controls 2 from 280 m to 2400 m above sea level the cli- were visited kruskal wallis cac2 3963.96 df 2 mate is seasonal and severe being warm P 0140.14 PROC univariate SAS 1987 26c26q in summer and cool lic1 11cI 1 OQ in win- we restricted our analyses to bird detections ter with an annual mean temperature of 17 50 m from the water points using a fixed 9cac s average rainfall is 12 cm for most of distance circular plot method reynolds et al the area with most precipitation occurring 1980 we counted all detections based on from december through march johnson either visual or aural observations during a 10 1968 vegetation consists of widely spaced min period and recorded behavior perched shrubs and the major floral communities in- flying singing feeding drinking observation clude alkali sink creosote bush larrea triden points were positioned so as to offer clear tatdtatatadd scrub shadshadscalescale atriplex conferticonfertifoliaconfertifolidfolia views of water sources following each count

department of fishery and wildlife biology colorado state university fortfoitfolt collins CO 80523 athe2thehe nature conservancy 633 S college fort collins CO 80524

393 394 GREAT BASIN naturalist volume 58 we surveyed an area within a 50som m radius from et al 1980 of the 7 counts at stock tanks when the water source or a designated center point cows were present we detected cowbirds only for control sites for sign droppings tracks once A more detailed study is required to doc- shells of wild burros equus asinusacinus and ument the association of cowbirds in mojave domestic cows each site was surveyed only desert rangelands with water and livestock if once cowbird distribution in the mojave desert is the total number of individuals seen during enhanced by the presence of stock tanks it each count was used to compute means sts would be worthwhile to investigate what lorfortor each species within each of the 3 site cate- impacts they are having on desert bird species gories A G statistic PROC FREQ SAS 1987 laymon 1987 in the east mojave desert we was used to test for independence of species encountered starlings only at stock tanks or in occurrences at control sites stock tanks and the vicinity of homes and towns and have yet springs to observe them at natural water sources or ravens starlings and cowbirds were not undisturbed parts of the desert seen equally at stock tanks springs and control when the US congress passed the cali- sites G 5745.745 74 df 2 P 00570.0570 057 ravens and fornia desert protection act in 1995 it rewrote starlings were seen only at stock tanks ravens the maps of southeastern california two na- 1001.00loo1 00 0260.260 26 inn 20 individuals at 12 sites star- tional monuments death valley joshua tree lings 0600.60ogo0 60 0410.410 41 inn 12 individuals at 4 sites were made national parks and the east mojave brown headed cowbirds were seen at stock national scenic area US bureau of land management made administrative tanks 04504500.4545 0290.290 29 inn 9 individuals 5 stock was an unit park tanks were never seen at control sites and of the national service and renamed the national preserve artificial water struc- were detected only once at springs olo0100.100 10 00700.0707 mojave are allowed in joshua tree national park inn 2 individuals of the ravens seen at stock tures discouraged valley national tanks all but 4 were seen drinking 80 and in death park is yet a policy regarding water only 17 n 2 of starlings were seen drink- there not in the mojave national preserve historically ing and only 1 cowbird was observed drinking has been viewed as a limiting factor for recent evidence of cattle use was observed water wildlife in much of the desert southwest this at all stock tanks at 10 control sites and at 11 belief has led to a virtually unquestioned belief burro sign was found at half of the springs that if some water is good for wildlife then at only 2 stock tanks and at none of springs more water is better broyles 1995 our study the control one tortoise shell was points hints at a possibility that there may be unan- found each of 2 different stock tanks no tor- at ticipated costs associated with water develop- shells found elsewhere toise were ment which in turn may have implications ravens are a species native to the mojave for desert wildlife communities though evidence suggests their ppopulationsopulations although our results are based on a single have increased substantially concurrent with field season and within only a portion of the human presence associated with linear right mojave desert our findings suggest that addi- of ways sanitary landlandfillsfills and agriculture tional research on bird communities associ- knight and kawashima 1993 knight et al ated with desert water sources is warranted 1993 ravens have been implicated as a more detailed studies might substantiate causative factor in the decline of the desert whether the pattern we present is indeed tortoise a federally threatened species boar- widespread in addition studies that quantify man 1993 and shells of tortoises found at bird use of different types of water and the stock tanks are within the size class suspected physical and vegetative differences associated of being consumed by ravens boarman 1993 with water types might reveal processes that brown headed cowbirds did not historically explain ecological patterns observed occur in the east mojave desert laymon 1987 we observed groups of cowbirds including we dedicate this paper to the late jack individuals copulatingcopulapopulatingting at stock tanks whereas kawashima an individual who cared more than we detected only 2 cowbirds at a spring as many for the integrity of the mojave desert they flew past cowbirds are stated to be asso- our work was made possible by southern cal- ciated with livestock mayfield 1965 rothstein ifornia edison and colorado state university 199811998 NOTES 395

literature CITED MAYFIELD HFH F 1965 the brown headed cowbird with old and new hosts living bird 4134 13 28 MUNZ AND KECK 1959 A flora BOARMAN WI 1993 when a native predator becomes a PA DDD D california uni- of california press pest a case study pages 186 201 in SSKK majumdar versity berkeley REYNOLDS SCOTT AND RAR A NUSSBAUM 1980 A EWE W miller JRJ R pratt RER F schmalz and EKE K brown RTJMRT JM editors conservation and resource management variable circular plot method for estimating bird pennsylvania academy of science press easton numbers condor 8230982 309 313 ROTHSTEIN S 1 VERNE STEVENS 1980 range BROYLES B 1995 desert wildlife watelwater developments s1saI J AND E and diurnal changes dispersion of the questioning use in the southwest wildlife society expansion in bulletin 2366323 663 675 brown headed cowbird in the sierra nevada auk 9725397 253 267 JOHNSON AWA W 1968 the evolution of desert vegetation in western north america pages 101 140 inm GWG W SAS INSTITUTE INC 1987 sasstatSAS stat guide for personal com- 6 edition SAS cary brown editor desert biology volume I1 academic puters version institute inc press new york NC 1029 appp TRAIL AND L BAPTISTA 1993 of brown KERPEZ TA AND NSN S SMITH 1990 competition between PW LFF the impact european starlings and native woodpeckers for nest headed cowbird parasitism on populations of the nuttall s crowned sparrow biol- cavities in saguaros auk 107367107 367 375 white conservation ogy 73097 309 315 KNIGHT RLR L AND JYJ Y KAWASHIMA 1993 responses of raven and red tailed hawk populations to linear WEITZEL NHN H 1988 nest site competition between the right of ways journal of wildlife management 57 european starling and native breeding birds in 266 271 northwesternnorthwestein nevada condor 9051590 515 517 KNIGHT RLR L HALH A L KNIGHT AND RJR J CAMP 1993 raven 13 september 1997 populations and land use patterns in the mojave received desert california wildlife society bulletin 21 accepted 26 january 1998 469 471 LAYMON SAS A 1987 brown headed cowbirds in california historical perspectives and management opportuni- ties in riparian habitats western birds 186318 63 70

askdadoashAsk dado

T H E GREAT BASIN naturalistIS mr W

F I1 40 vr m

I1 N D E X

VOLUME 58 1998

BRIGHAM YOUNG university gleatgreat basin naturalist 584 0 1998 appp 398 404

INDEX volume 58 1998

authorindexAUTHOR INDEX andersen W ralph 12 harperharpeikimbailt1198kimball T 1 198 ashley michael C 289 horton chris 292 austin dennis D 188 jacobi GZG Z 192 babbel david G 12 johansen jeffrey R 295 ballinger royce E 375 barnum andrew H 92 kilgore mary jane 282 baron jill S 250 klemmedson james 0 352 bates jon D 363 knight heather ALA L 393 baumann RW 192 knight richard L 393 belk mark C 390 kondratieff boris C 250 bell christopher J 82 kuenzi amy J 328 belthoff james R 167 killk6ylaoglukulkoyluoglu okan 380 blank robert R 217 bleich vernon C 265 lafrancois toben 250 lemos espinal juhojulio A 375 camp richard J 393 lynnsuellen328lynn suellen 328 cassidy kelly M 199 chace jameson FE 245 marcus W andrew 156 clark william H 295 mathews nancy E 90 conrad mark A 231 maxwell bruce 156 crawford john A 344 mcarthur E durant 1 12 cruz alexander 245 mead jim 1 82 curson david R 90 miller richard FE 363 milner gary 156 davis frank W 199 minshallMmshallshailshali G wayne 54 davison william B 285 morrisonmornson michael L 66 76 328 delk jennifer K 282 mudge joann 12 denoyelles frank jr 231 murray michael PF 199 dewey sharon L 231 donovan michael PE 87 neale jennifer CCC C 328 driese kenneth L 199 palmquist debra E 217 edge W daniel 344 pennuto christopher M 231 powers leon R 167 flechtner valerie R 295 fraas W wyatt 28 reynolds timothy D 167 frisina michael R 28 robinson christopher T 54 robinson megan 87 goguen christopher B 90 sacks benjamin N 328 hall linnea S 66 328 sanderson stewart C 1 12 hamlin robin 328 schauer andrew J 273

398 199811998 INDEX 399 schwaner terry D 87 taylor timothy J 265 severson kieth E 312 tiedemann arthur R 352 shannon joseph PE 97 147 trent james D 217 smith geoffrey R 375 smith robert L 292 uresk daniel W 312 sowell john B 273 urness philip J 188 spurrier margo frost 184 stark bill PE 282 van buren reneerenge 1 12 stevens lawrence E 97 147 vertucci frank A 231 stoms david M 199 vinyard gary L 380 sublette james E 97 147 sureda maite 76 wade brian K 273 svejcar tony 363 wambolt carl L 28 sveum colin M 344 warfa ahmed M 38 szewczak joseph M 66 welsh stanley L 45 386 szewczak susan M 66 young james A 217

KEY WORD INDEX

taxa described as new to science in this volume appear in boldface type in this index

aredusabedus herberti 292 biogeography 147 collections accepter 90 biomass 312 winter 231 aedes dorsalis 184 bird colorado 245 273 ageage390390 abundance 328 gunnison basin 273 algae species richness 328 river 97 147 soil 295 bisexual form 380 common raven 393 allenrolfea occidentoccidentalisoccidentalistalis 217 bison 245 community 147 alliance 199 bison bison 245 structure 231 allied taxa 38 bitterbitterbrushbrush 28 conservation assessment 199 altitude variation 375 black hills south dakota 312 Concontopustopus sordidulussordidulus 90 anthemideae I1 black billed magpies 289 crenobiont 380 aquatic broods 344 cricotopuscricotopnsCricotopus Cricocricotopuscricotopnstopus blinciblinni chemistry 250 ing 292 sublette 97 insect 292 parasitism 90 285 Cricocricotopustopus Cricocricotopuscricotoptistopus herr invertebrates 250 brown headed cowbird 90 manni sublette 97 andaridarld regions 66 245285393245285 393 cryptogamic crusts 295 arizona browsing 28 cylindrocystis brebissonii varvan glen canyon dam 9714797 147 bud development 28 desertidebertideserti 295 grand canyon 97 147 bufo boreas 87 cypridopsinae 380 artemisia 1 12 assessment california 265 demography 375 conservation 199 mojave desert 393 desert 66 astragalus 45 386 truckee river 328 rock pools 250 astragalus conconcordiuscordius welsh camp 156 diet 386 capitol reef national park fish292fish 292 atriplex lentiformislentiformis sppapp torresitorretorreyiyi utah 250 diploid 12 217 catavinacatavifiacanavinaCatavina 295 distribution 97 polyploidyautoautopolyploidy 12 cavemocyprisCavernocavernocypriscypris subterrane a 380 disturbance 156 250 avifauna 167 central desert mexico 295 diversity 363 Centrocentrocercuscercus urophasianus 344 floristic 312 baja california mexico 295 chasmistes liorusdiorus 390 bats 66 chironomidae 9714797 147 ecology 380 bioassessmentbioassessment 54 chiroptera 6 ecoregionecoregion 54 diversitybiobiodiversity 147 cladotanytarsus marki sub- ecotonesecotones lette 97 forest meadow 273 400 GREAT BASIN naturalist volume 58 egg predation 292 macromacroinvertebratesinvertebrates 54 predation 265 elakatothrix obtusata 295 management 156 egg 292 eolian dust 217 meadow prevalence 184 epiproctepiproct 282 subalpine 273 production 312 european starling 393 mertensia oblongifolia 38 purshia tridentata 28 euryeciouseurye cious species 97 metriocnemus stevensistevstevenskiensi sub- extinction 82 lette 97 range mexican spotted owl 76 expansion 245 fasciculochloris mexicanamexicansmexicana 295 mexico 295 summer 352 felis concolor 265 baja california 295 RAPD 1 12 feral horses 289 central desert 295 replacement rate 375 fish diet 292 microbiotic crusts 295 reproduction 375 floristic diversity 312 microtus 82 riparian habitat 328 flow regulation 147 midges 97 rodents 76 forest meadow ecoecotonestones 273 mojave desert california 393 functional feeding groups 231 molothrus ater 90 245 285 sage grouse344grouse 344 montana 28 sagebrush gap analysis 199 montane habitat 231 obligates 167 glen canyon dam arizona morphology 38 282 shrubsteppeshrubsteppe 167 97 147 97147 mortality 265 sarcobatusSarcobattis venniculatusvermiculatus 217 grand canyon 97 arizona mosquito parasitism 184 sceloporus grammicusgrammicus 375 147 dam mountain lion 265 seed germination 273 grazing mule deer 188 265 seedling establishment 273 livestock 188 great SEM 282 basin 66 national vegetation classifica- serial discontinuity concept 147 habits 289 grooming tion system 199 shoot growth 28 growth 390 nestling growth 285 small mammals 76 gunnison basin colorado 273 nevada soil habitat 54 66 344 truckee river 328 algae 295 montane 231 c352C 352 riparian 328 new k352K 352 use76use 76 species 97 386 moisture 273 habits taxa 45 n352N 352 grooming 289 nomenclature 45 386 P1352352 herbage yield 352 north america s352 hibernation 66 western 282 texture 273 horses south dakota feral 289 odocoileus hemionushemihemlomis 265 black hills 312 otolithsotolithus host parasite interaction 285 390 species hybridization 12 composition 363 Hydrachnid ia 184 parasitism euryeciouseuryecious 97 mosquito 184 sphaeromeria I1 idaho54380idaho 54380 park 273 spotted knapweed 156 idaho national engineering phosphorus 54 stocking levels 312 and environmental labora- pica pica 289 stomach contents 292 tory INEEL 167 picea engelmanniiengelengelmanniamannii 273 stonstoneflyefly 282 insects plant stream insects 231 stream 231 composition 352 subalpine meadow 273 cover 352 succession 188 june sucker 390 species 312 zonal 363 juniper debris 363 pleistocene 82 summer range 352 polyploid 12 survivorssurvivorshiphipbip 265 375 land ponderosa pine 312 sweltsaSweltsa 282 cover 199 population synonymiessynonymicssynonymies 97 management 199 density 375 lemniscuslemmiscusLemmis cus 82 size 87 tanacetum I1 life history 390 potential taxonomy 1 38 386 lizard 375 predator 87 tetraploid 12 litter 352 prey 87 thyasidesThyasides sphagnorum 184 livestock grazing 188 199811998 INDEX 401 trail 156 vegetation wilderness 156 tridentataeTrident atae 12 change188change 188 winter collections 231 truckee river california type328type 328 wyoming 231 nevada 328 zones66zones 66 typification 38 yield washington 344 herbage352herbage 352 understory 312 water 393 patterns 363 mites 184 zonal succession 363 utah 76 250 source 66 capitol reef national park western 250 juniper 363 utah lake 390 north america 282 wood pewee 90 402 GREAT BASIN naturalist volume 58

TABLE OF CONTENTS volume 58

no I1 january 1998

articles taxonomy of sphaeromeria artemisiaArtenusia and tanacetum compositae anthemideae based on randomly amplified polymorphic DNA RAPD E durant mcarthur reneerenge van buren stewart C sanderson and kimball T harper I1 randomly amplified polymorphic DNA analysis RAPD ofartemisiaofartemisia subgenus tridentataeTridentatae species and hybrids E durant mcarthur joann mudge renee van buren W ralph andersen stewart C sanderson and david G babbel 12 Bitterbitterbrushbrush purshia tridentata pursh growth in relation to browsing carl L wambolt W wyatt fraas and michael R frisina 28 identity of mertensia oblongifolia nutt G don boraginaceae and its allies in western north america ahmed M warfa 38 astragalus leguminosae nomenclatural proposals and new taxa stanley L welsh 45 regional assessment offadablewadableofwadable streams in idaho USA christopher T robinson and G wayne Minsminshallbailballhail 54 bats of the white and inyo mountains of california nevada joseph M szewczak susan M szewczak michael L morrison and linnea S hall 66 habitat use by small mammals in southeastern utah with reference to mexican spotted owl management maite sureda ndand michael L morrison 76 late pleistocene microtine rodents from snake creek burial cave white pine county nevada christopher J bell and jim I1 mead 82 notes western toad bufo boreas in southern utah notes on a single population along the east fork of the sevier river megan robinson michael P donovan and terry D schwaner 87 western wood peweespewzes accept cowbird eggs david R curson christopher B goguen and nancy E mathews 90

book review few and far between moments in the north american desert john martin campbell

I1 andrew H barnum 92

no 2 april 1998 articles chironomidae diptera of the colorado river grand canyon arizona USA I1 systematics and ecology james E sublette lawrence E stevens and joseph P shannon 97 chironomidae diptera of the colorado river grand canyon arizona USA IIIT11 factors influencing distribution lawrence E stevens james E sublette and joseph P shannon 147 spotted knapweed distribution in stock camps and trails of the S elwayselway bitterroot wilderness W andrew marcus gary milner and bruce maxwell 156 199811998 INDEX 403

breeding birds at the idaho national engineering and environmental laboratory 1985 1991 james R belthoff leon R powers and timothy D reynolds 167 mite parasitism of mosquitoes in central wyoming margo frost spurrier 184 vegetal change on a northern utah foothill range in the absence of livestock grazing between 1948 and 1982 dennis D austin and philip J urness 188

obituary arden R gaufin 1911 1997 obituary and list of publications RW baumann and gzjacobiGZ jacobi 192

book review wild plants and native peoples of the four corners william W dunmire and gail D tierney

I1 kimball T harper 198

no 3 july 1998 articles gap analysis of the vegetation of the intermountain semidesertsemi desert ecoregionecoregion david M stoms frank W davis kenneth L driese kelly M cassidy and michael P murray 199 natural history of a saline mound ecosystem robert R blank james A young james D trent and debra E palmquist 217 winter macromacroinvertebrateinvertebrate communities in two montane wyoming streams christopher M pennuto frank denoyelles jr mark A conrad frank A vertucci and sharon L dewey 231 range of the brown headed cowbird in colorado past and present jameson FE chace and alexander cruz 245 chemical and biological characteristics of desert rock pools in intermittent streams of capitol reef national park utah jill S baron toben lafrancois and boris C kondratieff 250 survivorship and cause specific mortality in five populations of mule deer vernon C bleich and timothy J taylor 265 persistence of subalpine forest meadow ecoecotonestones in the gunnison basin colorado andrew J schauer brian K wade and john B sowell 273 comparison of the epiproctepiproct structure of two closely related species sweltsafidelissweltsa fidelis banks and S revelrevelstokastoka jewett plecoptera Chlorochloroperlidaeperlidae jennifer K delk mary jane kilgore and bill PE stark 282

notes starvation and nestling ejection as sources of mortality in parasitized lazuli bunting nests

I1 william B davison 285 observations of black billed magpies pica pica grooming feral horses equus cacaballascaballusballus

1 1 I1 michael C ashley 289 fish predation on giant water bug heteroptera belostomatidae eggs in an arizona stream

I1 robert L smith and chris horton 292 404 GREAT BASIN naturalist volume 58

no 4 october 1998 articles algal composition of microbiotic crusts from the central desert ofbajaof baja california mexico

I1 valerie R flechtner jeffrey R johansen and william H clark 295 response understoryofunderstoryof species to changes in ponderosa pine stocking levels in the black hills

I1 I1 I1 I1 daniel W uresk and kieth E severson 312 bird use of riparian vegetation along the truckee river california and nevada suellen lynn michael L morrison amy J kuenzi jennifer CC neale benjamin N sacks robin hamlin and linnea S hall 328 use and selection of brood rearing habitat by sage grouse in south central washington

I1 colin M sveum john A crawford and W daniel edge 344 soil vegetation relations of recovering subalpine range of the wasatch plateau

I1 I1 I1 james 0 klemmedson and arthur R tiedemann 352 understory patterns in cut western juniper juniperus occidentoccidentalisoccidentalistalis sppapp occidentoccidentalisoccidentalistalis hook woodlands jon D bates richard FE miller and tony svejcar 363 comparative demography of the high altitude lizard sceloporus grammicusgrammicus phrynosomatidae on the iztaccihuatliztaccihuat1 volcano puebla mexico julio A lemos espinal royce E ballinger and geoffrey R smith 375 new bisexual form of Cavernocaoernocavemocypriscavernocypriscypris subterranesubterraneasubterraneana wolf 1920 crustacea ostracoda from idaho

I1 okan kalk6ymoglukulkoyluoglu and gary L vinyard 380 an undescribed astragalus leguminosae from southern utah a new subsection of the genus and validation of the combination sphaeralceajaneae welsh welsh stanley L welsh 386 notes age and growth ofjune sucker chasmistes horus from otolithsotolithus mark C belk 390 ravens cowbirds and starlings at springs and stock tanks mojave national preserve california richard L knight richard J camp and heather AL knight 393 information FOR AUTHORS the great basin naturalist welcomes previously VOUCHER SPECIMENS authors are encouraged to unpublished manuscripts pertaining to the biologi- designate properly prepare label and deposit cal natural history of western north america high quality voucher specimens and cultures docu- preference will be given to concise manuscripts of menting their research in an established permanent up to 12000 words simple species lists are dis- collection and to cite the repository in publication coucouragedraged references IN THE TEXT are cited by author and SUBMIT manuscripts to Rriehardrichard1 cbardwhardwbaraw W baumann date eg martin 1989 or martin 1989 multiple editor great basin naturalist 290 MLBM PO box citations should be separated by commas and listed 20200 brigham young university provo UT in chronological order use et al after name of 84602020084602 0200 an accompanying cover letter must first author for citations having more than two include phone numbers of the author submitting authors the manuscript and FAX number and emailE mail acknowledgments under a centered main address when applicable the letter must also pro- heading include special publication numbers when vide information describing the extent to which data appropriate text or illustrations have been used in other papers literature CITED also under a centered main or books that are published in press submitted or heading lists references alphabetically in the fol- soon to be submitted elsewhere authors should lowing formats adhere the following guidelines to manuscripts not mack GD and LD flake 1980 habitat relation- so prepared may be returned for revision great ships of waterfowl broods on south dakota manuscript preparation in general the stock ponds journal of wildlife management naturalist follows basin recommendations in 44695 700 style CBE scientific and format the manual for sousa WPWE 1985 disturbance and patch dynamics authorsors editors and publishers 6th edition aothauth ath on rocky intertidal shores pages 101 124 in council of biology editors 11 south inc lasalle STA pickett and PSES white editors the ecolo- street 1400 chicago IL 60603 USA PHONE suite gy of natural disturbance and patch dynamics 201 0101 FAX we 3122010101312 3122010214312 201 0214 do however academic press new york differ in our treatment of entries in cited literature coulson RN and JA witter 1984 forest ento- authors may consult the most recent issue of the mology ecology and management john wiley great for formatting guidelines basin naturalist and sons inc new york 669 appp TYPE AND DOUBLE SPACE all materials including literature cited table headings and figure legends TABLES are double spaced on separate sheets and avoid hyphenated words at the righthandright hand margins designed to fit the width of either a single column use WordwordperfectPerfect s italics feature for words to be 67676.7 cm or a page 14014014.0 cm use lowercase letters printed in italics use standard bond 22x28 cm to indicate footnotes leaving 25 cm margins on all sides photocopies OF FIGURES are submitted initially SUBMIT 3 COPIES 0off the manuscript 5 copies of with the manuscript editors may suggest changes fish manuscripts and the original on a 35 inch disk lettering on figures should be large enough to utilizing WordwordperfectPerfect 51slsi5.1 or above number all withstand reduction to one or two column width pages and assemble each copy separately title originals must be no larger than 22x2822 X 28 cm page abstract and key words text acknowledg- NOTES if the manuscript would be more appro- ments literature cited appendices tables figure priate as a short communication or note follow the legends figures above instructions but do not include an abstract TITLE PAGE includes an informative title no longer A CHARGE of 50 per page is made for articles than 15 words names and addresses of authors a published the rate for individual subscribers will running head of fewer than 40 letters and spaces be 35 per page however manuscripts with com- footnotes to indicate change of address and author plex tables andor numerous photographs may be to whom correspondence should be addressed if assessed an additional charge reprints may be pur- other than the first author chased at the time of publication an order form is ABSTRACT states the purpose methods results sent with the proofs and conclusions of the research it is followed by FINAL CHECK 6 12 key words listed in order of decreasing cover letter explaining any duplication of importance to be used for indexing information and providing phone numbers TEXT has centered main headings printed in all FAX number and emailE mail address capital letters second level headings are centered 3 copies of the manuscript 5 copies of fish in upper and lowercase letters third level head- papers and WordwordperfectPerfect diskette ings begin paragraphs conformity with instructions photocopies of illustrations issn0017ISSN 001736140017 3614 GREAT BASIN naturalist voissvolwolwoi 58 no 4 october 1998

CONTENTS articles algal composition of microbiotic crusts from the central desert ofofbajabaja california mexico valerie R flechtner jeffrey R johansen and william H clark 295 response of understory species to changes in ponderosa pine stocking levels in the black hills daniel W uresk and kieth E severson 312 bird use of riparian vegetation along the truckee river california and nevada suellen lynn michael L morrison amy J kuenzi jennifer CC neale benjamin N sacks robin hamlin and linnea S hall 328 use and selection of brood rearing habitat by sage grouse in south central washington colin M sveum john A crawford and W daniel edge 344 soil vegetation relations of recovering subalpine range of the wasatch plateau

I1 james 0 klemmedson and arthur R tiedemann 352 understory patterns in cut western juniper juniperusiurilurlfuniperus occidentoccidentalisoccidentalistalis sppapp occidenoccident talis hook woodlands jon D bates richard FE miller and tony svejcar 363 comparative demography of the high altitude lizard sceloporus grammicusgrammicus phrynosomatidae on the iztaccihuadiztaccihuati volcano puebiapueblaphebia mexicomaxico julio A lemos espinal royce E ballinger and geoffrey R smith 375 new bisexual form of cavemocypris subterranesubterraneasubterraneana wolf 1920 crustacea ostracoda from idaho okan kulkoyluoglukillkbyhloglu and gary L vinyard 380 an undescribed astragalus leguminosae from southern utah a new subsection of the genus and validation of the combination sphaeralceajaneaesphaeralcea janeaejaneace welsh welsh stanley L welsh 386 notes age and growth of june sucker chasmistes horushonusbonus from otolithsotolithus mark C belk 390 ravens cowbirds and starlings at springs and stock tanks mojave national preserve california richard L knight richard J camp and heather AL knight 393 index to volume 58 397