Full Issue, Vol. 56 No. 2

Great Basin Naturalist

Volume 56 Number 2 Article 16

4-29-1996

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VOLUME 56 n2na 2 APRIL 1996

BRIGHAM YOUNG university GREAT BASIN naturalist 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 8013785053801 378 5053 8013786688801 378 6688 FAX 8013783733801 378 3733 emailE mail nmshbllibyuedunmshbll1byuedu

associate editors MICHAEL A BOWERS PAUL C MARSH blandy experimental farm university of center for environmental studies arizona virginia box 175 boyce VA 22620 state university tempe AZ 85287

J R CALLAHAN STANLEY D SMITH museum of southwestern biology university of department of biology new mexico albuquerque NM university of nevada las vegas mailing address box 3140 hemet CA 92546 las vegas NV 89154400489154 4004 JEFFREY J JOHANSEN PAUL T TUELLER department of biology john carroll university department of environmental resource sciences university heights OH 44118 university of nevada reno 1000 valley road reno NV 89512 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 berranjerran T flinders chairman botany and range science duke S rogers zoology wilford M hess botany and range science richard R tolman zoology all are at brigham young university ex officio editorial board members include steven L taylor college of biology and agriculture H duane smith director monte L bean life science museum 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great basin naturalist the great basin naturalist founded in 1939 is published quarterly by brigham young university unpublished manuscripts that further our biological understanding of the great basin and surrounding areas in western north america are accepted for publication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 1996 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 pertaining 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 PO box 26889 brigham young university provo UT 84602688984602 6889

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copyright 0 1996 by brigham young university ISSN 001736140017 3614 official publication date 29 april 1996 4964 96 750 17922 the great basin naturalist PUBLISHED AT PROVO UTAH BY BRIGHAM YOUNG university

ISSN 001736140017 3614

VOLUME 56 30 APRIL 1996 no 2

great basin naturalist 562 0 1996 appp 95 118 SELECTING wilderness AREAS TO CONSERVE UTAHS biological DIVERSITY

diane W Davidson 1 william D Newmark2 jack W sites jrjrejr33 dennis K Shiozawa3 eric A Rickart 2 kimball T harper4harpere and robert B keiter5

ABSTRACT congress is currently evaluating the wilderness status of bureau of land management BLM public lands in utah wilderness areas play many important roles and one critical role is the conservation of biological diversity we propose that objectives for conserving biodiversitybiodiversity on BLM lands in utah be to 1 ensure the longtermlong term population viability of native animal and plant species 2 maintain the critical ecological and evolutionary processes upon which these species depend and 3 preserve the full range of communities successional stages and environmental gradients to achieve these objectives wilderness areas should be selected so as to protect large contiguous areas augment existing protected areas buffer wilderness areas with multiple use public lands interconnect existing protected areas with dispersal and movement corridors conserve entire watersheds and elevational gradients protect native communities from invasions of exotic species protect sites of maximum species diversity protect sites with rare and endemic species and protect habitats of threatened and endangered species we use a few comparatively well studied taxa as examples to highlight the importance of particular BLM lands

key words wilderness diversitybiobiodiversity conservation utah bureau of land management endemic species exotic species cryptocryptobioticbiotic soils plants bees vertebrates

THE wilderness ACT AND biodiversityBIO DIVERSITY historical value 16 US code 1131 cac41c4 ecological concerns have also figured promi- in the wilderness act of 1964 congress nently in several congressional wilderness endorsed the preservation of federal land in its bills for bureau of land management BLM natural state 16 US code sections 1131 36 public lands both the alaska national interest congress plainly anticipated that ecological lands conservation act 16 US code 3101 considerations were an important dimension b and the california desert protection act of the wilderness concept since the act pro- 103 public law 433 section 2 b 1 B 1994 vides that wilderness may contain ecological expressly acknowledge that wilderness designa- features of scientific educational scenic or tion is intended to protect important ecological

idepartmentepaitment of biology university of utah salt lake city UT 84112 2utahbutahtah museum of natural history university of utah salt lake city UT 84112 department of zoology brigham young university provo UT 84602 department4department of botany brigham young university provo UT 84602 5c1legescollegeSC ollege of law university of utah salt lake city UT 84112

95 96 GREAT BASIN naturalist volume 56 values among the significant ecological func- d tions of wilderness areas is their role in conserving biological diversity diversitybiobiodiversity loo100 km in utah undeveloped public lands admin- istered by the BLM fig 1 can potentially P play a key role in conserving the state s natural heritage the BLM is now pursuing an ecosystem management policy designed to ensure a sustainable ecological processes and biological diversity on lands under its jurisdiction department of the interior 1994 by using these same criteria to designate wilderness areas congress could not only advance the BLM s ecosystem management goals but also reduce conflict over the agency s multiple use lands 1aaa ege g by diminishing the risk of future endan- T gered species listings and the accompanying regulatory limitations over the long term it is both cheaper and easier to protect species in aggregate in their intact functioning ecosystems than to conserve them individually in fragmented and decimated populations under the endangered species act fig 1 map of the state of utah showing in black loca- proposed for BLM short the use of biological and ecologi- tions of all existing roadless areas in wilderness status the BLM formally studied a subset of cal criteria to designate BLM wilderness areas these areas and recommended a portion of studied lands in utah is consistent with the legal concept of for wilderness status data are from a department of inte- wilderness and would help to avoid future rior map of BLM wilderness study areas BLM proposed s BLM management wilderness and the utah wilderness coalition conflicts over resource wilderness proposal county boundaries also are shown isolated mountain ranges in utah s western deserts are biodiversityBIODIVERSITY DEFINED identified as follows a deep creek b fish springs c house range and d newfoundland range not for- designation on biological diversity the variety of life in a mally proposed or studied for wilderness the colorado plateau e the henry mountains given area includes three hierarchical components genetic diversity species diversity and ecosystem diversity ege g national research sity consists of the variety of major ecological council 1978 wilson 1988 reid and miller communities within areas that are heteroge- 1989 raven 1992 genetic diversity refers to neous in their physical attributes for example the variety of genes within species depletion in elevation or soil type of genetic diversity during population bottle- genetic species and ecosystem diversity necks or because of inbreeding within frag- all result from both interactions between organ- mented and isolated populations can threaten isms and their environments and interactions a species survival by reducing the capacity of of organisms with one another the physical organisms to adapt to changing environments environment sets limits on which species can souiesoule and wilcox 1980 frankel and soulesouie inhabit an area and interactions among those 1981 species diversity or the number of species determine which are most abundant species within a region species richness can strategies for preserving diversitybiobiodiversity must be divided into three major components therefore take note of all living things in the whittaker 1972 alpha diversity a the num- landscape and the linkages among them ber of species in a homogeneous habitat beta finally since different species specialize on diversity 83 the rate of species turnover different stages of natural disturbance cycles across habitats and gamma diversity y the it is important to preserve a range of commu- total number of species observed inm all habi- nities and ecosystems representing all stages tats within a region finally ecosystem diver in the disturbance cycle 199619961 wilderness SELECTION FOR biodiversityBIODIVERSITY 97

objectives have several of these traits on BLM lands in utah examples of such organisms are river the success of conserving biological diver- otter lutra canadensis and both bald and sity within a system of protected areas can golden eagles haliaeetus leucocephalus and only be assessed in relationship to a series of aquila chrysaetos risk prone plants include selected objectives we propose that the con- Hohnholmgrengren locoweed astragalus holmgrenio servation of utahutahs s biological diversity depends rum and jones cyccycladenialadenia Cyccycladenialadenia humilis on 1 ensuring the longtermlong term viability of native var jonesiijonejonesiiisii which have highly specific sub- plant and animal populations 2 maintaining strate requirements the critical ecological and evolutionary processes viability of populations depends on both upon which these species depend and 3 pro- the level of risk one is willing to accept and the tecting the full range of communities succes- time frame over which one wishes to conserve sional stages and environmental gradients eg the population shattershaftershaflershaffer 1981 schonewald cox IUCN 1978 mackinnon et al 1986 noss 1992 1983 soulesouie 1987 in general both survival both the size of the network of protected time and the likelihood of population persis- areas and the selection of individual wilderness tence increase with population size A level of areas should be guided by these 3 goals risk and persistence that is commonly pro- although it is possible to preserve a small sub- posed as a management goal is a 99 chance set of species and genotypes in zoological and of survival for 1000 years eg belovsky 1987 botanical gardens communities and species armbruster and lande 1993 interactions must be conserved in situ large for large carnivorescarnivores the minimum viable areas with minimal human intrusion and with population necessary to ensure a 99 chance natural processes reasonably intact are critical of survival for 1000 years is estimated to be elements of an in situ conservation strategy approximately 10000 100000 individuals be- they provide protection for fragile habitats such lovsky 1987 in habitat area this is equivalent as easily eroded soils and preserve habitat for to 100000 1000000 km2 or 252.5 25 million reclusive species moreover wilderness areas acres although this area requirement may offer natural ecosystems some protection from seem remarkably large documented losses of the biological invasions that have devastated mammalian species from among the largest of many communities especially plant communi- north american national parks eg the ties across utah 10328 km2 yellowstone grand teton park here we describe a strategy based upon assemblage during the last 90 years make widely accepted principles of conservation clear the importance of protecting large areas biology see eg primack 1993 meffe and newmark 198719951987 1995 for both selecting critical sites carroll 1994 maintenance of ecological for wilderness designation and determining and evolutionary processes the amount of habitat that should be pre- served as wilderness see also babbitt 1995 in selecting wilderness areas one must take care to ensure the maintenance of the CRITERIA FOR SELECTION ecological and evolutionary processes upon which all plant and animal species depend viable populations pickett and thompson 1978 kushlan 1979 utah contains approximately 3000 indige- among the most important of these processes nous plant species and varieties and about 584 are natural disturbance and recovery cycles vertebrate species viable populations for most ideally criteria for the selection of wilderness of these plants and animals can be ensured by areas should include information on fre- focusing within ecological communities on quency size and longevity of natural distur- species for which the risk of extinction is bances protected areas should be large greatest risk prone species typically include enough to contain minimum critical areas of those with small populations large home the entire range of recovery stages for each range requirements low reproductive poten- community type pickett and thompson tial restricted geographic ranges or large 1978 in western north america natural dis- temporal variation in population size brown turturbancebance regimes can encompass tens of thou- 1971 willis 1974 terborgh and winter 1980 sands to millions of acres as witnessed by the diamond 1984 pimm et al 1988 belovsky et recent and extensive wildwildfiresfires in yellowstone al 1994 newmark 1995 many top predators national park christensen et al 1989 98 GREAT BASIN naturalist volume 56

two other critical ecological processes are designating wilderness areas high priority migration and dispersal of terrestrial organ- should be given to lands whose selection ismsiams across landscapes and of aquatic species would enlarge and connect existing protected within watersheds the selection of wilder- areas eg national parks wildlife refuges and ness areas requires that attention be given to forest service wilderness areas and thus ensuring that migratory pathways are open to enhance the viability of animal and plant pop- organisms migrating seasonally along eleva- ulationsulations newmark 1985 salwasser et al 1987 tional gradients of particular importance is noss 1992 grumbine 1994 by themselves the need to maintain winter ranges and migra- BLM wilderness areas in utah clearly cannot tory routes of large maimmalmmammalsnalsnais such as mule deer satisfy the huge area requirements noted above odocoileus hemionus elk cervus claphuselaphus as requisite for maintaining viable populations and moose alces alces of large carnivorescarnivores however when linked to interactions among competitors and be- other public lands eg utah s national parks tween predators and piespresprey are integral aspects and wilderness areas in other states BLM of natural ecosystems and should be pre- wilderness in utah can be a key component of served for example in the southwestern strategies for longtermlong term preservation of biolog- deserts of the united states the direct and ical diversity indirect effects of seed predation on plant other high priority areas are those which community structure have been documented alone or together with other protected areas in longtermlong term experiments manipulating densi- encompass entire watersheds in addition to ties of rodent and ant granivoresgranivores davidson et affording direct benefits to humans watershed samson al 1984 et al 1992 these effects protection is the most effective means of con- include transformation of a shrubland into a serving the aquatic and riparian communities grassland biome brown and heske 1990 that account for a disproportionate fraction of special care must be taken to conserve popu- both species diversity and endangered and lations of predators with large area require- threatened species in andaridarld western north because of ments extinctionsextinctions these species america miller 1961 minckley and deacon whole g by leading can alter communities ege 1968 1990 holden et al 1974 johnson et al to outbreak densities of prey which then over 1977 cross 1985 knopf 1985 moyle and exploit some of the their plant resources williams 1990 moreover since populations evidence for such trophic cascades strongest of riparian species are usually isolated from comes from the greater yellowstone ecosys- similar communities in other drainage systems where browsing by elk has tem intensive species losses from these environments are greatly altered many zones by the re- riparian not easily remedied by natural recolonization moval of willows genus salix and has elimi- A ard3rd priority in selecting wilderness sites nated aspen seedlings populus tremultremuloidesoides is land that forms or helps to complete the pro- recruiting from seeds and rhirhizomeszomes shortly tection of entire elevational gradients for after the extensive 1988 fires contem- huge example in isolated mountain ranges of the porary elk herds numbering 40000 individ- great basin scant attention paid to conserving uals in the paripaikpark and 20000 in the northern in in these gradients in the past is evident in the herd alone are likely the result of reductions restriction of most national parks and wilder- in the full complement of large predators kay in ness areas in western north to higher 1990 wagner et al 1995 considerable america evi- elevation of wilderness in dence also suggests that deer and elk herds in sites designation in comparatively low elevation BLM lands would utah average significantly larger at present afford protection to regions of greatest species than during any extended period in the histor- in richness for organisms mammals ical past durrant 1950 julander 1962 harper many eg amphibians and whose 1986 birds insects trees diversity generally declines with elevation much of strategies FOR SELECTING throughout western north america 1984 stevens 1992 wilderness AREAS harris landscape wide priorities optimal design goals given the large area requirements of many if BLM wilderness areas are to contribute extinction prone utah species it is important substantially to the preservation of biodiver to protect large contiguous land blocks in sity in utah then site selection must take into 199619961 wilderness SELECTION FOR biodiversityBIODIVERSITY 99 account the 3 general goals outlined above BLM wilderness ideally BLM wilderness lands should form an forest service wilderness interconnected core zone of roadless lands when combined with other federal wilderness national park areas national and state parks and wildlife wilderness multiple use refuges fig 2 special attention should be public given to linking roadless lands so as to pre-preclude further fragmentation of natural habitat fragmentation or the transformation of an unbroken block of natural habitat into a number of smaller patches separated by altered habitats reduces population sizes increases core zone their isolation and threatens their longtermlong term viability is one of the bio- it greatest threats to buffer zone logical diversity worldwide wilcox and murphy 1985 wilcove et al 1986 saunders et al iifhllhlfbif I1 1991 across diverse habitats there are numerous examples of species extinctionsextinctions precipi- core zone tated by both natural and human induced habitat fragmentation eg brown 1971 terborgh and winter 1980 diamond 1984 fig 2 an example of a preferred arrangement of heaney 1984 patterson 1984 newmark 1987 wilderness and multiple use federal and state lands to 1991 1995 case and cody 1988 soulesouie et al conserve biological diversity wilderness areas adminis- by the management 1988 bolger et al 1991 tered bureau of land forest service national park service and fish and wildlife service adjacent multiple use lands can buffer should form a contiguous core zone in which the most human impacts on biological diversity within extinction prone species in utah can be protected multi wilderness areas such lands can be expected pie use lands can effectively buffer this core zone and to provide marginal habitat for the many species provide additional marginal habitat to species that are pri- that are restricted primarily to more pristine marily restricted to roadless areas wilderness regions thus proposed wilderness areas surrounded by public lands should receive high priority for protection their genes move about only through the processes of seed dispersal and pollen transport EXAMPLES OF RARE AND therefore it is not surprising that many plants ENDEMIC SPECIES have narrowly restricted ranges are locally adapted to conditions within those ranges and the design advocated above is based isolated often by distances from largely on conservation strategies for preserv- are great other sites similar prevail ing wide ranging vertebrate species although where conditions although locally endemic plants such strategies can help to ensure the long- can often be relatively inside ranges term viability of most species within a given abundant their their region exclusive reliance on such approaches populations are easily jeopardized by habitat may well overlook and endanger many locally alteration eg by all terrain vehicles within isolated rare and endemic plants and animals their narrow distributions of utah s approxi- we cannot give a comprehensive treatment of mately 2600 plant species and 400 named this subject here but we discuss 3 taxonomic varieties albee et al 1988 welsh et al 1993 groups of organisms for which especially high about 180 or 7 of species are currently clas- rates of endemism or existing threats to iso- sified by federal or state agencies as endan- lated populations present particular manage- gered threatened or sensitive A majority of ment dilemmas that should be taken into these 133 or 74 definitely or probably account in wilderness decisions in most cases occur on BLM lands atwood et al 1991 and specific habitats must be protected to assure a substantial subset of the classified species the preservation of these species are narrow endemicsendemics shultz 1993 provides a useful summary of plants of special concern endemism in the utah flora approximately unlike the wide ranging animals discussed 240 species or 10 of all utah plant species are above plants occupy fixed positions they and endemic to the state this rate of endemism 100 GREAT BASIN naturalist volume 56 the percentage of the flora considered for list- plant species are endemic to the region in and ing as threatened or endangered and the per- around the proposed wilderness area PWA centage of rare species inm the flora are among near the white river south of vernal UWC the highest in the continental united states 1990 and most of these are confined to the the vast majority 86 of utah endeendemicsmics reside parachute and evacuation creek members of in and and semiarid regions of the state and the green river shale formation another 90 are edaphicedaphicallyedapbicallyallyaily restricted to finehine textured dozen endeendemicsmics occur in a diversity of habi- andor high ph substrates limestone clay silt tats in and around the san rafael swell here mudstone and shaleshaie that magnify drought the most important habitat is a beige rather stiessstress plant distributions generally appear to than red Moenmoenkopikopi formation spatially iso- respond more to edaphic topographic and lated from other Moenmoenkopikopi outcrops and un- geologic features of the environment when usual in its soil chemistry A few endemicsendemics also drought is a factor stebbins 1952 because occur on the younger carmel and summer- most endemicsendemics live in close proximity to mor- ville formations surrounding the core of the phologicallyphologically similar species albee et al 1988 swell especially between muddy creek and these species appear to be mainly neoendemics crack canyon S welsh personal communica- that have evolved since the last glacial maxi-maxi tion wilderness designation in these 2 regions mum 18000 arsyrs BP or in the bonneville basin the san rafael PWA and the white river during the past 10000 arsyrs PWA of the uinta basin fig 3 see UWC geographically endemism of utah plants is 1990 could afford significant protection to some highest in the Canyoncanyonlandslands phytogeographic of utah s endemic plants south and east of section of the colorado plateau division of the san rafael in the dirty devil PWA UWC the intermountain region cronquist et al 1990 are the distinctive flora of the orange 1972 fig 3 modified from shultz et al 1987 cliffs region fig 3 and some additional nar- an unusual diversity of substrates occurs here row endeendemicsmics deserving protection in the main and these substrates are more apt to be exposed and south forks of happy canyon shultz et rather than covered with alluvium as in other al 1987 areas of semiandsemianasemiarid utah welsh et al 1993 thus the Moenmoenkopikopi formation is also important fully 50 of utah s 240 rare and endemic as a substrate for endeendemicsmics elsewhere in semi- plant species occur on the colorado plateau arid utah two federally listed endangered whereas just 15 occur in the great basin species arctomecon humilis the dwarf bear 11 in the mojave desert and 10 in the claw poppy and Pediopediocactuscactus filerisileri a cactus uinta desert welsh 1978 shultz 1993 and several other species are endemic to par- about half of utah s endemicsendemics belong to just 5 ticular Moenmoenkopikopi outcrops in southwestern genera that are both common and physiologi- utah wherever possible the boundaries of cally adapted to aridity total utah species and wilderness areas and other protected areas peipercentper cent endeendemicsmics in parentheses astragalus should encompass these specialized habitats fabaceae 114 3683636.88 Penpenstemonpentstemonstemon scrophulanscropbulari aceae 106 264 cryptanthaCryptantha boraginaceae aceal bees and wasps in the 61 36136.136 1 eriogonum polygonaceae 60 san rafael desert 23323.323 3 and erigeronengeron asteraceae 54 2412424.11 welsh et al 1975 welsh 1978 shultz 1993 because of their capacity for directed move- because most of the state s endemic plants ments animals are less likely than plants to are restricted to particular geologic formations exhibit high rates of endemism nevertheless and because multiple endemicsendemics often occur on since insects often tend to be host or habitat the same formation groups of endemicsendemics gen- specific eg in pollinatorspollinators herbherbivoresivores or sub erally can be protected simultaneously by safe- strate specific ground destersnenesterssters endemism can guarding those soil formations and surround- often be high in insect taxa bees and wasps ing areasai easaas two regions where largelailalial ge numbers of order hymenoptera are examples of such endeendemicsmics stand to benefit from wilderness insects here as elsewhere bees and preda- protection of BLM lands are the uinta basin tory wasps are especially diverse in andaridarld regions and the san rafael swell and surrounding san michener 1979 the state supports a mini- rafael desert fig 3 table 1 M windham mum of 950 species of native bees roughly personal communication no fewer than 15 25 of the total number of species known 199611996 wilderness SELECTION FOR biodiversityBIODIVERSITY 101

1I va

V laintaljinta in

swell t

san rafael4 el desert

orange cilnis

fig 3 satellite image of utah showing the positions of the san rafael swell the san rafael desert and the orange cliffs all within the Canyoncanyonlandslands phytogeographic section outlined in bold the arrow in the uinta basin shows the approximate position of the white river PWA utah wilderness coalition 1990 from america north of mexico and 50 of the Nefneffleffandneffandfandand simpson 1993 and many of the areas utah species are currently undescribed T currently under consideration for wilderness griswold FE parker and V tepedino personal designation in utah are centers of endemism communication many areas especially in the for both groups although we lack extensive in- southern part of the state have not been formation on bees of the canyonlandsCanyon lands section explored intensively and undoubtedly harbor fig 3 where endemism is highest for plants many additional undescribed species see above intensive collecting in that small bees and plants often show comparable geo- part known as the san rafael desert has graphic patterns in diversity and endemism yielded a total of 316 species of bees 42 of 102 GREAT BASIN naturalist volume 56

TABLE 1 plants endemic to the 2 areas with the highest endemism on utah BLM lands

endemicsEnde mics of the southern uinta basin endemicsEndemics oftheodtheof the san rafael swell aquilegia barnebarnebyibyi munz ranunculaceae astragalus rafaelensis jones fabaceae astragalus equisolensiseqmsolensis neese & welsh fabaceae cryptanthaCryptantha creutzfeldtcreutzfeldiicreutzfeldn welsh boraginaceae A hamiltoniiharmltomihamiltonii C porter C johnjohnstoniijohnstomijohnstoniistomi higgins A lutosuslukosus jones C jonesianajone siana payson payson A sausaurinussaturinussaunnusrinus barneby erigeronengeron maqwreimaquireimaquireifelfei cronquist asteraceae cirsiumcirstum barnebyiharnebyibarneharnebyi johnst asteraceae lomatiumjunceumlomatium junceum barneby & N Hohnholmgrengren apiaceae Cryptcryptanthaantha barnebyibarnebyi johnst boraginaceae lygodesmia intradaentrada welsh & goodrich asteraceae C gragrahatniigraharmiharmi johnst Pediopediocactuscactus despaidespaldespainiidespaimidesp aimiaimoatmoniinil welsh & goodrich cactaceae cymopteriscymoptens duchesnensis jones apiaceae Penpenstemonpentstemonstemon marcumarcummarcomsiislisll keck N holmgren scrophulanaceaescrophulariaceae penstemonflowersiipenstemonflowersti neese & welsh schoencrambeSchoencrambe barnebarnebyibyi welsh & atwood rollins brassicaceae scrophulanaceaescropbulariaceaescrophulariaceae talinum thompsothompsonnnii atwood & welsh portulacaceae P goodrgoodrichiigoodnchiiichii N Hohnholmgrengren townsendia abricaaprica welsh & reveal asteraceae P grahgragrahamiigraharmiharmiamiiamli keck schoencrambeSchoencrambe argillaceaargillacea welsh & atwood rollins brassicaceae S suffrutescentsuffrutescens rollins welsh & chatterly sclerocactus glaucus K schum L benson

which are presently undescribed T griswold in semiarid utah assuredly depends on their E parker and V tepedino personal communi- relationships with bees for example a rare cation thus 33 of the state s total species species of perdita found in utah only at the count and 84 of utah s undescribed but beehive dome site southeast of st george cataloguercatalocataloguedgued species are endemic to a region pollinatespollinates the rare and endangered dwarfbeardwarf bear comprising just 202.0 of the state s land area claw poppy V tepedino personal communica- furthermore a significant portion of this fauna tion bees that have specialized by collecting 24 occurs only on the colorado plateau pollen only from flowers of a particular plant the remainder of the Canyoncanyonlandslands phytogeo- family or even from a single genus within a graphic section in which the san rafael desert family are termed oligoleges such bees tend is embedded is likely to be equally diverse to be most common in andaridarndannd regions neff and and to have as many new species simpson 1993 and generally are regarded as other hymenopteran groups such as the being closely adapted to the phenology and aculeate wasps also are highly diverse in the floral traits of the plants on which they spe- san rafael desert T griswold E parker and cicialize such adaptations tend to make them V tepedino personal communication for ex- superior pollinatorspollinators squash bees and squash ample with a total of 22 species there the cir flowers are examples of such a co adapted pair cumcumglobalglobal genus philanthus is more diverse in in the americas tepedino 1981 some oligoaligo the san rafael desert than anywhere else in leges may one day prove to be useful as crop north america and probably the world these pollinatorspollina tors the legume specialist osmiacosmia san predatory digger wasps nest in the soil and rafaellerafaelaerafaelae a native of the san rafael desert has may have diversified in response to the varied been investigated as a potential pollinator of substrates present in this desert clearly des- alfalfa medicago sativadativasatwa L an important for- ignation of wilderness in the san rafael region age crop parker 1985 1986 many of the see UWC 1990 could afford significant pro- species of the san rafael desert appear to be tection to an area of very high endemism and oligoleges A brief list of some of the unde- diversity for the order hymenoptera scribed and recently described bee species bees and wasps are among the most benefi- and their host plants is provided in table 2 cial insects predatory and parasitic wasps help these entries were chosen only to illustrate to control populations of pest species eg the variety of plant taxa upon which native grasshoppers aphids etc below outbreak bees specialize densities an estimated 67 of flowering plants and depend on insects primarily bees for pollen native endemic fishes transfer and sexual reproduction axlerod freshwater ecosystems are natural habitat 1960 and the welfare of many plant species islands as such their longtermlong term isolation by 199619961 wilderness SELECTION FOR biodiversityBIODIVERSITY 103

TABLE 2 pollen preferences for representative oligolectic bees in the san rafael desert data from T griswold FE parker and V tepedino personal communication plant family plant genusspeciesgenus species bee species asteraceae helianthus anoanomolusmolus perdita nr laticincta hesperapisHesperapis sp wyethia scabiascabra perdita boharbohartorumtorum boraginaceae coiColcoldeniacoldemacordeniadeniadenla perdita heteroperditaHeteroperdita sp stanleyaStanstanleybleya perdita nr zebrata euphorbiaceae euphorbia parryiparrys perdita ninr labergei fabaceae astragalus ashmeadiellaAshmeadiella ninr mimichcheneneneri loasaceae mentzelia multifloramultiflora perdita multifloraemultiflorae onagraceae camissoniaCamis sonia dufoureaDufourea sp papaveraceae argemone perdita ute polemoniaceae gilia perdita ninr giliaegildae perdita elongaticeps scrophulariaceae Penpenstemonpentstemonstemon anthocopaAntho copa sp undescribed species intervening terrestrial habitats or by unsuitable bonneville cutthroat trout oncorhynchus clarki aquatic habitats often promotes local special- utah once thought to be extinct behnke izationization evolutionary diversification and endem- 1992 survives in populations in trout creek ism in aquatic organisms seven centers of and birch creek within the deep creek PWA endemism are recognized for fishes of western UWC 1990 north america miller 1959 and utah includes where protection of whole watersheds is substantial portions of 2 of these centers the not possible wilderness that includes key habi- bonneville basin and the colorado river tats may help to stabilize declining populations basin collectively 28 fish species are native of native fishes preclude new listings and draft to these basins smith 1978 and 27 are extant ings of recovery plans and promote recoveries because of their limited distributions en- and delistings this should be the case most demic species are easily endangered by both often for fishes living in headwater streams habitat alterations and introductions of nonna- protected by natural and artificial downstream tive competitors and predators seven species barriers from unintended invasions of alien and subspecies from the bonneville and col- coldwatercold water species for example habitat in the orado basins are now federally listed as endan- upper book cliffs desolation canyon PWA gered US fish and wildlife service 1993 A may support the colorado river cutthroat trout further 11 species and subspecies are consid- oncorhynchus clarki pleuriticpleuriticusus considered ered by fishery specialists to be endangered the rarest of the cutthroat taxa behnke and threatened or of special concern in utah war- zamzarn 1976 and federally listed as a category 2 ren and burr 1994 the decline of native species kerchner 1995 although the region fishes has been associated with both water- has not been surveyed for this subspecies shed development eg reservoirs irrigation native populations occur in streams entering diversions channelization floodplainfloodplain drainage the deschesneduschesneDuschesne river from the north shiozawa and the introduction of alien species and evans 1994 and have recently been found conservation of endemic fish populations in streams of the western book cliffs closer to has been especially successful when much of price and soldier summit shiozawa and evans the watershed has been protected williams unpublished data given these observations it 1991 but adherence to strict legal definitions is likely that streams flowing into the book of wilderness often precludes such wide- cliffs desolation canyon PWA will also con- spread protection in utah opportunities for tain this subspecies protecting entire watersheds are limited to in relatively large downstream systems relatively small drainage systems extending secondary and tertiary streams key habitats from stream headwaters in mountain ranges of include floodfloodplainplain wetlands among the first the bonneville basin to dry or saline lake beds habitats to be lost due to human activities at lower elevations A particularly important although wetlands have been viewed tradi- case is in the deep creek range where the tiotionallynally either as breeding sources for insect 104 GREAT BASIN naturalist volume 56

pests or as waterfowl production sites periodic argentissimus and the virgin river chub gila or continuous connection to rivers renders lobustarobusta semiseminudaseminudeseminuddnudanudd are endemic to this system them important appendages to lotic systems two additional species the flannelmouth sucker densities of aquatic invertebrates are signifi- and the desert sucker catostomus clarki cantly higher in wetlands than in mainmaln river have evolved very slender caudal pedunclespedpeduncleduncles channels over 100 fold in some cases wolz possibly as a response to occasional high flows and shiozawa 1995 mabey and shiozawa in the virgin river smith 1966 unpublished data floodplamfloodplainFloodplainplam wetlands can the health of this unique fish fauna already therefore serve as important nursery grounds is cause for concern two of the endeendemicsmics the for larval and immature native fishes woundwoundfinfin and the virgin river chub are feder- the loss of wetlands may be a significant ally listed as endangered although the desert factor endangering several native fishes in the sucker occurs in arizona nevada and new colorado river tyus and karp 1989 fishes mexico this species merits special concern in native to the larger streams and rivers of the utah utah division of wildlife resources colorado river basin are predominantly minmin- UDWR 1992 where it is limited to the virgin nows cyprinidae and suckers catostomidae river drainage loss of either this species or that have evolved in isolation are adapted to the flannelmouth sucker from the virgin river unique local conditions of this drainage ege g system would eliminate only a subset of their heavy silt loads and wide fluctuations in dis- existing populations and is unlikely to move charge and temperature and are the most either species to endangered status however morphologically distinct fishes in north amer- the uniqueness of these populations smith icaleaiea hubbs 1940 1941 deacon and minckley 1966 may warrant their designation as sepa- 1974 minckley et al 1986 four of these rate subspecies this together with the concern native species the colorado squawfish ptyapty now evidenced for the flannelmouth sucker chochchocheiluseilus lucius the humpback chub gila throughout its range could easily translate into cypha the bonetailbonytailbonytail chub cliacilagila elelegansdelegansegans and candidacy for listing if existing populations are the razorback sucker xyrauchen tetanustexanustexanus are not protected now federally listed as endangered the decline concern for native fishes of the virgin river of both the blueheadbluebeadbluehead sucker catostomus pan drainage has already constrained water devel- tosteustosteus discobolus and the flannelmouth sucker opment in washington county utah any actions catostomus latiiatilatipinnislatipmmspinnis within the mainmaln stems that would help preserve the integrity of ripar- of the colorado and green rivers may result in ian habitat and stream channels would also their listings as threatened especially if popu- reduce stress for these fishes since the integ- lations in tributary streams are not stabilized rity of riparian habitats is best maintained over several of these species occur in areas under large areas wilderness designation in PWAs of consideration for wilderness status both the the beaver dam slope and the greater zion price river in the book cliffs desolation can- area would serve this purpose yon PWA and the san rafael river in the san finally protection of utah s rare and en- rafael PWA have populations of roundtail dangereddangered fishes would likely also afford signif- chub flannelmouth sucker and blueheadbluebeadbluehead sucker icant protection to other aquatic organisms blueheadbluebeadBluehead sucker are also known from the for example utah s diverse communities of dirty devil and muddy creek drainages smith aquatic insects reciprocally the maintenance 1966 and both flannelmouth sucker and round- of high species diversity in stream insect com- tail chub are likely to occur there wilderness munitiesmunities is critical to assuring a continuous designation could broaden the protected ranges food supply to fishes in rivers with wide sea- of several of these species by stabilizing wet- sonal and annual fluctuations in flow rates land habitats in the dirty devil san rafael Maymayfliesflies ephemeroptera are among the best and book cliffs desolation canyon PWAs studied stream insects in utah and 16 18 although the virgin river drainage is also genera 22 24 species are known from warm part of the colorado river basin it has a water tributatributariesries of the colorado river system unique fish fauna that appears to have evolved G edmunds personal communication con- in isolation from populations min other parts of structionst of reservoirs on these rivers has the basin the virgin river spinespinedacedace lepilepf already inundated many river miles and altered bomeda mollispinusmollispinus the woundwoundfmwoundfinfin plagoptentsplagopterus flow rates sediment loads and downstream 199619961 wilderness SELECTION FOR biodiversityBIODIVERSITY 105 temperatures Maymayfliesflies and other aquatic insects reptiles and amphibians for example the great are highly sensitive to all these variables basin spadefoot toad scaphiopus intermanintermon unnaturally constant temperatures in tailwaterstailwaters tanus the western whiptail lizard cnemi- beneath dams can lead to depauperate com- dophorusdophorus tigris and possibly the rubber boa munitiesmuni ties of maymayfliesflies and other stream insects charina bottaibottae reach their eastern distribu- for example below flaming gorge reservoir tional limits here three additional species edmunds 1994 1995 four mayfly genera the longnoselongnose leopard lizard camgamGamCamcambeliacammeliagambeliabelia wis from this area of extremely high natural diver- lizeniilizenielizenii the collared lizard crotaphytus col sity have not been collected since the dam was laris and possibly the plateau striped whip- built habitats rich in maymayfliesflies and other tail cnemidophorus velox are represented aquatic insects and most in need of protection here by edge populations at the periphery of from future impoundments include the green their respective ranges other species such as river from the colorado border to ouray the northern leopard frog rana pipiens east- utah and the colorado river from the colo- ern fence lizard sceloporus undulatesundulatus great rado border to moab utah relatively warm plains ratratsnakesnake elapheelephe gutguttaguttatdguttataguttatedtatatd and the utah sections of the duchesne uintah white milk snake lampropeltis triangulum have escalante virgin and santa clara rivers their westernmost limits in this region steb- would also be sensitive to manipulations of bins 1985 unpublished BYU museum records stream flows while none of these species is federally listed as threatened or endangered a few are so EXAMPLES OF biologically listed by the state UDWR 1992 moreover IMPORTANT SITES LANDS ON BLM geographically peripheral populations such as these are particularly important as dynamic floras and faunalfaunas in different parts of the foci of evolutionary change eg brown 1995 utah have unique evolutionary histories determined by the geography and topography of lesica and allendorf 1995 book cliffs tavaputsTavaputs also sup- the lands they inhabit in this section we dis- the region a rich mammalian fauna although cuss 4 such sites in the context of important ports our is far from complete scientific criteria outlined above for wilder- knowledge the area con- at least 62 rel- ness selection we also review various sci- tains native species including a site ursus entific and educational values of these same atively stable population of black bear sites americanosamericanusameric anus H black personal communication recent fieldwork has resulted in records for 6 book cliffs and the tavaputsTavaputs plateau species previously unreported from the region for several reasons the book cliffs and D rogers personal communication these tavaputsTavaputs plateau areas along both sides of the include merriamsMerrmerriam7siamslams shrew sorex merrimerriamiami green river are critical for the longtermlong term con- dwarf shrew S nanus water shrew S palus servation of biological diversity in utah this tris big free tailed bat nyctinomops mamacrotismacrotincrotis region contains some of the largest remaining northern flying squirrel ciaugiauClauglaucomyscomys sabrinussabrinus roadless areas on BLM lands in utah fig 1 and western jumping mouse zapus princeps and therefore provides important habitat for of these species S merrimerriamiami S nanus and N sensitive species with large area requirements macrotistnacrotismacrotinmacrotis appear to be rare throughout their it includes broad elevational gradients with known distributions more fieldwork is likely the potential to protect a wide range of natural to produce additional records for this region communities and to maintain crucial routes isolated desert mountain ranges for seasonal wildlife migration between high and low elevation furthermore it constitutes the isolated mountain ranges in utah s great a vital dispersal corridor linking the uinta moun- basin and colorado deserts are extremely tains to the north and the colorado plateau to important biologically because of their role in the south maintaining critical ecological and evolution- because of both the high habitat diversity ary processes because of their broad eleva- and the central location of the book cliffs tional gradients extending from high peaks to tavaputsTavaputs region the biota is unusually diverse desert valley floors these ranges support a and compositionally unique and includes many wider variety of habitats and a greater diver- species at their distributional limits among sity of species than do areas of comparable 106 GREAT BASIN naturalist volume 56

size but less elevational relief this character- mojave desert inm southwestern utah istic also enables them to support the seasonal washington county includes utah s only migrations of animals from large anguungu ranging representative of the mojave desert a warm lates to small passerine birds furthermore desert commonly recognized by biogeogra these ranges have outstanding sciseisel mountain langes sci- ahersphers as lying between the great basin desert entific value because they represent cool and to the north and the sonoran desert to the islands in otherwise mesicmeslemesie habitat in an warm south shreve 1942 jaeger 1957 rowlands et al and landscape their natural communities have 1982 macmahon 1986 the mojave desert is developed through intermittent periods of physically part of the basin and range geol- extreme isolation grayson 1993 coupled with ogical province but it is characterized by rel- the geological diversity the great of region atively low elevation over most of its area 600 this isolation has led to the formation of to 1500 inm above sea level and by both limited plant assemblages often including rare unique precipitation 100 275 mm annually in most al al local endemicsendemics albee et 1988 welsh et places and warm summers 35 mean 1993 illustrating how populations and 40c by maxima for july see macmahon 1986 the communities of habitat islands are modified uniqueness of the physical environment of the through colonization and these extinction mojave is reflected in its biota characteristic have role mountain ranges played a major in plants include the joshua tree yucca brevifoliabrevifohdbrevifolia the development of theories of geographical creosote bush larrea tntritrldentata white bur ecology and biogeography brown 1971 1995 sage ambrosia dubosadudumosamosa brittle bush encelia grayson 1993 E rickart in preparation farfanfahfannorafannosafarinolafarinosajahnarinosanosa and several species of saltbush attiattlatri- portions of several isolated mountain ranges plex of these the joshua tree can be consid- are represented within PWAs on BLM lands ered endemic and if the distribution of this UWC 1990 such ranges include the henry species is used to define the boundaries of the mountains of the colorado plateau and the mojave desert then the desert covers a sub- deep creek fish springs house and new- stantial portion of southeastern california the foundlandfoundland ranges of utah s west deserts aixtigfiggix southern cone of nevada the northwestern 1 As the most isolated range in utah the and west central parts of arizona and the ex- newfoundland mountains in box elder county treme southwestern corner of utah are especially distinctive at 2129 in above sea judicious designation ofnew wilderness areas level desert peak and a considerable area of in this corner of the state could help to safe- surrounding uplands would have existed as an guard the many components of utah s biologi- island throughout the history of ancient lake cal diversity that are endemic to the mojave bonneville currently the range forms a 154 desert and the associated virgin mountains of km2 island of and to semiarid vegetation northwestern arizona and adjacent nevada immersed in a salt playa sea no doubt salt figure 4 details land ownership in this region marshes have covered the present salt flats of washington county because so much of periodically as the lake has advanced or this land is already in the public domain there receded in response to glacial and interglacialmterglacial is opportunity for diversitybiodiversitybio conservation climates the range has theithereforeebore been an eco- with minimal disruption of economic activity logical island thithl oughoutthroughout nearly 2 million years protected areas include zion national park a of pleistocene and quaternary time given such substantial wilderness in the pine valley long isolation these mountains have much to mountains of the dixie national forest no I1 teach scientists about the persistence local in fig 4 the upper virgin river desert wild- extinction vagivasilityvagilitylity and evolutionary dynamics life management area or DWMA a reserve of a variety of animal and plant species that for the desert tortoise copherusgopherusGopCop herusberusbefus agassizagassiznagassianii nos either live there now or have lived there in the 2aaa and ab2b in fig 4 the existing beaver dam past in utah and elsewhere in the intermoun- wilderness areas that extend into utah from tain legionregion knowledge of these topics will be arizona 4 and the lytle ranch preserve 5 important in the future as land managers try to although all of these protected areas play anticipate plant and animal responses to the important roles in conserving regional blodi increasing fragmentation and isolation of nat- versityhersityversity 2 of the largest areas zion national ural habitats within the human dominated park and designated forest service wilderness landscape brown 1995 in the pine valley mountains are generally z 1

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fig existing within lands arewaewau 108 GREAT BASIN naturalist volume 56 too high in elevation andor too far to the tree and beaver dam wash wilderness areas northeast to include many mojave desert would thus provide an economical way to aug- species the upper virgin river DWMA will ment conservation of tortoise populations con- protect lower elevation communities and will fined to the south facing slopes of the beaver include some mojave desert taxa however dam mountains many mojave desert species in utah do not of the 13 squamate reptiles listed in table 3 extend northeast of the beaver dam moun- nine are confined to either the mojave habitats tains and existing protected areas on the proper sites 3aaa ab3b 4 and 5 in fig 4 or to beaver dam slope are relatively small and iso- these sites plus the upper virgin river DWMA lated from each other fig 4 by virtue of both sites 2aaa and ab2b in fig 4 four species have size and location 2 PWAs the beaver dam more extensive distributions because they are wash and joshua tree units nos 3aaa and ab3b also recorded from zion national park among respectively in fig 4 see UWC 1990 could the 9 squamatedsquamsquamatesates with restricted distributions make important contributions to diversitybiodiversitybio the lizards heiHelhelodermaodenna suspectumsuspectum and xantusiaXantusia conservation in utah together these 2 units vigilis and the snakes crotalus cerastes and cover a range of elevations include several leptotyphlops humilis may occur at all 5 mojave distinctive plant communities not represented sites although this needs to be confirmed in the upper virgin river DWMA and are through additional fieldwork xantusiaXantusia vigilis close enough to one another and to the exist- also occurs further east in isolated populations ing protected areas to serve as stepping stones in garfield and san juan counties and previ- for animal movement ous molecular studies by bezy and sites 1987 we illustrate the conservation value of show deep genetic divisions among many iso- these 2 PWAs through an example the herpeto lated populations many of these isolates would fauna of the mojave desert includes 3 anu qualify as full species following the criteria of rans 1 tortoise 16 lizards 18 snakes and about davis and nixon 1992 but the specific status 28 additional species whose distributions are of the isolated utah populations remains un- peripheral but extend into this desert along known the lizard callisaurus draconoidesdraconoides one of its edges stewart 1994 the portion of occurs with certainty in the upper virgin river this fauna ranging into utah includes 2 anu DWMA in snow canyon state park beaver rans the turtle and 13 squamatedsquamsquamatesates 5 lizards dam wash PWA and lytle ranch preserve and 8 snakes their distributions across exist- sites 2aaa 3aaa and 5 in fig 4 the iguana dip- ing or proposed protected areas are summa- sosaurusso dorsalis is known confidently from rized in table 3 of this total the relict leop- only the lower beaver dam wash PWA ard frog rana onca apparently is extinct in although it may occur at low densities in the utah platz 1984 jennings and hayes 1994 other 3 mojave sites among the snakes cro- and therefore absent from all existing and pro- talus scutulascutulatustus is confined to the 4 strict posed protected areas in washington county mojave desert areas and C mitchelmitchelliimitchellislii is known the other anuran confined to this part of utah with certainty from only the higher elevation is the southwestern toad bufo micromicroscaphusscaphus mojave sites ab3b and 4 although the other 2 it is known to exist with certainty in several locations are possible based on a new snake areas and is likely widespread throughout the record for utah phyllorhynchus decurtatusdecurtatus is region where appropriate aquatic habitats known from a specimen BYU 45605 taken on exist table 3 11 july 1995 ca 151.5lsis mi N of the utah arizona the desert tortoise copgopcopherusgopherusCopGop herus agassizagassiziiii has border along the beaver dam slope road based been studied extensively over the past decade on this record the species likely occurs in the and intermittently for a much longer period of beaver dam wash and joshua tree areas aa3a time woodbury and hardy 1948 bury and and ab3b which are similar in vegetative struc- germano 1994 grover and defalco 1995 ture to the collecting site and possibly at the while utah populations have apparently de- other mojave desert sites as well regardless clined in the beaver dam slope area they of exact distributions all 9 squamate species persist at high densities north of st george with the most restricted distributions would data summarized in bury and germano 1994 benefit by wilderness designation of the pro- and are now protected in the virgin river posed beaver dam wash and joshua tree units DWMA protection of the proposed joshua UWC 1990 and for 7 species C draconoidesdraconoides 199619961 wilderness SELECTION FOR biodiversityBIODIVERSITY 109

TABLE 3 distribution of amphibians and reptiles restricted to southwestern utah relative to existing protected areas and beaver dam wash and joshua tree units of proposed BLM wilderness included in HHRR 1500 the areas numbered are shown in figure 421 the proposed red mountain and cottonwood canyon wilderness areas UWC 1990 are not illustrated because they are largely red mountain or entirely cottonwood canyon contained within the upper virgin river DWMA beaver dam dixie NFNE upper virgin wash joshua tree beaver dam lytle zion national wilderness river DWMA wilderness wilderness wilderness ranch taxon park 1 aa2a ab2b aa3a 313 4 5

ANURA rana onca bufo micromicroscaphusscaphus testudinesTESTUDINES GopCopcopherusgopherusherus agassiz ii

SQUAMATA callisaurus draconoidesdraconoides leonyxCocoleonyx varievariegatusgatus dipsosaurus dorsalis heloderma suspectsuspectumum xantusiaXanxantusiavigilistusia vigilis crotalus cerastes crotalus mitchelmitchelliimitchellffmitchellisliiilllffiff crotalus scutulascutulatustus leptotyphlops humilis masticophisMastic ophis flagellum phyllorhynchus decurtatusdecurtatus sonora semisemiannulataannulata trimorphodonTrimorph odon biscutatus distributions were inferred from locality records available in seaichsearchreresearch collections of california academy of sciences M L bean life sciences museum brigham young university provo utah museum of vertebrate zoology university of california berkeleybelBei keleskeley utah museum of naturalnatuial history university of utah salt lake city species listed as present if they 1 exist as museum voucher specimens 2 have been documented photographically but not collected because of threatened or endangered status or01 3 have been collected near a protected aleaarea and are known to occupy the appropriate habitat for example stewart 1994 summarizedsumman zed distributions of all mojave desert amphibians and reptiles on the basis of thentheir occurrence in distinct habitat types and we used these data as an indication of the likely plesencepresencepie sence of a species inin an area if not actually documented doubts about any occurrences aiedreare indicated by

D dorsalis the 3 species of crotalus L species furlow and armijo prewitt 1995 lesica humilis and P decurtatusdecutfatusdecurtatus these 2 PWAs would and allendorf 1995 lomolino and channell constitute the largest blocks of protected area 1995 designation of the beaver dam wash in the utah portions of their distributions and joshua tree PWAs as wilderness would the biological significance of the mojave provide an extremely economical preactiveproactiveproactive desert region could be illustrated with com- conservation strategy for many species parable examples involving native birds small mammals and vascular plants literally scores IMPACT OF ROADS ON PLANT of species are restricted to the low elevation AND ANIMAL communities joshua tree habitats on the southwestern slopes of the beaver dam mountains see behle et al by definition under the 1964 wilderness 1985 albee et al 1988 and zeveloff 1988 for act wilderness areas must be large at least recent species compilations although most 5000 acres and roadless because even some are on the periphery of their ranges it is increas- remote and pristine areas contain primitive ingly apparent that such peripheral popula- roads or tracks roadlessness is often an issue tions are critical to maintaining genetic diver- in debates over wilderness designation enviedvi sity and to ensuring the longtermlong term survival of ronmentalists tend to argue that the existence 110 GREAT BASIN naturalist volume 56

of minor roads or dirt tracks is not contradic- 12 in the depths of some naturally shallow tory to wilderness but that no new roads should lakes have already increased winter fish kills be built wilderness opponents respond that finally if efforts were made to reintroduce any road no matter how primitive disqualifies some of the large mammals considered above PWAs for wilderness status decision makers these efforts might be greatly facilitated by the may be pressured to make exceptions to allow protection of large blocks of roadless lands new roads and water development within that experience minimal human intrusion wildernesswildeinesselness boundaries heiehere we review the in summary if travel on minor roads and objective evidence bearing on the importance tracks were to be permanently restricted most of roadlessness from a purely biological per- but not all of the negative effects on wildlife spectivespective we deal with the effects of roads on would likely be ameliorated similar reasoning animals and plants independently would suggest that the effects of any new un- paved roads or tracks might be effects of roads on animals minor minimal if the roads were used briefly and sporadically roads affect wildlife in many ways both eg to carry communications equipment direct and indirect among the more com- effects of plant monly reported adverse impacts of roads on roads on communities animal populations are road mortalitiesmortalities animal the most compelling argument for large avoidance of roads isolation of populations by roadless areas is probably the protection of plant roads acting as barriers to animal movement communities from disturbances that can even- reductions in natural habitats increased poach- tually transform whole ecosystems through ing and elevated erosion leading to siltation of both direct and indirect effects roads tend to aquatic habitats on utah BLM lands large disrupt native communities of both microphytes mammals such as bighorn sheep ovis cana and macrophytes increased off road vehicle denbensisdensissis black bear and river otter are gener- traffic in roaded areas directly harms cryptobi ally intolerant of human disturbance and activ- otic soil crusts which play a key role in main- ities these and other mammals are known taining healthy ecosystems in semiarid and also to avoid habitat adjacent to roads oxley andaridarndannd lands and kills or injures plants and per- et al 1974 rost and bailey 1979 mader 1984 haps soil nesting insects like bees and wasps witmer and calesta 1985 van dyke et al indirect effects include the introduction of 1986 and can therefore be displaced by the nonnative pest plants which have gradually presence of roads historically humans in replaced many native species and drastically western north america have also persecuted a altered features of certain habitats the eco number of contemporary or former occupants systemwidesystem wide effects of these exoticsexotica are well of BLM lands such species include golden illustrated by asian tamarisk tamarix chinen and bald eagles gray wolf and grizzly bear sis which has channelized rivers and streams bortolottiBoi tolotti 1984 mechmeehmeeb 1995 in utah the in-in throughout the colorado drainage and thereby cidence of poaching is considerably higher in altered the characteristics flow regimes tem- regions adjacent to roads than in roadless areas peraperaturestures and sediment loads of both aquatic W woody UDWR personal communication and riparian habitats to the detriment of num- the negative effects of roads on wildlife can erous native fishes insects birds mammals generally be ameliorated by closing the roads and plants loope et al 1988 sudbrock 1993 to traffic road mortality and the advance of below we elaborate on the direct and indirect habitat alteration along roads should stop en- effects of roads on plant communities and on tirely and poaching should be sharply cur- the maintenance of both diversitybiodiversitybio and nat- tailed for larger animals roads would likely ural networks of interactions in utahutahs s native cease to act as barriers to animal movement ecosystems and gene flow however this might not be true THREATS TO cryptobioticCRYPTO BIOTIC SOILS across for some smaller species whose movements utah s andaridarld rangelands a collection of cyanoayano are more restricted generally significant ero- bacteria algae lichens and mosses form micro- sion and siltation of aquatic habitats might be phytic or cryptobioticcrypto biotic crusts on soil surfaces reduced only slightly siltation can be an impor- in pristine plant communities these crusts often tant consideration for example on the aquar- account for at least as much soil surface cover iuslus plateau where reductions by as much as as do vascular plants the cryptophytes provide 199619961 wilderness SELECTION FOR biodiversityBIODIVERSITY iiiili111 a number of valuable ecosystem services re- the many consumers depending directly or in- viewed in harper and marble 1988 west directly on these producers for food harper and 1990 and johansen 1993 including stabiliza- pendleton 1993 have suggested that destruction of soils against wind and water erosion tion of soil crusts and associated changes in enhancement of water retention and infiltra- forage quality may be related to a decline in tion brotherson and rushforth 1983 harper the health of desert tortoise populations in and st clair 1985 harper and marble 1988 southwestern utah grover and defalco 1995 and nitrogen fixation by autotrophic bacteria if that suggestion is supported by empirical including both free living and symbiotic cyanoayano evidence in the future then destruction of crusts bacteria eg snyder and wullstein 1973 west may account in part for the 10 million cost and skujins 1977 klubek and skujins 1980 to date T esque personal communication of terry and burns 1987 their contribution to the desert tortoise recovery program the nitrogen economy of these andaridarndannd ecosystems ROADS AS CORRIDORS FOR INVASIONS OF is substantive in southern utah grasslands and introduced SPECIES possibly the greatest cold deserts dominated by pinyon pine and adverse impact of roads on biological commu- juniper nitrogen fixation by crusts is demon- nities in utah is the aggravation of invasions of strably the dominant source of nitrogen for aggressive weeds along road corridors where vascular plants evans and ehleringer 1993 disturbance from road construction has elimi- the greater soil moisture and fertility associ- nated native competitors these introduced ated with biotic crusts have been shown to plants now form the dominant cover on many result in higher tissue nutrient levels belnap andaridarld and semiarid landscapes in western north and harper 1995 and references therein america and are widespread in utah mack higher seedling survivorship in associated vas- 1981 morrow and stahlman 1984 young et cular plants st clair et al 1984 harper and al 1987 papers in mcarthur et al 1990 and st clair 1985 belnap 1994 and greater aot monsen and kitchen 1994 habitat degrada- floristic diversity kleiner and harper 1972 tion by nonnativenormative congregating glazersgrazers un- herbivoresHerbivores and other consumers may benefit doubtedly aided the initial spread of brome indirectly from the enhanced nutrient status of grasses genus bromus and other european or these ecosystems harper and pendleton 1993 asian annuals into native habitats including belnap and harper 1995 grasslands previously dominated by caespitose growing recognition of the importance of or tussock grasses young and evans 1971 cryptocryptobioticbiotic crusts to ecosystem processes has loope 1976 mack 1981 1989 billings 1990 led to concern about the impact of disturbance 1994 brome grasses red brome B rubens by recreational users and nonnativenormative glazersgrazers on japanese brome B japojaponicusnicus downy brome such surfaces anderson et al 1982 johansen B mollismomsMOWS ripgut brome B diandrus and et al 1984 terry and burns 1987 cole 1991 especially cheatcheatgrassgrass B tectoriumtectorumtectorum have greatly evans and ehleringer 1993 belnap et al increased fire frequency from an average of 1994 belnap 1995 on most semiarid utah 60 110 yr to 5 yr in sagebrush steppe as lands a single pass of an off road vehicle will well as altered the pattern and dynamics of reduce nitrogen fixation by cyanobacteria and fires eg whisenantwbisenant 1990 invaded lands suf- increase wind and water erosion of surface fer declining productivity stewart and young soils williams et al 1995 estimates of time 1939 and watershed damage buckhouse 1985 to fallfullfuli recovery of disturbed biotic crusts includ- and become drastically depleted in both native ing nitrogen fixing capacity range up to 50 years plant species and cryptocryptobioticbiotic soil crusts young in the great basin or 100 years on the colorado and evans 1978 whisenant 1990 billings 1990 plateau J belnap personal communication 1994 rosentreter 1994 fig 5 treatments to the full biological and economic conse- restore these lands often involve introductions quences of disturbing biotic crusts remain to of still other exoticsexotica eg agropyron cristatumcristatum be quantified however in semiarid ecosys- kochia proprostrataprostratestrata see contributions to mcarthur tems where plant productivity is limited by et al 1990 and monsen and kitchen 1994 availability of water and nitrogen even small the influx of invading weedy annuals has reductions in these resources can be expected profound effects on genetic species and eco- to diminish primary productivity to the detri- system diversity although such effects remain ment of both the producers themselves and poorly documented in some parts of utah 112 GREAT BASIN naturalist volume 56

harner quadratsquadransQuadrats io10101.0 yd circumference to oararea ratios might protect andaridanidannd 124 quadratsQuadquadransrats and semiarid western ecosystems against whole- 12 sale habitat conversion exotic weeds tend to total species invade native plant communities mainly along roadsides railroad right of ways and other 10 highly disturbed sites forcella and harvey 1983 hunter 1990 literature cited in billings 1990 1994 also bergelson al 1993 8- and see et favorably wet drainage ditches provide inroads to new habitat and invaders spread outward from the ditches during particularly 6 wet years although systematic surveys of nonnativesnonnatives do no of natives samplesampie size not presently exist for PWAs and are sorely

13 5 16 3 1 37 16 4 2 needed is 4 there evidence that invasions of 4 6 8 exotic weeds may be prevented by restricting no aliensquadrataliens quadrat access on existing roads thus of the replicate roadsides studied by hunter 1990 introduced fig 5 relationship of both total species richness and species including not only brome grasses but numbers of native species per quadrat to the number of erodium cicutarium salsola sppapp and sisym- individuals of introduced per quadrat plotted species brium altissimum all from raw data in harner and harper 1973 data are dominated but the one flomfrom sagebrush grasslands on private and BLM foothill that had been closed to traffic and left undis- lands in salt lake davis and thoeletooele counties turbed for many years prior to censuringcensusingcensusing the effects of roads on plant communities brome grasses form virtual monomonoculturescultures en- appear to differ importantly from those on animal tirely replacing native communities especially communities construction of new roads in wet years eg pellant and hall 1994 and especially those with drainage ditches may authors observations in other western states hasten longtermlong term and permanent changes to local brome grass invasions threaten state or feder- floras and these changes may eventually effects ally listed plant species rosentreter 1994 have markedly adverse on whole eco- california native plant society personal com- systems existing dirt tracks are probably less municationmunication effects of habitat conversion threatening to plant communities although radiate upward through the food chain and moisture conditions on the tracks may be as favorable as drainage adverse effects have been documented on here in ditches soil com- pronghorn antilocapra americana and deer paction appears to retard growth of most plants given costliness pellant 1990 roberts 1994 small vertebrate the of aggressive fire sup- vail 1994 and habitat prey of eagles and other raptorsraptores kochert and pression eg restoration al pellant 1986 nydegger and smith 1986 native measures see reports in mcarthur et 1990 and monsen 1994 birds dobler 1994 and insects fielding and and kitchen the most economical strategy for the of brusven 1994 As summarized by billings preventing spread introduced still 1994 exotic annual grasses could constitute grasses to areas that are relatively pristine may be to maintain their road- a genuine threat to the existence of large inte- less character this also would provide oppor- grated ecosystems that have existed since the tunitunitiesties for investigating the effects of roads pleistocene in the relatively andaridarld lands between or lack thereof on the advance of the rocky mountains and sierra nevada these exotic operational ecosystems could disappear over large plants on andaridarld lands in utah areas of thousands of square kilometers conclusions A very high priority for future ecological work in utah will be to determine the extent wilderness serves many purposes and its to which the remote BLM lands being consid- designation involves many and varied considered for wilderness status might serve as ref- erationserations the technical issues and evidence uges for native flora and fauna seeds of brome presented here demonstrate that BLM wilder- grass dispersed by animal vectors certainly ness lands can play a major and perhaps pre- travel over long distances and into wilderness dominant role in safeguarding genetic species areas however large roadless areas with low and ecosystem diversity across much of andaridarld 199611996 wilderness SELECTION FOR biodiversityBIODIVERSITY 113 utah over the long term large contiguous ATWOOD D J HOLLAND R BOLANDER B FRANKLIN D E networks of wilderness and other protected HOUSE L ARMSTRONG K THORNE AND L lands can provide for ENGLAND 1991 utah threatened endangered and sanctuary populations of sensitive plant field guide animals with large area requirements and can AXLEROD D I1 1960 the evolution of flowering plants help maintain natural processes and interac- pages 227 305 in S tax editor evolution after dar- tions that sustain healthy biotic communities winwm volume 1 the evolution of life university of chicago press in many situations wilderness designation can chicago BABBITT B 1995 science opening the next chapter of provide low cost protection for rare and en- conservation history science 267 1954 1956 dangereddangered species BLM lands in geographi- BEHLE W H E D SORENSEN AND C M WHITE 1985 cally diverse regions of utah all offer unique utah birds a revised checklist occasional publica- ecological scientific and educational values tions no 4 utah museum of natural history university of utah salt lake city an extent so far to unmeasured wilderness BEHNKE R J 1992 native trout of western north amer- lands may protect native ecosystems from icaiealea american fisheries society monograph 6 275 wholesale transformation by invasions of exotic PP species clearly if biological considerations BEHNKE R J AND M ZARN 1976 biology and management and are taken into account in wilderness decisions of threatened endangered western trouts USDA forest service technical report RMGTRRM GTR wilderness can play a critical role in the long- 28 fort collins CO term preservation of utah s biological heritage BELNAP J 1994 potential value of cyanobactenalcyanobacterial inocula- tion in revegetation efforts pages 179 185 in S B acknowledgments monsen and S G kitchen editors proceedings ecology and management of annual rangelands report INT GTR 313 ogden UT jayne belnap phyllis coley donald duff 1995 soil surface disturbances thentheir role in sharon emerson donald feener bruce how- accelerating desertificationdecertification environmental moni- lett thomas kursar and samuel rushforth toring and assessment 37 1 17 contributed to an earlier nontechnical version BELNAP J AND K T HARPER 1995 the influence ofcrypof crapcryp tobiotic soil crusts on elemental content of of this paper in addition to these individuals tissue in two desert seed plants andaridarld soil researchReseaich and and the authors patricia berger lynn bohs rehabilitation 9 107 115 rex catesgates steven clark susan fairbanks jer BELNAP J K T HARPER AND S D WARRENWARRLN 1994 sur- ran flinders sarah george james harris face disturbance of cryptocryptobioticbiotic soil crusts nitrogenitrogen richard hildreth carl marti brian maurer nase activity chlorophyll content and chlorophyll norman negus rogers seger degradation andaridarld soil researchReseaichsealch and rehabilitation duke jon john 818 1 8 sperry richard tolman delbert wiens BELOVSKY G E 1987 extinction models and mammalian michael windham and samuel zeveloff en- persistence pages 35 57 in M E soulesouie editor dorsed the viable populations for conservation cambridge uni- earlier paper klancy de nevers press and jon seger rescued corrupted versity cambridge UK computer BELOVSKY G E J A bissonette R D DUESER T C files griswold parker T F and V tepedino EDWARDS JRJRCC M LUECKE M E RITCHIE J B allowed us to use their unpublished data table SLADE AND F H WAGNER 1994 management of 2 and section on bees J belnap and L shultz small populations concepts affecting the recovery of commented a endangered species wildlife society bulletin 22 on previous draft of the manu- 307 316 and garrison script carla helped us obtain BERGELSON J J A NEWMAN AND E M floresrouxFLORES ROUX BLM maps 1993 rates of weed spread in spatially heteroge- neous environments ecology 74 999 1011 BEZY R L AND W SITES JR 1987 A preliminary study CITED J literature of allozymeallozyme evolution in the lizard family xantusipantusi idae reptilia sauriasauna Herpetologicherpetologicaherpetologicala 43 280 292 ALBEE B L M SCHULTZ S GOODRICH J AND 1988 BILLINGS W D 1990 bromus tectoriumtectommtectorumtectommtorum a biotic cause of atlas of the vascular pub- plants of utah occasional ecosystem impoverishment in the great basin licationslications no 7 museum utah of natural history pages 301 322 in G M woodell editor the earth in university of salt 670 utah lake city appp transition patterns and processes of biotic impoveiimpoverimprover ANDERSON D C K T HARPER AND S R RUSHFORTH ishmentashment cambridge university press cambridge 1982 recovery of cryptogamic soil crusts from graz- UK ing on utah winter ranges journal of range man- 1994 ecological impacts of cheatcheatgrassgrass and resul- agement 35 355 359 tant fire on ecosystems in the western great basin armbruster P AND R LANDE 1993 A population viability pages 22 30 inm S B monsen and S G kitchen analysis for african elephant loxodonta africana editors proceedings ecology and management of how big should reserves be conservation biology annual rangelands USDA forest service technical 7 602 610 report INT GTR 313 ogden UT 114 GREAT BASIN naturalist volume 56

bolcerBOLGIRBOLGER D T A C ALBERTS AND M E SOULESOULL 1991 DURRANT S D 1950 mammals of utah taxonomy and occurrence patterns of bird species in habitat frag- distribution museum of natural history university ments sampling extinction and nested species sub- ofkansasof kansas publications 6 sets american naturalist 137 155 166 EDMUNDS G F 1994 tail water insects the landing net bortolottiBoRroloaoLom G BR 1984 trap and poison mortality of summer 6 7 golden and bald eagles journal of wildlife man- 1995 tail water insects revisited the landing agement 48 1173 1179 net fall 4 11 brotherson J DANDSD AND S BR rushforth 1983 influ- EVANS R D AND J R ehleringerEHLERingerINGEB 1993 A break in the ence of cryptogamic crusts on moisture relationships nitrogen cycle in andandlandsaridarldaridlandssandlandslands evidence from on of of soils in navajo national monument arizona soils oecologiaoecologia9494 314 317 great basin naturalist 43 73 78 FIELDING D J AND M A BRUSVEN 1994 grasshopper BROWN J H 1971 mammals on mountainmountaintopstops nonequi- community responses to shrub loss annual grass- librium insular biogeography american naturalist lands and crested wheatwheatgrassgrass seedings manage- 105467105 467 478 ment implications pages 162 166 in S B monsen 1995 maclomacroecologymacioecologyMacioMacromaero ecology university of chicago press and S G kitchen editors proceedings ecology chicago and management of annualofannual rangelands USDA for- BROWN J H AND E J HESKE 1990 control of a desert est service technical report INT GTR 313 ogden grassland transition by a keystone rodent guild sci- UT ence 250 1705 1707 FORCELLA FE AND S J HARVEY 1983 eurasian weed BUCKHOUSE J C 1985 effects of fire on rangeland water- infestation in western montana in relation to vegeta- sheds pages 58 60 in K sandeisandelsanders s and J durham edition and disturbance madronoMadro fiogio 30 102 109 tors proceedings of a symposium rangelandranBangeland fire FRANKEL 0 H AND M E SOULESOULS 1981 conservation effects USU S department of the interior bureau of and evolution cambridge university press cam- land management boise ID bridge UK BURY BR B AND D J GERMANO EDITORS 1994 biology FURLOW FE B AND T ARMIJO PREWITT 1995 peripheral of north american tortoises USDI fish and wild- populations and range collapse conservation biol- life researchBeseaich report ISSN 104024111040 2411 no 13 204 ogy 9 1345 PP GRAYSON D K 1993 the desert s past a natural prehis- gaskCASECASLgase T J AND M L CODY 1988 testing theories of tory of the great basin smithsonian institution island biogeography american scientist 75 402 411 press washington DC christensenchrlsCHRIS 1 ensiLNSI N N lelalleralL ET AL 1989 interpreting the yellow- GROVER M C AND L A DEFALCO 1995 desert tortoise stone fires of 1988 biosciencebioscience3939 678 685 copgopcopherusgopherusCopGop herus agassizagassiziiagassizziziii status of knowledge outline goleCOLECOLL D N 1991 trampling disturbance and recovery of with references USDA forest service technical cryptogamic soil crusts in grand canyon national report INT GTR 316 ogden UT 134 appp park great basin naturalist 50 321 325 GRUMBINE R E 1994 what is ecosystem management cronquistCRONQUISI A A H HOLMGREN N H HOLMGREN AND conservation biology 8 27 38 J L REVEALBEVCAL 1972 intermountain floiafiolaflora volume 1 HARNER R F AND K T HARPER 1973 mineral composi- hafner publishers new york tion of grassland species of the eastern great basin CROSSgross S P 1985 responses of small mammals to forest in relation to stand productivity canadian journal of perturbations pages 269 275 min riparian ecosys- botany 51 2037 2046 tems and thentheir management reconciling conflicting HARPER K T 1986 historical environments pages 516351 63 uses USDA foiestforest serviceSeivice technical report RM- in W L dazevedo editor handbook of north GTR 120 foitcolhnscofort collins CO american indians volume 11 great basin smith- DAVIDSON D W R S INOUYE AND J H BROWN 1984 sonian institution washington DC giamvoiygranivoreGrangranivoryivory in a desert ecosystem experimental evi- HARPER K TANDJT AND J R MARBLE 1988 A role for nonvas- dence foifol indirect facilitation of ants by rodents cular plants in management of and and semiarid ecology 65 1780 1786 rangelands pages 135 169 in P T tuellerthellertheiler editor DAVIS J I1 AND K C NIXON 1992 populations genetic vegetation science applications for rangeland analy- variationvanation and the delimitation of phylogenetic sis and management kluwer academic publisher species systematic biology 41 421 435 dordrechtDordrecht DEACONDLACON J E AND W L MINCKLEY 1974 desert fishes HARPER K T AND R L PENDLETON 1993 cyanobacte pages 385 488 inCT G W brown editor desert biol- riana and cyanolichenscyanolicbens can they enhance availability ogy volume II11 academic press new york of essential minerals for higher plants great basin department OFTHEor THE INTERIOR BUREAU OF LAND manage- naturalist 53 59 72 ment 1994 ecosystem management in the BLM HARPER K T AND L L ST CLAIR 1985 cryptogamic fromflomelom concept to commitment US government soil crusts on and and semiarid rangelands in utah printing office 573183573 183 washington DC 16 appp effects on seedling establishment and soil stability DIAMOND J M 1984 normal extinctionsextinct ions of isolated final report bureau of land management utah populations pages 191 246 in M H nitecki editor state office salt lake city extinctionsExtinct ions university of chicago press chicago HARRIS L D 1984 the fragmented forest island bio- DOBLERDOBLLR F C 1994 washington state shrub steppe eco- geography theory and the preservation of biotic system studies with emphasis on the relationship diversity university of chicago press chicago between nongame budsbirds and shrub and grass cover HEANEY L R 1984 mammalian species richness on densities pages 149 161 in S B monsen and S G islands on the sunda shelf southeast asia decolooecolo kitchen editorsediedl toistols proceedingspioceedmgs ecology and man- gia berlin 61 11 17 agement of annual rangelands USDA forest ser- HOLDEN P B R STONE W WHITE G somerville D vice tecltecitechnicalTeel micalmicai report INT GTR 313 ogden UT DUFF R GERVAIS AND S GLOSS 1974 threatened 199611996 wilderness SELECTION FOR biodiversityBIODIVERSITY 115

fishes of utah proceedings of the utah academy of LESICA P AND F W ALLENDORFALLENDORE 1995 when are periph- science arts and letters 51 46 65 eral populations valuable for conservation conser- HUBBS C L 1940 speciation of fishes american natu- vation biology 9 753 760 ralist 74 198 211 LOMOLINO M V AND R CHANNELL 1995 splendid iso-iso 1941 the relation of hydrological conditions to lation patterns of the geographic range collapse in speciation inm fishes pages 182 195 in J G need- endangered mammals journal of mammalogy 76 ham editor A symposium on hydrobiology univer- 335 347 sity of wisconsinofwisconsin press madison LOOPE L L P G SANCHEZ P W TARR W L loopeLOOPL HUNTER R 1990 recent increases in bromus on the AND R L ANDERSON 1988 biological invasions of nevada test site pages 22 25 in E D mcarthur and land nature reserves biological conservation E M romney S D smith P T tuellertheller editors 449544 95 118 proceedings symposium on Cheatcheatgiasscheatgrassgrass invasion LOOPE W L 1976 relationships ofvegetationof vegetation to enviedvienvi- shrub die off and other aspects of shrub biology ronmentronment canyonlandsCanyonlands national park utah unpub- and management USDA forest service technical lished doctoral dissertation utah state university report INT GTR 276 ogden UT logan IUCN 1978 categories objectives and criteria for pro- MACK R N 1981 the invasion of bromus tectoriumtectorumtectorum L tected areas moimolmorgesges switzerland into western north americaamenea an ecological chronicle JAEGER E C 1957 the north american deserts stan- agro ecosystems 7 145 165 ford university press palo alto CA 308 appp 1989 temperate grasslands vulnerable to plant JENNINGS M R AND M P HAYES 1994 decline of native invasions characteristics and consequences pages ranid frogs min the desert southwest pages 183 211 155 179 in J A drake H A mooney F di castri in P R brown and J W wright editors herpetol- R H groves FE J kruger M bejrejmanekrejmdnekRej manek and M ogy of the north american deserts southwestern williamson editors biological invasions john wiley herpetological society special publication no 5 and sons new york van nuys CA 311 appp MACKINNON J K MACKINNON G CHILD AND J THOR- JOHANSEN J R 1993 cryptogamic crusts of semiarid and SELL 1986 managing protected areas in the tropics and lands of north america journal of phycology IUCN gland switzerland 2914029 140 147 MACMAHON J A 1986 deserts audubon society nature JOHANSEN J R L L ST CLAIR B L WEBB AND G T guides alfred knopf new york 638 appp NEBEKER 1984 recovery patterns of cryptocryptogamicgamiegamic MADER H J 1984 animal habitat isolation by roads and soil crusts in desert rangelands following fire distur- agricultural fields biological conservation 29 bance bryologist 87 238 243 81 96 JOHNSON R R L T HAIGHT AND J M SIMPSON 1977 MCARTHUR E D E M ROMNEY S D SMITH P T endangered species vs endangered habitats a con- TUELLER EDITORS 1990 proceedings symposium cept pages 68 79 in importance preservation and on Cheatcheatgrassgrass invasion shrub die off and other management of riparian habitat USDA forest ser- aspects of shrub biology and management USDAUS DA vice technical report RM GTR 43 fort collins CO forest service technical report INT GTR 276 JULANDER 0 1962 range management in relation to ogden UT mule deer habitat and herd productivity in utah MECH L D 1995 the challenge and opportunity of journal of range management 15 278 281 recovering wolf populations conservation biology KAY C E 1990 yellowstone s northern elk herd a critical 9 270 274 s evaluation of the natural regulation paradigm MEFFE G K AND C R CARROLL 1994 principles of con- unpublished doctoral dissertation utah state uni- servation biology sinauer sunderland MA 600 appp versityversity logan MICHENER C D 1979 biogeography of the bees annals KERSHNERKERSHNEK J L 1995 bonneville cutthroat trout pages of the missouri botanical garden 66 277 347 28 35 in M K young editor conservation assess- MILLER R R 1959 origin and affinities of the freshwater ment for inland cutthroat trout USDA forest ser- fish fauna of western north america pages 187 222 vice technical report RM GTR 256 fort collins in C L hubbs editor zoogeography AAAS publi- CO cation 51 KLEINER E F AND K T HARPER 1972 environment and 1961 man and the changing fish fauna of the community organization in grasslands of canyon american southwest papers michigan academy of lands national park ecology 53 299 309 science arts and letters 46 365 404 KLUBEK B AND J SKUJINS 1980 heterotrophic nitrogen MINCKLEY W L AND J E DEACON 1968 southwestern fixation in and soils crusts soil biology and bio- fishes and the enigma of endangered species sci- chemistry 12 229 236 ence 159 1424 1432 KNOPF F L 1985 significance of riparian vegetation to MINCKLEY W L AND J E DEACON EDITORS 1990 battles breeding birds across an altitudinal dinehinecline pages against extinction native fish management in the 105 111 in riparian ecosystems and their manage- american west university ofarizonaof arizona press tucson ment reconciling conflicting uses USDA forest ser- MINCKLEY W L D A hendrickson AND C E BOND vice technical report RM GTR 120 fort collins 1986 geography of western north american fresh- CO water fishes description and relationships to intra- KOCHERT M N AND M PELLANT 1986 multiple use in continentalcontmental tectonismteutonismtectomsm pages 519 613 in C H the snake river birds of prey area rangelands 8 hocutt and E 0 wiley editors the zoogeography 217 220 of north amerleanamerican freshwater fishes john wiley & KUSHLAN J A 1979 design and management of conti- sons new york nental wildlife reserves lessons from the everglades MONSEN S B AND S G KITCHEN EDITORS 1994 pro- biological conservation 15 281 290 ceceedings ecology and management of annual 116 GREAT BASIN naturalist volume 56

rangelands USDA forest serviceSeivice technical report PIMM S L H L JONES AND J DIAMOND 1988 on the INT GTR 313 ogden UT risk of extinction american naturalist 132 757 785 MORROW L A AND P W STAIILMAN 1984 the history PLATZ J 1984 status report for rana onca cope pre- and distributiondistl ibution of downy brome bromus tectoriumtectorumtectorum pared for office of endangered species USU S fish in north americaamenea weed science 32 supplement 1 and wildlife service albuquerque NM 27 appp 2 7 PRIMACK R B 1993 essentials of conservation biology moylMOYIMOYLE L P B AND J E WILLIAMS 1990 Biodiversity loss sinauer sunderland MA 561 appp in the temperate zone decline of the native fish RAVEN P R 1992 the nature and value of bio diversity fauna of californiacalifoicalifor nia conservation biology 4 275 284 pages 1 18 in global diversitybiodiversitybio strategy guide- NATIONAL RESEARCHrl SLARLII COUNCIL 1978 conservation of lines for action to save study and use earth s biotic geigelgermgermplasmplasm resourcesrisouiresoui eesces an imperative national acad- wealth sustainably and equitably WRI IUCN emy of sciences washington DC UNEP nefrigefrinl 11 J L AND B B SIMPSON 1993 bees pollination REID W V AND K R MILLER 1989 keeping options systems and plant diversity pages 143 167 in J alive the scientific basis of conserving biodiversity lasalle and I1 E gauld editorsediedltois hymenoptera and world resources institute washington DC diversitybiodiversitybiodiveisitybioblo CAB intelinternationalnational wallingford UK RICKART E A in preparation elevational diversity gradi- NIWMARKNEWMARK W D 1985 legal and biotic boundaries of ents biogeography and the structure of montane westeinwestern northnor th american national parks a problem mammal communities in the intermountain region of congruence biological conservation 33 197 208 journal of biogeography in preparation 1987 A land bridge island perspective on mam- ROBERTS T C JR 1994 resource impacts of cheatcheatgrassgrass malian extinctionsextinctions in western north american parks and wildwildfiresfires on public lands and livestock grazing nature 325 430 432 pages 167 169 in S B monsen and S G kitchen 1991 tropical forest fragmentation and the local editors proceedings ecology and management of extinction of understory budsbirds in the eastern usam annual rangelands USDA forest service technical baiabalabara mountains tanzania conservation biology 5 report INT GTR 313 ogden UT 67 78 rosentreterROSENTreterKETER R 1994 displacement of rare plants by 1995 extinction of mammal populations in west- exotic grasses pages 170 175 min S B monsen and ern northnoi th american national parks conservation S G kitchen editors proceedings ecology and biology 9 512 526 management of annual rangelands USDA forest NOSS R FE 1992 the wildlandswildlands project land conservation service technical report INT GTR 313 ogden strategy pages 10 25 inOT the wildlandswildlands project wild UT eartheaieal th special issue cenozoic society ROST G R AND J A BAILEY 1979 distribution of mule NYDEGGERNYDLGGLB N C AND G W SMITH 1986 pages 152 156 deer and elk in relation to roads journal of wildlife in D E mcarthur and B L welch editors pro- management 43 634 641 ceccceedings symposium on the biology of artemisiaofartemisia ROWLANDS P H JOHNSON E RITTER AND A ENDO and chrysothamnus USDA forest service techni- 1982 the mojave desert pages 103 162 in G L cal report INT GTR 200 ogden UT bender editor reference handbook on the deserts oxioxlOXLEYLY D J M B pentonFENTONFLNTON AND G R CARMODY 1974 of northnoi th america greenwood press westwood CT the effects of i oadsroads on populations of small mam- 594 appp mals journaljoinjoun nalnai ofappliedofapplied ecology 11 51 59 SALWASSERSALWASSFR H C schonewald COX AND R BAKERBAKEB PARKERPAHKLR F D 1985 A candidate legume pollinator osmiacosmia 1987 the role of interagency cooperation in manag- sanrafaelae parker journal of apiculture research ing viable populations pages 159 173 in M E 2413224 132 136 soulesouie editor viable populations for conservation press 1986 field studies with osmiacosmia sanrafaelae parker cambridge university cambridge UK a pollinatorpoll matoi of alfalfa hymenoptera megachilidae SAMSON D A T E PHILIPPI AND D W DAVIDSON 1992 journal of economic entomology 79 384 386 GrangranivorygranivoreGramgramvoryivoryvory and competition as determinants of annual diversity the PAHLRSONPATTERSON B D 1984 mammalian extinction and bio- plant in chihuahuanchihuabuan desert oikosbikos 65 61 80 geographygeogiaphy in the southernsouthein rocky mountains pages SAUNDERS D A R J HOBBS AND C R MARGULES 1991 247 293 in M H nitecki editor extinctionsExtinctions uni- biological ecosystem versity of chicago pipressess chicago consequences of fragmentation a review conservation biology 5 18 32 PELLApellaniPLLLANINr M 1990 the cheatgrasscheat grass wildfire cycle are schonewald COX C M 1983 guidelines to manage- there any solutions pages 11 18 in E D mcarthur ment a beginning attempt pages 414 445 in C M E M romney S D smith P T tuellertheller editors schonewald S M chambers B macbryde and L proceedings symposium on cheatgrassCheatgrass invasion thomas editors genetics and conservation benja- shrub ofe and aspects of biology diedle off other shrub min cummings menlo park CA and management USDA forest technical service SHAFFER M L 1981 minimum population size for report INT GTR 276 ogden UT species conservation bioscience 31 131 134 PELLANTPLLLANT M AND C HALL 1994 distribution of two SHIOZAWA D K AND R P EVANS 1994 the use ofofdnaDNA pages ofena exotic glassesgrasses on intermountain rangelands to identify geographical isolation in trout stocks 109 112 in S B monsen and S G kitchen editors pages 125 131 in R barnhart B shake and R H proceedings ecology and management of annual hamre editors wild trout V wild trout inm the 21st rangelands USDA forestfoiest service technical report century INT GTR 313 ogden UT SHREVES H REVE F 1942 the desert vegetation of north america pickettPICKLTIPIGKETT S T A AND J N THOMPSON 1978 patch botanical reviews 8 195 246 dynamics and the design of nature reserves biologi- SHULTZ L M 1993 patterns of endemism in the utah cal conservation 13 27 37 flora pages 249 263 in R sivinski and K lightfoot 199611996 wilderness SELECTION FOR biodiversityBIODIVERSITY 117

editors southwestern rare and endangered plants TYUS H M AND C A KARP 1989 habitat use and new mexico department of forestry and resources streamstreamflowflow needs of rare and endangered fishes conservation division miscellaneous publication yampa river colorado USU S fish and wildlife ser- no 2 santa fe NM vicevlee biological report 8914 27 appp SHULTZ L M E E NEELY AND J S TUHY 1987 flora of USU S FISH AND WILDLIFE SERVICE 1993 endangered and the orange cliffs of utah great basin naturalist 47 threatened wildlife and plants 50 CFR 171117.1117 11 and 287 298 171217 12 august 23 1993 division of endangered SMITH G R 1966 distribution and evolution of the species USU S fish and wildlife service washington north american catostomid fishes of the subgenus DC Pantospantosteusteus genus catostomus miscellaneous publi- UTAH DIVISION OF WILDLIFE RESOURCES 1992 native cations of the museum of zoology university of utah wildlife species of special concern draft docu- michigan no 129 ment utah division of wildlife resources salt lake 1978 biogeography of intermountain fishes inter- city 31 appp mountain biogeography a symposium great basin UTAH wilderness COALITION UWC 1990 wilderness naturalist memoirs 2 17 42 at the edge peregrine smith books layton UT SNYDER J M AND L H WULLSTEIN 1973 the role of VAIL D 1994 symposium introduction management of desert cryptocryptogamscryptogramsgams in nitrogen fixation american semiarid rangelands impacts of annual weeds on midland naturalist 90 257 265 resource values pages 3 4 in S B monsen and S G soultSOULE M E EDITOR 1987 viable populations for con- kitchen editors proceedings ecology and man- servation cambridge university press cambridge agement of annual rangelands USDA forest ser- UK vicevlee technical report INT GTR 313 ogden UT soultSOULE M E D T BOLGER A C ALBERTS R SAUVAJOT VAN DYKE F G R H BROCKE H G SHAW B B ACKER- J WRIGHT M SORICE AND S HILL 1988 recon- MAN T P HEMKER AND F G LINDZEY 1986 react- strucstructedted dynamics of rapid extinctionsextinctions of chaparral ions of mountain lions to logging and human activ- requiring birds on urban habitat islands conserva- ity journal of wildlife management 50 95 102 tion biology 2 75 92 WAGNER F H R FORESTA R B GILL D R mccul- SOULESOULS M E AND B A WILCOX 1980 conservation lough M R PELTON W F PORTER AND consulta- biology an evolutionary ecological perspective sin- tion ON LAW AND POLICY BY J L SAX 1995 wildlife auer sunderland MA policies in the USU S national parks island press 242 ST CLAIR L L B L WEBB J R JOHANSEN AND G T PP NEBEKER 1984 cryptogamic soil crusts enhance- WARREN M LLANDBAND B M BURR 1994 status of freshwater ment of seedling establishment in disturbed and fishes of the united states overview of an imperiled undisturbed areas reclamation and revegetation fauna fisheries 19 6 18 research 3 129 136 WELSH S L 1978 endangered and threatened plants of STEBBINS G L JR 1952 aridity as a stimulus to plant utah a reevaluation great basin naturalist 38 1 18 evolution american naturalist 86 33 44 WELSH S L N D ATWOOD S GOODRICH AND L C STEBBINS R C 1985 A field guide to western reptiles HIGGINS 1993 A utah flora and2nd edition print ser- and amphibians houghton mifflin co boston MA vices brigham young university provo UT STEVENS G C 1992 the elevational gradient in altitudi- WELSH S L N D ATWOOD AND J L REVEAL 1975 nal range an extension of rapoport s latitudinal rule endangered threatened extinct endemic and rare to altitude american naturalist 140 893 911 or restricted utah vascular plants great basin natu- STEWART G R 1994 an overview of the mohave desert ralist 35 327 326 and its herpetofauna pages 55 69 in P R brown WEST N E 1990 structure and function of microphytic and J W wrghtwnghtwright editors herpetology of the north soil crusts in wildlandwildwindlandland ecosystems of and to semiarid american deserts southwestern herpetological soci- regions advances in ecological research 20 ety special publication no 5 van nuys CA 311 appp 179 223 STEWART G AND A E YOUNG 1939 the hazard of basing WEST N E AND J SKUJINS 1977 the nitrogen cycle in permanent grazing capacity on bromus tectoriumtectorumtectorum north american cold winter semisemidesertdesert ecosys- american society ofagronomy journal 31 1002 1015 tems oecologia Planplantarumplantariumtarum 12 45 53 SUDBROCKSUDBBOCK A 1993 tamarisk control I1 fighting back an WHISENANT S G 1990 changing fire frequencies on overview of the invasion and a low impact way of idaho s snake river plains ecological and manage- fighting it restoration and management notes 11 ment implications pages 4 10 in E D mcarthur 31 34 E M romney S D smith P T tueller editors TEPEDINO V J 1981 the pollination efficiency of the proceedings symposium on cheatgrassCheatgrass invasion squash bee peponakispeponapisPeponapis pruinosepruinosapruinosa and the honeybee shrub die off and other aspects of shrub biology apis mellifera on summer squash cucurbita pepo and management USDA forest service technical journal of the kansas entomological society 54 report INT GTR 276 ogden UT 359 377 WHITTAKER R H 1972 evolution and measurement of TERBORGH J AND B WINTER 1980 some causes of species diversity taxon 21 213 251 extinction pages 119 134 in M E soulesouie and B A WILCOVE D S C H MCLELLAN AND A P DOBSON wilcox editors conservation biology sinauer sun- 1986 habitat fragmentation in the temperate zone derland MA pages 237 256 in M E soulesouie editor conservation TERRY R E AND S J BURNS 1987 nitrogen fixation in biology the science of scarcity and diversity sinauer cryptogamic soil crusts as affected by disturbance associates sunderland MA pages 369 372 in proceedings pinyon juniperjumper WILCOX B A AND D D MURPHY 1985 conservation conference USDA forest service technical report strategy the effects of fragmentation on extinction INT GTR 215 ogden UT american naturalist 125 879 887 118 GREAT BASIN naturalist volume 56

WILLIAMS J D J P dobrowolski N E weslWESIWEST AND D A river at the ouray national wildlife refuge uinta cilletteGILLLTIEGILLETTE 1995 microphytic crust influence on wind county utah great basin naturalist 55 213 224 erosioneiosion transactions of the american society of WOODBUWOODBURYllykryiRy A M AND R HARDY 1948 studies on the agricultural engineers 38 131 137 desert tortoise CopcopherusgopherusGopherus agassizagassiziiagassiznagassianii ecological mono- willlamsWILLIAMSwllwil LIAMS J E 1991 preserves and refuges for native graphs 18 145 200 western fishes history and management pages YOUNG J AAANDRAND R A EVANS 1971 invasion of medusa 171 189 in W L minckley and J E deacon edi- head into the great basin weed science 18 89 97 tors battle against extinction native fish manage- 1978 population dynamics after wildwildfiresfires in sage- ment in the american west university of arizona brush grasslands journal of range management 31 press tucson 517 appp 283 289 wiluswilmswllwil 1 IS E 0 1974 populations and local extinctionsextinctions of YOUNG J A R A EVANS AND B L KAY 1987 cheatgrassCheatgrass birds on barro colorado island panama ecological rangelands 9 266 270 monographsmonogiaphs4444 153 169 ZEVELOFF S I1 1988 mammals of the intermountain west WILSON E 0 1988 diversitybiodiversityBio national academy university of utah presspi ess salt lake city press washington DC WITMERWITMLR G WwandoWANDDAND D S CALESTA 1985 effect of forest received 27 september 1995 roads on habitat use by roosevelt elk northwest accepted 21 march 1996 science 59 122 125 WOLZ E R AND D K SmSHIOZAWAozAwA 1995 soft sediment benthic macromacroinvertebrateinvertebrate communities of the green great basin naturalist 562 C 1996 appp 119 128 NUTRIENT distribution IN QUERCUS CAMBELIIGAMBELII STANDS IN CENTRAL UTAH

A R tiedemannlTiedemanntiedemann1Tiedemannlanni1 and W P claryaclary2

ABSTRACT gambel oak quercus gambelligambeliigambelii nutt is increasingly recognized as a valuable fuelwood throughout ari-an- zona colorado new mexico and utah knowledge of the distribution of nutrients among biotic and abiotic components is an important step in developing prescriptions for managing these stands for tamablesussustainable productivity eight Q gambelligambeliigambelii stands were sampled for concentrations and accumulations kg ha I1 of total nitrogen N phosphorus P sulfur S calcium ca magnesium mg potassium K and sodium na among aboveground and belowbelowgroundground biomass components and the upper 30 cm of soil highest concentrations of N P and S occurred in oak leaves understory leaves and the forest floor layer generally highest concentrations ofcaof ca mg K and na occurred in the soil the greatestgi eatesteanest proportion of the total capital of individual nutrients was contained in the soil 82 99 aboveground components of live biomass standing and down dead and forest floor contained 10 14 and 8 respectively of total capitals of N P and S the forest floor had the largest accumulation 63 of total nutrients N PE S ca mg K and na of live and dead aboveground components nutrient accumulation in live biomass was heavily weighted to the belowbelowgroundground component the dense system of roots rhirhizomeszomes and lignotubers comprising 56 of total biomass contained 62 of the total accumulation of nutrients in live biomass low levels of total P in the soil and accumulation of 14 of the ecosystem total of P in aboveground biomass components suggest the need for a better understanding of the role of P in productivity of these stands inm development of pre- scriptionsscriptions for management of residues after harvest

key words nutrient cycling soil nutrients nitrogen phosphorus sulfur cationscanionscations quercus gambeliigambelligamb elii utah

gambel oak quercus gambehigambeliigambgambelineliieill nutt is in which nutrients are distributed among the found as a small shrub or large tree on about abiotic and biotic components of the system 383.8 million ha in colorado arizona new this information will help develop manage- mexico and utah it is a clonal species that ment guidelines so that harvest activities do sprouts readily after harvest or other distur- not deplete nutrients to the extent that future bance from a dense belowbelowgroundground system of site productivity may be jeopardized lignotubers and rhirhizomeszomes tiedemann et al our objectives were to determine the con- 1987 the lignotubers are similar to those centrationscentrations and total amounts of major plant found on eucalyptus carrodus and blake nutrients nitrogen N phosphorus P sul- 1970 rhizomesRhizomes belowbelowgroundground stems are also fur S calcium ca potassium K magne- common in oaks muller 1951 sium mg and sodium na in live and dead with increasing demands for fuelwood Q gambehigambeliigambgambelineliihill biomass components and in soil throughout its range Q gambeliigambehigambgambelineilieliieill is coming understory and forest floor of a representative under close scrutiny for its initial value as a portion of the Q gamberigambehigambeliigambelligambeliihill ecosystem in central fuelwood source and for continued fuelwood utah and to relate findings to similar studies production potential wagstaff 1984 clary and in other hardwood stands this study was a tiedemann 1992 the density of the wood its companion to a study of biomass distribution superior heat yielding qualities compared with clary and tiedemann 1986 softsoftwoodswoods barger and ffolliott 1972 and its sprouting nature tiedemann et al 1987 make STUDY AREAS AND METHODS this species ideal for fuelwood management in the development of management strate- eight Q gambehigambeliigambgambelineliihill stands plots were selected gies for sustainable productivity of Q gambehigambeliigambgambelinehieblebieliieilieill near ephraim in central utah the stands were an important step is to determine the manner on slopes with gradients from 5 to 40 soils

pacificipacific northwest research station 1401 gekeler la grande OR 97850 antei21nteimountainntei mountain research station 316 east myrtle street boise ID 83702

119 120 GREAT BASIN naturalist volume 56

are typic calcixerolls formed on alluvium and to mineral soil no separation into litter L colluvium derived from limestone sandstone fermentation F and humus H layers was and shale swenson et al 1981 soils are cob- made hence the forest floor includes plant bly loamscoams in the surface 50 cm and very stony detritus accumulated above mineral soil in- clay loamscoams in the substratum to depths of 150 cluding down and dead oak 05os050.5 cm all sam- cm elevations of the 8 stands range from ples were oven dried at 70c70 C and weighed to 2089 to 2480 in average annual precipitation determine mass per unit area kg hahatihaci1 of the ranges from 36 to 51 cm and the annual frost forest floor weight of down and dead oak 05os050.5 free period is 90 to 110 d swenson et al cm was assigned to the category of down and 1981 dead oak trees A small sample of each compo- plot sizes varied in approximately inverse nent from each 1 m2ma plot was used for nutri- proportion to tree stem density clary and ent analysis forest floor samples contained tiedemann 1986 we attempted to obtain a some soil as a result of wind deposition and sample of the range of stand densities and the fact that sampling results in collection of a stem heights A 3 X 3 m plot was used for the small amount of soil from the forest floorsoilfloor soil densest stand 34444 stemshastemskastemsha a 10 X 10lom m interface therefore weights of forest floor plot for the least dense stand 5000 stemshastemskastem sha samples were adjusted for content of soil by mean ages of stems ranged from 37 to 109 yr determining weight loss on combustion of clary and tiedemann 1986 small samples in a muffle furnace at 900c900googooc C at each plot all live stems were counted combustion of organic materials results in a and numbered and 5 were selected at random small amount of mineral ash residue of 5 g per for measurement of height diameter biomass 100 g of forest floor tiedemann 1987b we and nutrient concentration sample stems adjusted forest floor weights by this amount were cut about 4 cm above the ground parti- soil volume weight bulk density was tioned into 60 cm sections and weighed in determined by collecting a 15 to 20 cm diam the field live and dead branches and leaves eter sample to a depth of 30 cm at each of the were removed A 10 cm portion of each bole subplotssubplots after vegetation was harvested and section was placed in a plastic bag sealed and the forest floor sampled this was the maxi- returned to the laboratory for determination of mum depth feasible to collect without using moisture content and nutrient concentrations mechanized digging apparatus because of the live branches dead branches and leaves from increased rocks cobbles roots and rhirhizomeszomes each tree were bagged returned to the labora- at greater depths the soil hole was lined with tory and oven dried at 70c70 C to constant weight plastic and the volume determined by measur- after weighing a sample was taken from each ing the quantity of water to the nearest 10 ml component foiforholbol analysis of nutrient concentra- required to fill the hole soil was oven dried at tion standing dead trees were counted on 70c70 C weighed and retained for nutrient each plot and 5 were randomly selected to be analysis this method of bulk density determi- cut and weighed in the field A section was nation compares favorably with the paraffin taken from each including any attached clod technique howard and singer 1981 branches for determination of moisture and one plot plot 8 was hydraulically exca- nutrient concentrations vated to a depth of I1 in by use of a hydraulic understory biomass including Q gambe pump capable of supplying 114 lminalmin tiede- lnhi 1 in other shrubs herbaceous plants for- mann et al 1987 all roots rhirhizomeszomes and ilg est floor and down and dead oak was sam- tubersnonotubers were removed and transported to the pled on three 1 m2ma subplotssubplots randomly located laboratory for drying dissecting weighing within each plot except plot 8 where only I1 and nutrient analysis weight of roots at subplot was sampled plot 8 was sampled at a depths I1 in was estimated from taper weight different time from plots 1 7 with the main relationships established for the first I1 in of objective of excavation to determine charac- vertical roots A composite sample of the roots teristicste of the underground system tiede- io10 101.0 cm 101.0io 252.5 cm and 25252.5 cm and rhi mann et al 1987 we inadvertently collected zomes was taken for nutrient analysis the only 1 subplot for determination of understory proportion of each component in the sample biomass forest floor down and dead oak and was weighted on the basis of its proportion of soil on all subsubplotsplots forest floor was collected total weight 199611996 NUTRIENT distribution IN QUERCUS CAMBELIIGAMBELH 121

each 10 cm bole portion was separated of nutrients to a kg ha 1 basis extrapolation into 8 equal radial segments one of these was based on application of taper weight rela- from each portion was further separated into tiontionshipsships for each root heartwood sapwood and bark samples from for purposes of data presentation nutrient each radial segment were then comcompositescompositedposited for contents kg ha 1 of individual aboveground each tree prior to analysis all vegetation sam- biomass components were grouped into three ples were ground to 025 mm fineness in categories 1 aboveground live overstory and preparation for analysis of nutrient concentra- understory vegetation 2 standing and down tion soil samples were sievedsievek through a 2 dead that includes standing dead trees dead mm mesh screen and ground to 0125 mm branches on live trees down and dead trees fineness prior to analysis and dead branches on the ground 05os050.5 cm all samples were analyzed for total N by and 3 the forest floor that includes all plant kjeldahl digestion followed by titrimetric deter- detritus above mineral soil except for quercus minationmination of distilled ammonium bremner branches 05os050.5 cm 1965 for total P by sulfuric acid selenium analysis of variance in a randomized com- digestion parkinson and alienallenailen 1975 followed plete block design with the 8 individual plots by molybdenum blue determination of P olsen as blocks was used to determine differences in and dean 1965 for total S by the procedure concentration among aboveground biomass of tiedemann and anderson 1971 and for components for each nutrient constituent total cationscanions ca mg na and K by atomic steel and torrie 1960 biomass component absorption spectroscopy jones and isaac 1969 was the main effect term in the analysis val- on the sulfuric acid selenium digest used for ues for the 5 individual trees and for the 3 for- total PE est floor and understory subsubplotsplots in each of mass per unit area kg ha 1 of individual the 8 plots blocks were pooled and the means plot values for each individual biomass com- were used in the analysis of variance statistical ponent of trees leaves live branches standing comparison with underground biomass com- dead etc from the study of clary and tiede- ponents was not possible because this was de- mann 1986 were used to convert concentra- terterminedmined on only I1 plot where the F test was tions of individual nutrients to mass per unit significant differences among individual bio- area kg ha 1 in the biomass determination mass components were determined using the clary and tiedemann 1986 stems were not LSD test carmer and swanson 1971 signifi- partitioned into bark heartwood and sapwood cant differences are expressed at FP ooi0010.01 no we determined the percentage by weight of statistical tests were applied to kg ha 1 nutri- these 3 components for each bole and con- ent content data because individual compo- verted weights to kg ha 1 for each plot using nents were summed to provide more inclusive values from clary and tiedemann 1986 these groupings for example live aboveground bio- values were then multiplied by concentrations mass includes oak leaves live branches heart- of individual nutrients for determination of wood sapwood bark and understory leaves mass per unit area kg haglhagiha 1 content of nutri- and stems ents mass per unit area kg ha 1 values for understory vegetation down dead oak and RESULTS AND discussion forest floor were multiplied by concentra- the nutrient concentrations tion values for individual nutrients to determine mass per unit area of each nutrient bulk there were no significant differences in con- density of the upper 30 cm of soil minus par- centrationscentrations of nutrients in biomass FP 0010.01ooi ticles 2 mm was used to develop mass per among plots blocks for any nutrient con- unit area kg ha 1 values for soil so we could stituent except ca differences among bio- convert nutrient concentration values to mass mass components were highly significant for of individual nutrients per hectare mass per every nutrient constituent unit area values of quercus roots rhirhizomeszomes nitrogen concentrations in the forest floor and lignotubers in the upper I1 m of the exca- and in quercus leaves were significantly higher vated plot plus the extrapolation of larger than in any other component table 1 under- 25 252.5 cm vertical roots to their extinction story leaves were significantly lower in N con- point was used to convert concentration values centrationcentration than the forest floor or quercus 122 GREAT BASIN naturalist volume 56

lableIABLLTABLE 1 concentration percent of nutrient constituents in biotic and abiotic components of quercus gambellgambehigambeln live dead standing nutrient leaves branches heartwood sapwood bark branches dead trees N 157 056 015 027 062 055 035 lsd001LSD ooi001 008

P 021 003 0003 002 002 002 001 LSD ooi0010.010 01 0024

S 008 003 003 002 004 004 004 LSDlsd001ooi0010.01 0014 ca 091 090 017 017 155 098 100 lsd001LSD 001 030 mg 035 016 002 004 020 014 008 LSD ooiool0010.010 01 029

K 068 036 033 015 032 026 021 lsd001LSD ooi0010.01 018 na 004 001 001 0002 001 001 001 lsd001LSD ooi0010.01 0007 coinacompnisonscoinp arisonsaniongimong aboveground hiomissmoimot nass components only leaves we did not observe increases in N con- ha 1 at this site clary and tiedemann 1986 centrationcentration of the forest floor that usually provides a continuous supply of N to the soil accompany decomposition mineralization and through decomposition and leaching leaching of other constituents from the fallen leaves of understory plants 0270270.27 quer- overstory leaves bocock 1963 gosz et al cus leaves 0210210.21 and forest floor 0120120.12 had 1973 in a litter bag study klemmedson 1992 highest concentrations of PE differences among measured a 60 increase in N concentration these 3 components were significant reduced in Q gamberigambehi leaves in the litter layer over a concentration of P in the forest floor compared 750 d time span differences between our to quercus leaves corresponded to observa- observations and those of klemmedson were tions of klemmedson 1992 concentration of probably because we report comparisons be- P in Q gambehigambeliigambgambelineliieill leaves at the surface of the tween quercus leaves and the entire forest forest floor began to decrease shortly after floor whereas his comparisons were for the lit- deposit and declined steadily for 500 d to ter layer only lowest concentrations of N were about 60 of original concentration concen- observed in the heartwood standing dead and tration then leveled off for the remaining 250 down dead trees were both higher in N con- d of the experiment our lowest levels of P centrationscentrations than were heartwood and sapwood occurred in the heartwood 000300030.003 although of living stems this probably resulted from there were some significant differences among selective decomposition and loss of other ele- other biomass components the actual differ- ments causing an increase in the concentration ences were slight and probably of little biolog- of N in standing dead and down dead trees ical significance total P in soil 0020020.02 was concentration of N in the upper 30 cm of substantially below normal levels which are soil 0420420.420 42 was greater than would be expected 0090.09 0130.13 for soils of the united states for this site according to jenny 1941 the parker et al 1946 normal range of soil N for semiarid sites is concentrations of S were greatest in forest 0100.10olo0 10 0250.25ogs0 25 for the surface 10 cm the high floor 0120.12 and understory leaves 0110110.11 content of N in these soils can probably be and there was no significant difference between attributed to 2 principal factors 1 the high these 2 components however S concentra- clay content is conducive to retention of high tion in both was significantly higher than in levels of organic N klemmedson and jenny quercus leaves lowest S concentrations in 1966 millar et al 1966 and 2 the extraordi- aboveground components were in the sapwood nary accumulation of forest floor 37348 kg and heartwood our comparisons of N and S 199619961 NUTRIENT distribution IN QUERCUS cambelligambelCAMBELgambelliGAMBELII11 123 ecosystems in cental utah a understory understory down dead forest roots and leaves stems trees floor rhirhizomeszomes lignotubers soil 146 054 043 166 044 033 042

027 005 001 012 003 002 002

011 004 006 012 004 003 004

0980.980 98 0610.610 61 0760.760 76 2672.672 67 0970 97 1151.151 15 129

0400 40 017 011 1151.151 15 014 0090 09 192

114 064 007 043 021 014 087

0008 002 0005 006 002 0008 008

levels in the forest floor with quercus leaves tionseions were higher concentrations of ca in the presented an anomaly we would expect S forest floor and in the bark of quercus trees comparisons between forest floor andquercusand quercus and K in understory leaves leaves to be similar to those for N because S calcium concentrations in the forest floor is a companion to N in several amino acids layer were more than 252.5 times greater than allaway and thompson 1966 coleman 1966 quercus leaves the content ofcaof ca in bark was klemmedson s 1992 observations bear this nearly 10 times greater than heartwood or sap- out because both N and S concentrations in wood quercus leaves live branches dead quercus leaves increased about 60 over a branches standing dead trees and down dead 750 d period after deposition at the surface of trees were all comparable in ca concentration the forest floor however when we compared magnesium concentrations in biomass com- quercus leaves and the entire forest floor it ponents were highest in the forest floor appeared that N and S responded differently layer approximately 3 times greater than in over the long periods required for develop- quercus and understory leaves in contrast to ment of the forest floor nitrogen concentra- ca patterns mg concentrations in live branches and S tion tended to remain constant concen- and standing dead and down dead trees were increased over time mineralization of tration significantly lower than in quercus leaves S deeper layers of the forest floor may pro- in understory leaves were significantly higher ceed more slowly than mineralization of N in K concentration 1141141.14 than were quercus thereby resulting in an increase in S concen- leaves 0680.68 or understory stems 0640.64 trationtration products of decomposition for N may 068 064 potassium were about equal for also be more mobile than those for S concentrations live branches heartwood and bark and about total S concentration in soil 0040040.04 was in the middle of the range reported for US soils half the concentration found in quercus leaves ooi0010.01 0060.06 burns 1968 the ratio of NS of concentration of K in forest floor was substan- 101 in soil indicates that the S level is great tially lower than in quercus leaves and may enough that N will be efficiently utilized for reflect the ease with which K is leachedbeached from the formation of plant proteins black 1968 the forest floor relative to the other cationscanions burns 1968 attiwillAttiwill 1968 concentrations of the 4 measured cationscatcanionsions highest concentrations of na occurred in ca mg K and na were generally higher in quercus leaves and in the forest floor differ- the soil than in any plant component excelexcep ences among other biomass components were 124 GREAT BASIN naturalist volume 56 minorminon even though some were statistically comparable except for higher concentrations significant ofofnoanN 25252.5 in leaves of P tremultremuloidesoides concen- comparisons of cation levels in quercus trattrationsions of P K and ca were similar for all tree leaves with levels min the forest floor were varivarlvari- components sodium concentrations were gen- able between our study and results of the literally greater in Q gambgambehigambeliigambelineliiellieili than in P tremuteemu ter bag study of klemmedson 1992 we loides concentrations of N P and S in live showed significantly greater ca and mg in the aboveground biomass of Q gambehigambeliigambgambelinelii were forest floor than in quercus leaves klemmed- comparable to those reported for Q robur in son 1992 found similar increases in ca in russia rodin and bazilevich 1967 and in quercus leaves over 750 d however mg con- belgium duvigneaud and denaeyer de smet centrationcentration in his study declined to about 80 1970 concentrations of N in forest floor and of the level in fresh leaves over the 750 d dead branches also were comparable to values study differences in K concentration that we for southern and eastern US quercus stands found between quercus leaves and the forest lang and forman 1978 concentrations of floor were not nearly as great as the decline inm cationscanions in our study did not agree as well with K concentration over time in the litter layer those presented in the literature as for N P measured by klemmedson 1992 potassium and S for example Q gambehigambeliigambgambelineliieill forest floor concentration in quercus leaves declined concentrations of K and mg were 3 and 8 about 70 in 500 d and then stabilized to the times greater than those reported for Q robur end of the 750 d study differences between calcium concentrations in Q gambehigambeliigambgambelineliieill were klemmedson s observations and ours were substantially greater than those observed in probably a result of the fact that he studied other studies in forest floor live branches changes in nutrient concentration in the litter dead branches standing dead trees and layer and our comparisons were with the down dead trees forest floor entire distribution of nutrient capital is little information on the there is concen- among components trationstrations of nutrients in biomass components in western hardwood stands there are 2 appar- comparisons of nutrient distribution be- ent reasons for this compared with the east- tween above and belowgroundbelowground components ern united states the area occupied by stands must be considered from the perspective that of hardwood species min the west is minor our soil sampling was restricted to the upper therefore until recently western hardwoods 30 cm because of rock and the massive under- have not been viewed as an economically im-im ground structures of Q gambehigambeliigambgambelinehlehieliihill the actual portant resource rather they were considered zone of rooting and nutrient acquisition was weed species because they were assumed to undoubtedly much greater than the area we compete with marketable coniferous trees or sampled therefore our estimates of the pro- with understory forage producing species portions of nutrients in aboveground compo- with emerging demands for fuelwood and nents were likely to be higher than if the new markets for unique woods for furniture entire rooting zone had been sampled also there is increased awareness of the value of the kg hahai 1 estimates were for the area of the western hardwoods and especially Q gambgambelngambeangambelffeiffelff actual clone sampled clones of Q gambehigambeliigambgambelinelii do wagstaff 1984 clary and tiedemann 1992 not occupy the entire area of the sites on nutrient concentrations of leaves agreed which they occur most studies take into closely with those reported by klemmedson account the high and low density areas of 1992 for Q gamberigambehigambehz in northern arizona tree occupancy in determining nutrient distri- bartos and johnston 1978 determined the bution therefore in making projections to an concentrations and proportions of individual areal basis the actual area occupied by Q nutrients in the various components of 3 gambehigambeliigambgambelineilieliieill clones must be considered clones of populus tremultremuloidesoides michxmicha quaking the greatest proportion of total nutrient aspen trees in utah and wyoming but did not capital sampled was contained in the soil consider the forest floor understory and down table 2 of the total capitals of individual dead components of the nutrient pool con- nutrients 82 99 were contained in the centrationscentrations of N in the various tree compo- soil aboveground accumulations of individual nents of Q gamberigambehi and P tremultremuloidesoides were nutrients in live biomass standing and down 199611996 NUTRIENT distribution IN QUERCUS GAMBELII 125

TABLE 2 distribution of nutrients among biomass forest floor standing plus down dead and soil components of Q gamberigambehi stands livel standing liveclive0levec above- plus total below totalstotaldtotal1 ground down forest aboveground live totalftoralftotalc nutrient biomass1biomass3iomass dead floor ground biomass biomass soile capital nitrogenNiti ogen kg ha I1 245 140 654 1039 270 515 9500 10810 of total aboveground 24 13 63 of total capital 10 2 88 phosphorus kg ha 1 19 4 48 71 19 38 410 500 of total aboveground 27 5 68 of total capital 14 4 82 sulfur kg ha 1 19 13 46 78 22 41 946 1046 of total aboveground 24 17 59 of total capital 8 2 90 calcium kg harlhariha 1 334 303 1167 1804 924 1258 28844 31571 of total aboveground 18 17 65 oftotalof total capital 6 3 91 magnesium kg ha I1 62 35 381 478 63 125 42485 43023 of total aboveground 13 7 80 oftotalof total capital I1 1 99 potassium kg harlhariha 1 201 72 144 417 116 317 20268 20801 of total aboveground 48 18 34 of total capital 2 1 98 sodium kg harlhariha 1 7 4 20 31 7 14 1765 1804 of total aboveground 22 13 65 oftotalof total capital 2 1 98 total kg ha 1 887 571 2460 3918 1421 2308 oftotal aboveground 23 14 63 of total in living biomass 38 62

includesalncludes living aboveground overstoryover tory and understory vegetation includesbincludes rilriiillallaliail forestfoiroiboi est floor layers above mineral soil includeincludes roots rhiwmesrhirhizomeszomes and lignotubers in the upper 100 cm of soil standing crop plus belowgroundbelow ground biomassbioblomasimass eupperaupper1 30 cm of01 soil standing crop plus standing and down dead plus forest floor plus belowbelowgioundbelowgroundground biomass plus soil

dead and forest floor ranged from 31 kg ha 1 forest and woodland ecosystems klemmedson for na to 1804 kg ha 1 for ca proportions of 1975 brown 1977 tiedemann 1987a total capitals of N P and S in aboveground the forest floor was the most important components were highest with 10 14 and aboveground reservoir of nutrients with 63 8 respectively the proportion of N the most of the total accumulation above ground accu- widely reported nutrient in aboveground mumulationslations of individual nutrients in the forest components 10 was comparable to that floor ranged from 20 to 1167 kg ha I1 and con- described for other semiarid and temperate stituted 34 80 of the aboveground capitals 126 GREAT BASIN naturalist volume 56 total nutrient content of the forest floor in our ativelyactively constant narrow range of 200 300 kg Q gambgambehigambeliigambelineliieilieilf clones 2460 kg hatihatlha 1 substantially ha 1 normally found in leaves reported by exceeded the range described by lang and rodin and bazilevich 1967 forman 1978 in their summary for US quercus forests 206 kg ha 1 yount 197519751 to conclusions 1462 kg hahaglhagi1 gosz et al 197619761 greater accu- mulationmulation of ca in the forest floor layer 1167 gambel oak appears to be unique from kg haba 1 compared with that reported by other other deciduous forests in the accumulation of observers 98 400 kg ha 1 lang and forman nutrients in the forest floor and in below 1978 accounted for much of the difference in ground biomass components both were major leaves total accumulation of nutrient elements in Q areas of nutrient accumulation the in gambehigambeliigambgambelineliihill compared with other quercus stands contrast were a minor storage area of aboveground also forest floor biomass accumulation in our accumulation nutrients in 1 living and dead components expressed as a Q gambgambehigambeliigambelineliihill stands 37348 kg ha clary and proportion of total site nutrients was similar to tiedemann 1986 was near the upper limit that reported for other semiarid and temper- 46800 kg ha 1 of that presented for US ate forest habitats quantity ofoanN the most quercus forests lang and forman 1978 the ofn commonly measured nutrient stored in the massive belowbelowgroundground system of lignobigno the forest floor also agreed well with this litera- tubers rhirhizomeszomes and roots comprised 56 of ture it should be noted that had we been able the total biomass of Q gambeliigambelffgambelligambeliieillelffeiffeluf clary and to sample a larger proportion of the total root- 1986 and contained 1 to 4 of tiedemann ing zone the proportion of the total nutrient of individual the total of the capitals nutrients capital aboveground would likely have been however relative to the total nutrient accu- smaller mumulationlation in live biomass the live belowgroundbelowground low levels of P in the upper 30 cm of soil component was an important storage area con- suggest that this element may limit productiv- taining 37 74 of the individual nutrient ity of Q gamberigambeliigambehigambelligambehichicblelii because of potential limita- accumulations the proportion of total nutri- tions in the soil accumulation of 14 71 kg ents in belowgroundbelowground biomass 61 substan- halhaxha 1 of the total ecosystem P in aboveground tially exceeded the range for deciduous forests living and dead components we suggest cau- worldwide 30 40 summarized by rodin tion in the way the forest floor and residues and bazilevich 1967 this finding supported are managed elwoodfullwoodfuelwoodFu harvest followed by the conclusions of chattaway 1958 robbinsbobbins et removal of residues by broadcast burning could al 1966 and blake and carrodus 1970 that cause large losses of P depending on degree storage of nutrients is an important function of of consumption of organic matter and fire belowbelowgroundground components such as lignotubers temperatures covington and debano 1988 total content of nutrients in the entire debano 1988 this loss may reach 60 of 71 organic component total live and dead above- kg ha 1 if fuels are totally consumed raison ground and belowbelowgroundground biomass of our Q et al 1985 however such losses need to be gambehigambeliigambgambelineillelii stands 5339 kg ha 1 was in the mid- weighed against changes in P availability that dle of the range for deciduous forests world- result from burning in his summary of plant wide 2000 7500 kg ha 1 summarized by and litter contained nutrients debano 1988 rodinbodin and bazilevich 1967 similarly total indicated that fire induced increases in P avail- nutrient content of live biomass 2308 kg ha 1 ability decline and reach pre fire levels within was comparable to values for oak forests in I1 yr debano and klopatek 1988 showed that russia 2600 3400 kg ha I1 rodin and bazile- inorganic P is released by prescribed burning vich 1967 but is quickly immobilized and may not be worldwideWorld addeAide leaves usually constitute aclo8clo8 10 readily available for plant growth of the store of mineral elements in plant bio- although there are also substantial accu- mass rodin and bazilevich 1967 mineral mumulationslations of N and S in aboveground biomass element accumulation in Q gambehigambeliigambgambelineillelii leaves and these are sensitive to losses from vola- and understory leaves 245 kg ha 1 not shown tilization knight 1966 tiedemann 1987b in table 2 comprised 11 of the total mineral they are not limiting in the soil and quantities content of live biomass and was within the rel are likely sufficient to replenish losses 199611996 NUTRIENT distribution IN QUERCUS GAMBELII 127

fertilizer amendment with P may warrant COLEMAN R 1966 the importance of sulfur as a plant soil 101 consideration as a means of Q gam nutrient in world crop production science improving 230 240 behlbebi productivity after harvest this decision COVINGCOVINGTONTON W W AND L FE DEBANO 1988 effects of however should be based on soil tests to fire on pinyon juniperjumper soils pages 78 86 in J S determine the availability of PE krammes technical coordinator effects affireoffireof firerire management of southwestern natural resources USDA forest service general technical report RM 191 literature CITED rocky mountain forest and range experiment station fort collins CO ALLAWAY W H AND J FE THOMPSON 1966 sulfur in the DEBANO L F 1988 effects of fire on the soil resource in nutrition of plants and animals soil science 101 arizona chaparral pages 65 77 in J S krammes 240 247 technical coordinator effects of fire management of ATriATTIWILLwILL P M 1968 the loss of elements from decom- southwestern natural resources USDA forest service posing litter ecology 49 142 145 general technical report RM 191 rocky mountain BARGERBARCER R L AND P FE FFOLLIOTT 1972 the physical forest and range experiment station fort collins characteristics and utilization of major woodland CO tree species in arizona USDA forest service DEBANO L F AND J M KLOPATEK 1988 phosphorus research paper RM 83 80 appp rocky mountain for- dynamics of pinyon juniperjumper soils following simulated est and range experiment station fort collins CO burning soil science society of america journal 52 BARTOS D L AND R S JOHNSTON 1978 biomass and 271 277 nutrient content of quaking aspen at two sites in the duvigneaud P AND S DENAEYER DE SMET 1970 bio- western united states forest science 24 273 280 logical cycling of minerals in temperate deciduous forests pages 199 225 D E editor BLACK C A 1968 soil plant relationships 2ndand edition in reichle spnngerspanger john wiley and sons inc new york 792 appp analysis of temperate forest ecosystems springer verlag berlin heidelberg germany BLAKE T J AND B B CARRODUS 1970 studies on the gosz J R G E LIKENS AND F H BORMANN 1973 lignotubers of eucalyptus obliquaobhquaoblique II11 endoge- dhen nutrient release from decomposing leaf and branch nous inhibitor levels correlated with apical domi- litter in the hubbard brook forest new hamp- new 69 1073 1079 nance phytologist shire ecological monograph 43 173 191 BOCOCK changes amounts of dry matter K L 1963 in the 1976 organic matter and nutrient dynamics of carbon and energy decomposing wood- nitrogen in the forest and forest floor in the hubbard brook for- land leaf litter in relation to the activities of soil est Occooecologiaoccologialogia berlin 22 305 320 273 284 fauna journal of ecology 52 HOWARD R F AND M J sincerSINGERSINGEK 1981 measuring forest BREMNER J M 1965 total nitrogen pages 595595624624 in soil bulk density using irregular hole paraffin clod C A black D D evans L E ensmingerensmmger et al and air permeability forest science 27 316 322 editors methods of soil analysis part 2 chemical JENNY H 1941 factors of soil formation mcgraw hill and microbiological properties agronomy 9 ameri- book co inc new york 281 appp can society of agronomy inc madison WI JONES J G JR AND R A ISAAC 1969 comparative ele- BROWN T H 1977 nutrient distribution of two eastern mental analysis of plant tissue by spark emission and washington forested sites unpublished master s the- atomic absorption spectroscopy agronomy journal sis washington state university pullman iiiliiI1111 I1 I1 appp 6139361 393 394 BURNS G R 1968 oxidation of sulfur in soils sulphur klemmedson J 0 1975 nitrogen and carbon regimes in institute technical bulletin 1 41 appp an ecosystem of young dense ponderosaponde iosalosa pine inm ari- CARMER S G AND M R SWANSON 1971 detection of zona forestfoiest science 21 163 168 differences between means a monte carlo study of 1992 decomposition and nutrient release flomfrom gambel oak and leaf lit- five pairwise multiple comparison procedures mixtures of ponderosa pine agronomy journal 63 940 945 ter forest ecology and management 47 349 361 J AND H JENNY 1966 nitrogen avail- CARRODUS D D AND T J BLAKE 1970 studies on the hgilg119 klemmedson 0 ability inm california soils in relation to precipitation notuber of eucalyptus obliquaobhquaoblique ehen I1 the nature and parent material soil 102 215 222 ofthehgnotuberof the lignotuber new phytologist 69 1069 1072 science KNIGHT H 1966 loss of nitrogen from the forest floor by CHATTAWAY M M 1958 bud development and lignotulignottibignotti burning forestry chronicle june 149 152 beibelber formation in eucalyptseucalyptus australian journal of LANG G E AND R T T FORMAN 1978 detrital dynamics botany 6 103 115 in a mature oak forest hutcheson memorial forest CLARY TIEDEMANN of W P AND A R 1986 distribution new jersey ecology 59 580 595 biomass within small tree and shrub form quercus MILLAR C E L M TURK AND H 0 FOTH 1966 funda- 242 gambelligambeliigamb elii forest science 32 232 mentals of soil science john wiley and sons inc 1992 ecology and values of gambel oak wood- new york 491 appp lands pages 87 95 in P F ffolliott et al technical MULLER C H 1951 the significance of vegetation repro- coordinators ecology and management of oak and duction in quercus madronoMadrofioflo 11 129 137 associated woodlands perspectives in the south- OLSEN S R AND L A DEAN 1965 phosphorus pages western united states and northern mexico 27 30 1035 1049 in C A black D D evans L E ens- april 1992 sierra vista AZ USDA forest service minger et al editors methods of soil analysis part general technical report RM 218 rocky mountain 2 chemical and microbiological properties agron- forest and range experiment station fort collins omy 9 american society of agronomy inc madi- CO son WI 128 GREAT BASIN naturalist volume 56

PARKLRFWJPARKER E W J R ADAMS K G CLARKETALCLARK ET AL 1946 fer site productivity pages 352 359 in R L everett tilizerstiliutilizerszers and lime in the united states resources pro- compiler proceedings pinyon juniper conference duction marketing and use USDA miscellaneous reno NV 13 16 january 1986 USDA forest ser- publication 586 washington DC 94 appp vicevlee general technical report INT 215 intermoun- PARKINSON J S AND S E ALLEN 1975 A wet oxidation tain research station ogden UT procedure suitable for the determination of nitrogen 1987b combustion losses of sulfur from forest and mineral elements in biological material com- foliage and litter forest science 33 216 223 munications in soil science and plant analysis 6 TIEDEMANN A R AND T D ANDERSON 1971 rapid analy- 1 11 sis of total sulfur in soils and plant material plant ralsonraibonRAISON R J P K KHANNA AND P V WOODS 1985 and soil 35 197 200 mechanisms of element transfer to the atmosphere TIEDEMANN A R W P CLARY AND R J BARBOUR 1987 during vegetation fires canadian journal of forest underground systems of gambel oak quercus gam research 15 132 140 beill1 in central utah american journal of botany ROBBINS W W T E WEIERWLILR AND C R STOCKING 1966 47106547 1065 1071 botany an introduction to plant science john wiley WAGSTAFF F J 1984 economic considerations in use and and sons inc 614 appp management of Cambelgambel oak for fuelwood USDA for- RODIN L E AND N I1 bazilevich 1967 production and est service general technical report INT 165 8 appp mineral cycling in terrestrial vegetation english intermountain forest and range experiment sta- translationtiantran slation by G E fogg editor oliver and boyd tion ogden UT edinburgh london 288 appp YOUNT J D 1975 forest floor nutrient dynamics in STEELSTLEL R G D AND J H TORRIE 1960 principles and southern appalachian hardwood and white pine plan- procedures of statistics mcgraw hill new york tation ecosystems pages 598 608 in FE G howell 481 appp J B gentry and M H smith editors mineral cycling SWENSON J L D beckstrand D T ERICKSON ET AL in southeastern ecosystems energy research and 1981 soil survey of sanpete valley area utah USDA development administration symposium series soil conservation service and USU S department of technical information center springfield VA the interior bureau of land management in cooperation with utah agriculture experiment station received 17 october 1995 and utah state division of wildlife resources 179 accepted 11 march 1996 PP TIEDEMANN A R 1987a nutrient accumulations in pinyon juniper ecosystems managing for future great basin naturalist 562 D 1996 appp 129 134

comparison OF TWO ROADSIDE SURVEY procedures FOR DWARF mistletoesMISTLETOES ON THE SAWTOOTH NATIONAL FOREST IDAHO

robert L Mathiasenmathiasenlmathiasen11 james T Hoffman 2 john C guyon3guyons and linda L Wadleigh 4

ABSTRACT two roadside surveys were conducted for dwarf mistletoesmistletoes parasitizing lodgepole pine and douglas fir on the sawtooth national forest idaho one survey used vanablevariable radius plots located less than 150 m from roads the 2ndand survey used vanablevariable radius plots established at 200 m intervals along 1600 m transects run perpendicular to the same roads estimates of the incidence percentage of trees infected and percentage of plots infested and severity average dwarf mistletoe rating for both lodgepole pine and douglas fir dwarf mistletoesmistletoes were not significantly different for the 2 survey methods these findings are further evidence that roadside plot surveys and transect plot surveys conducted away from roads provide similar estimates of the incidence of dwarf mistletoesmistletoes for large forested areas

key words dwarf mistletoesmistletoes surveys lodgepole pine douglas fir

dwarf mistletoesmistle toes arceuthobium sppapp are naissance information with data collected using damaging disease agents in many western variable radius or fixed area plots located near forests hawksworth and wiens 1995 in the roads roadside plot surveys for estimating intermountain west lodgepole pine pinus con the incidence percent of trees or plots in- torta dougl ex loud and douglas fir pseudo- fected and severity intensity of infection in tsuga menziemenziesiimenziesiasii birbmirb franco are the most individual trees hawksworth 1956 1958 commonly infected trees hawksworth and hawksworth and lusher 1956 andrews and wiens 1972 1995 hoffman 1979 each of daniels 1960 graham 1960 1964 dooling these hosts is parasitized by a different dwarf 1978 hoffman 1979 johnson et al 1980 mistletoe lodgepole pine dwarf mistletoe A johnson et al 1981 hoffman and hobbs 1985 americanumamericanum nutt ex engelm and douglas merrill et al 1985 maffei and beatty 1988 fir dwarf mistletoe A dougladouglasiisii engelm roadside reconnaissance surveys consist of severe infection by these parasites is often driving roads at slow speed and recording visual associated with tree mortality reduced growth estimates of dwarf mistletoe infection within a and cone production tree deformity and pre- short distance from the roadside usually 20 m disposition to attack by other diseases andor dwarf mistletoe incidence is estimated by insects hawksworth and wiens 1995 there- determining the ratio of the number of kilo- fore resource managers in many private state meters surveyed adjacent to infected trees to and federal land management agencies imple- the total kilometers surveyed adjacent to stands ment management activities designed to reduce predominated by host trees dooling 1978 the damage associated with dwarf mistletoesmistletoes roadside plot surveys involve locating plots because information on the incidence and near roads at specific intervals and collecting severity of these pathogens is required by tree data including species diameter height resource managers for making decisions regard- age and mistletoe severity on each plot dwarf ing dwarf mistletoe management surveys are mistletoe incidence has typically been repre- commonly conducted in designated manage- sented by the percentage of plots infested ment units stands and over larger areas such with mistletoe rather than the percentage of as national forests trees infected in all plots dooling 1978 surveys of dwarf mistletoe infection over roadside surveys have the benefit of allow- large areas frequently combine roadside recon ing large areas to be surveyed rapidly and

1 idaho department of lands box 670 coeur dalene ID 83816 forestmorest pest management USDA forest serviceSe ivice 1750 front street boise ID 83702 forestmorest pest management USDA forest service 4746 south 1900 east ogden UT 84401 USDA forest service 524 25th street ogden UT 84403

129 130 GREAT BASIN naturalist volume 56

inexpensively in addition roadside surveys starting reference points were landmarks that concentrate efforts in areas that are accessible could easily be relocated such as a bridge and more likely to be considered for manage- stream crossing or road junction crews drove ment actions concerns about the reliability of a distance of 800 m from the starting reference roadside survey methods are primarily related point toward the center of each township to the bias that may be encountered by sam- they then selected a compass bearing perpen- pling mistletoe incidence and severity near dicular to the righthandright hand side of the road and roads because roads are typically constructed located an end point 120 m from the road according to topographic features in drainages three 20 basal area factor variable radius plots or along ridgetopsndgetopsridgetops rather than randomly or point samples avery and burkhart 1983 were systematically located throughout the survey established 40 m from this end point at com- area since there is evidence that dwarf mistle- pass bearings of 240 120 and 0 from the toe distribution is i elatedrelated to topography hawks- compass bearing used to locate the end point worth 1959 1968 these concerns need to be crews then drove another 800 m down the considered when conducting dwarf mistletoe road and established a and2nd cluster of 3 vari- surveys over large forested areas able radius plots using the same procedure because few surveys have compared data for each plot tree the following information collected from roadside reconnaissance or was recorded plot number species diameter roadside plot surveys with data collected from at 1371.371 37 m aboveground nearest 0250.250 25 cm sta- more intensive random or systematic surveys tus live or dead and dwarf mistletoe rating for dwarf mistletoesmistletoes over large areas hawks- DMR 6 class system hawksworth 1977 if a worth 1956 1958 johnson et al 1981 merrill plot did not contain trees it was recorded as et al 1985 we initiated this study to compare nonnonstockedstocked dwarf mistletoe incidence and severity estimates obtained from roadside plot surveys transect plot survey with those from transect plot surveys that A 1600 m approximately 1 transect sampled areas at greater distances from roads imimi along we surveyed 3 districts of the sawtooth national perpendicular to the road was run the bearing for establishing forest idaho because this national forest is same compass used roadside plots 800 from the representative of forests in the intermountain the ist set of m west where lodgepole pine and douglas fir starting reference point in each township sur- are the predominant tree species and dwarf veyed A 20 basal area factor variable radius mistletoesmistletoes are common hoffman 1979 hoff- plot was located every 200 m along each tran- man and hobbs 1985 sect for a total of 8 plots information recorded for plot trees was the same as above METHODS analyses we a used a roadside plot survey and tran the incidence of each species of dwarf sect plot survey to collect dwarf mistletoe in-in mistletoe percentage of trees infected was 3 districts cidence and severity data in adjacent calculated for each set of roadside plots up to ranger fairfield ranger ketchum district 6 plots and each set of transect plots up to 8 and sawtooth district national recreation plots for each township incidence was calcu- area of the sawtooth national forest idaho lated on a per hectare basis by multiplying by in 1990 we surveyed each district by arbitrar- per hectare conversion factors based on 2542.542 54 ily selecting a major road system in each town- classes for 20 basal area factor ship 10 of federally man- cm diameter containing sections avery aged land townships with no roads or with few vanablevariable radius plots and burkhart 1983 roads were not sampled road systems were weighted dwarf mistletoe ratings were calcu- chosen before fieldwork began and adjustments lated by multiplying the DMR of each tree by also were made in the field only when selected road the per hectare conversion factors these systems weiewelewere closed or impassable weighted values were used to calculate the mean percentage of trees infected and mean survey roadside plot dwarf mistletoe rating for each survey proce- field crews arbitrarily chose a starting ref- dure in each township on a per hectare basis erence point on each selected road system these values were then used to calculate the 199619961 ROADSIDE SURVEYS FOR DWARF mistletoesMISTLE TOES 131 percentage of trees infected and a mean DMR RESULTS AND discussion for each tree species and survey method data from townships where the surveys did mean diameters for trees sampled using not sample at least 3 douglas fir or lodgepole each survey method were approximately the pine for each of the survey procedures were same for lodgepole pine and douglas fir not included in the analyses only living trees table 1 sampled tree diameters were clearly were used in the analyses for calculating mean skewed toward larger trees table 1 because DMR because it was not always possible to both survey methods used variable radius accurately assign a DMR to dead trees inci- plots that sample large trees more often than dence values were calculated for 9 townships small trees avery and burkhart 1983 for lodgepole pine and for 17 townships for because both survey methods sampled trees douglas fir the roadside plot survey sampled in the same way the survey results should be a total of 206 lodgepole pine and 357 douglas comparable however it is probable that the fir in 46 and 75 plots respectively the tran percentage of infected trees and mean DMR sect plot survey sampled 171 lodgepole pine would have been lower for both lodgepole and 342 douglas fir in 42 and 87 plots pine and douglas fir had more small trees respectively A one way analysis of variance been sampled because small trees are typically ANOVA FP 0050.05 was used to determine if less often and less severely infected parmeter the mean values for incidence and severity were 1978 significantly different between the 2 survey estimates of incidence for douglas fir dwarf procedures percentages were converted using mistletoe using the 2 survey methods were arcsin transformations before ANOVA analyses within 3 of each other based on the percent- were performed snedecor and cochran 1989 age of trees infected table 2 estimates of to compare our results with those of other douglas fir dwarf mistletoe severity were sim- dwarf mistletoe surveys we determined inci- ilar also the differences between douglas fir dence of both dwarf mistletoesmistletoes for both survey dwarf mistletoe incidence and severity for the procedures by calculating the percentage of 2 survey methods were not statistically signifi- plots infested if a plot had at least I1 infected cant the differences between estimates of the tree it was considered infested this method incidence and severity of lodgepole pine of reporting dwarf mistletoe incidence has dwarf mistletoe for the 2 survey methods were been applied in the majority of roadside plot larger than for douglas fir dwarf mistletoe surveys conducted for dwarf mistletoesmistletoes in the table 3 however the differences were not western united states significant therefore the 2 survey methods

TABLE 1 distribution of lodgepole pine and douglas firhbl sampled by diameter classes for the roadside plotpiot and transect plot surveys on the sawtooth national forest idaho lodgepoleoieole pine douglasdouglas fir roadside plot transectti ansedtansect plotpiot roadside plotpiot transect piotplot diameter mean mean mean mean class diameter diameter diameter diameter cm cm N cm N cm N cm N

2 13 91 47 86 39 106 17 91 10 14 25 196 108 198 92 206 90 201 108 26 38 292 39 300 34 318 98 320 99 39 51 419 7 429 5 437 85 447 60 52 64 607 5 511 1 566 32 574 24 64 a 965 35 894 41

TOTAL 208 206 201 171 395 357 409 342

noano trees sampled inin this sizesi7esibestze class 132 GREAT BASIN naturalist volume 56

TABLE 2 incidence and severity of douglas fir dwarf mistletoe estimated from roadside plot and transect plot surveys on the sawtooth national forest idaho incidence severity mean survey percent 95 mean 95 mean method infectinfectededa1 confidence limit mean darbdmrbdmr11 confidence limit roadside plot 284c 110 458 0oge 020.2 151.5 transect plot 258 10010.0loo 41541.5 080.8os 02 141.4 basedbased oilollon011 the peipercerltagpeicentagepetpercentagecentage of individual trees infected on a peiper hectaiehectareliectare basis idwarfinistletoedwaildwaif mistletoe ratinglilting hawksworthHawkswoiwor lil 1977 means in thistins column aieateare not significantly different one way ANOVA P 005oos0 05

TABLE 3 incidence and severity of lodgepole pine dwarf mistletoe estimated from roadside plot and transect plot surveys on the sawtooth national forest idaho incidence severity mean survey percent 95 mean 95 mean method infectinfectededa confidence limit mean darbdmrbdmr13 confidence limit roadside plot 485c485g 29429.4 675 121 060.6og 181.8 transect plot 55755.7 35335.3 76176.1 16 11 21

Bbasedsed on011 the percentagecentage ofindividnalindividualofmdividiiiil trees infected on a per hectare basis i oiloll peipet dwailDdwaifwaifacif mistletoe lataglatmgrating hawksworth 1977 means inin tinithistins column aieare not significantly different one way ANOVA P 005oos0 05 provided equivalent estimates of dwarf mistle- our surveys in the sawtooth national forest toe incidence based on the percentage of approximately 80 is higher than for most trees infected and severity for both dwarf national forests surveyed thus far an earlier mistletoesmistletoes dwarf mistletoe survey of the sawtooth national dwarf mistletoe incidence based on the forest hoffman and hobbs 1985 reported percentage of plots infested is presented in the incidence of lodgepole pine dwarf mistle- table 4 both survey methods provided esti- toe as 71 however that survey did not in- mates that were within 2 of each other for clude the sawtooth national recreation area both dwarf mistletoesmistle toes calculating dwarf mistle- the district in which we detected a very high toe incidence based on the percentage ofplotsof plots incidence of lodgepole pine dwarf mistletoe infested greatly increases the estimates of 83 therefore the sawtooth national for- dwarf mistletoe incidence when compared to est probably does have a higher incidence of the incidence based on the percentage of trees lodgepole pine dwarf mistletoe than many infected because it requires only I1 infected tree other western national forests for a plot to be treated as infested an earlier estimate of the incidence of lodgepole pine dwarf mistletoe is one of douglas fir dwarf mistletoe based on the per- the most widely distributed dwarf mistletoesmistletoes centage of plots infested for the sawtooth in the western united states hawksworth national forest was 53 hoffman 1979 and wiens 1995 the incidence of this mistle- although that survey sampled only the south- toe based on the percentage of plots infested ern districts of the sawtooth national forest has varied between approximately 40 and and did not include the districts we surveyed 70 for the majority of national forests sur- our estimate for douglas fir dwarf mistletoe veyed and averages about 50 hawksworth based on the percentage of plots infested is 1958 graham 1960 1964 johnson et al 1980 approximately the same almost 50 1981 hoffman and hobbs 1985 the incidence our findings provide additional evidence of lodgepole pine dwarf mistletoe based on that estimates of incidence and severity of the percentage of plots infested estimated from dwarf mistletoesmistletoes using roadside plot surveys 199619961 ROADSIDE SURVEYS FOR DWARF mistletoesMISTLE TOES 133

TABLE 4 incidence of douglas fir and lodgepole pine DOOLING 0 J 1978 survey methods to determine the dwarfmistletoesdwarf mistlemistiemistletoestoes based on the percentage ofplotsof plots infested distribution and intensity of dwarf mistletoe pages estimated from roadside plot and transect plot surveys on 36 44 in proceedings of the symposium on dwarf the sawtooth national forest idaho mistletoe control through forest management USDA forest service general technical report fir douglas lodgepole pine PSW 31 dwarf mistletoe mistletoe dwarf GRAHAM D P 1960 surveys expose dwarfdwarfmistletoemistletoe prob- percent percent lem in inland empire western conservation jour- survey method plots infested plots infested nanal1175617 56 58 1964 dwarfmistletoeDwarfmistletoe survey in western mon- roadside plot 75 47 46 80 tana USDA forest service research note INT 14 7ppapp7 appp plot 42 transect 87 48 78 hawksworth F G 1956 region 3 dwarfmistletoedwarfmistletoe survey progress report on the 1954 55 field workwoik mimeo- graphed USDA forest service special report approximate those of similar surveys con- rocky mountain forest and range experiment sta- ducted away from roads hawksworth 1956 tion fort collins CO 5 appp 1958 survey of lodgepole pine dwarfdwarfmistletoemistletoe on reported similar results based on a more in- the roosevelt medicine bow and bighorn national tensive comparison of roadside plot and tran forests USDA forest service rocky mountain for- sect plot surveys for dwarf mistletoesmistletoes on the est and range experiment station paper 35 13 appp mescalero apache indian reservation new 1959 distribution of dwarfmistletoes in i elationrelation to topography on apache reserva- 1980 the mescalero mexico partridge and canfield com- tion new mexico journal of forestry 57 919 922 pared the incidence of several forest pests in 1968 ponderosa pine dwarf mistletoe in relation southern idaho estimated using roadside plot to topography and soils on the martoumanitoumamtou experimen- surveys and plots randomly located in areas tal forest colorado USDA forest service research without roads they reported no discernible note RM 107 4 appp 1977 the 6 class dwarf mistletoe rating system differences between the incidence of the pests USDA forest service research note RM 48 7 appp detected including dwarf mistletoesmistle toes for the 2 hawksworth F G AND A A LUSHER 1956 dwarf survey procedures because this study and oth- mistletoe survey of the mescalero apache indian ers indicate that roadside plot surveys provide reservation new mexico journal of forestry 54 390 similar estimates of dwarf mistletoe incidence 384 hawksworth FE G AND D WIENS 1972 biology and to surveys conducted away from roads we classification of dwarf mistletoesmistle toes arceuthobium recommend that resource managers continue USDA forest service agricultural handbook 401 to use roadside plot surveys for estimating 234 appp dwarf mistletoe incidence for national forests 1995 dwarf mistletoesmistle toes biology pathology and or other large forested areas however because systematics USDA forest service agricultural handbook 709 410 appp these surveys sample only a small fraction of HOFFMAN J T 1979 dwarfdwarfmistletoemistletoe loss assessment sur- the survey area they will provide only rough vey in region 4 1978 USDA forest service inter- estimates of the incidence and severity of mountain region forest pest management report dwarf mistletoesmistletoes R 4 79 4 12 appp HOFFMAN J T AND L HOBBS 1985 lodgepole pine acknowledgments dwarf mistletoe survey in the intermountain region plant disease 69 429 431 JOHNSON D W FE G hawksworth AND D B DRUM- we appreciate the field assistance provided MOND 1980 1979 dwarf mistletoe loss assessment by al dymerski valerie deblander carl survey on national forest lands in colorado USDA koprowski and lia spiegel reviews of the forest service forest pest management methods original manuscript by ralph williams greg application group report 80680 6 18 appp filip and catherine parks are appreciated JOHNSON D W FE G hawksworth AND D B DRUM- MOND 1981 yield loss of lodgepole pine stands to also dwarf mistletoe in colorado and wyoming national forests plant disease 65 437 438 literature CITED MAFFEI H M AND J S beatrybearryBEATTY 1988 changes in the incidence of dwarf mistletoe over 30 years in the ANDREWS S R AND J P DANIELS 1960 A survey of southwest pages 80 90 in proceedings of the 36th dwarfmistletoes in arizona and new mexico USDA western international forest disease work confer- forest service rocky mountain forest and range ence park city UT 19 23 september 1988 experiment station paper 49 17 appp MERRILL L M F G hawksworth AND D W JOHN- AVERY T E AND H E BURKHART 1983 forest measure- SON 1985 evaluation of a roadside survey proce- ments mcgraw hill book company new york 331 dure for dwarf mistletoe on ponderosa pine in col- PP orado plant disease 69 572 573 134 GREAT BASIN naturalist volume 56

PARMETERPARMFTER J R 1978 forest stand dynamics and ecologi- university of idaho moscow forestry wildlandWildwindlandland cal factors in relation to dwarf mistletoe spread and range experiment station note 34 7 appp impact and control pages 16 30 in proceedings of SNEDECOR G W AND W G COCHRANCOCHBAN 1989 statistical the symposium on dwarf mistletoe control through methods iowa state university press ames forestfoiest management USDAUS DA forest service general technical report PSW 31 received 13 february 1995 PARTRIDGE A D AND E R CANFIELD 1980 frequency accepted 23 october 1995 and damage by forest tree pests in southern idaho great basin naturalist 562 0 1996 appp 135 141

EFFECTS OF DOUGLAS FIR FOLIAGE AGE CLASS ON WESTERN SPRUCE BUDWORM oviposition CHOICE AND LARVAL performance

kimberlykimberlyakimberlynKimberiyalyAA doddsdodds1I1 karen M clancy2 kathryn J leyvasleyva3 david Greenberg3 and peter W price3priced

ABSTRACT the western spruce budworm chonstoneurachoristoneura occidentoccidentalisoccidentalistalis freemanFipreemaneeman prefers to feed on flushing buds and current year needles of douglas fir pseudotsuga menziemenziesnmenziesiimenziesiasnsiislisll birbmirb franco budworm larvae will not typically consume older age classes of needles unless all current yeaiyear foliage is depleted we tested the following null hypotheses 1 budworm larvae can feed on foliage with a wide range of qualities iei e current year versus 1 2 or01 3 year old needles without measurable effects on fitness and 2 budworm adults do not show any oviposition preference linked to the age of the foliage they fed on as larvae we used both laboratory and field experiments there was strong evidence to support rejection of hypothesis 1 budworm larvae had greater survival from the ath4th instar to pupal stage when they fed on current year foliage 43 52 survival versus older age classes of foliage 0 25 survival pupae from current year foliage were also heavier than pupae from 1 year old foliage there was weak evidence to support rejecting hypothesis 2 budworm adults that had fed on current year or 3 year old foliage as larvae preferred to oviposit on current year foliage similar conclusions were drawn from the laboratory and field experiments

key words chonstoneurachoristoneura occidentoccidentalisoccidentalistalis western spruce budworm oviposition preference needle age foliar quality eruptive species

the western spruce budworm choris of needles impact the budbudwormwornss fecundity toneura occidentoccidentalisoccidentalistalis freeman is a major defo- growth rate and survivorship mattson and liator of douglas fir pseudotsuga menziemenziesiimenziesiasii senberscriber 1987 clancy et al 1988 clancy 1991991bib birbmirb franco trees in western north amer- 1991c and may influence female oviposition ica fellin and dewey 1982 wulf and catesgates choices 1987 clancy et al 1988 budworm larvae pre- however the budbudwormwolasworaswormss oligophagous feed- fer to feed on the flushing buds and current ing behavior and eruptive population dynamics year needles of their host trees however if all suggest it is unlikely that there is a tight link- current year foliage is depleted larvae will age between female oviposition preference feed on older needles fellin and dewey 1982 and larval performance price et al 1990 talerico 1983 blake and wagner 1986 previ- female moths do not determine where their ous experiments by talerico 1983 and blake offspring will feed budworm adults lay eggs and wagner 1986 show older foliage is sub- on mature foliage in late summer furniss and optimal resulting in reduced fecundity higher carolin 1977 brookes et al 1987 upon hatch- mortality rates and impaired development ing the ist instarsmstars which do not feed dis- when budworm larvae are forced to feed on perse to sheltered locations ege g beneath bark only mature foliage they have reduced growth scales where they spin a hibernaculum and lower pupal weights and decreased survival overwinter when larvae emerge from their or they may not survive at all blake and wag- hibernaculahibernacular the following spring they disperse ner 1986 again typically on silken threads to find appro- variations in host foliage quality may influ- priate food sources the budworm s life his- ence the feeding and oviposition behavior of tory suggests that neither adults nor larvae the western spruce budworm clancy et al actively select host foliage based on differ- 1988 differences in levels of foliar nutrients ences in nutritional quality among individual water content needle toughness etc between host trees instead larvae passively disperse current year and older 1 year old age classes from their overwinteringoverwintering sites and may land

department of forestry northern arizona university box 15018 flagstaff AZ 86011 2rockybrockyocksoekyocky mountain forest and range experiment station USDA forest service researchReseaich 2500 S pine knoll drive flagstaff AZ 86001 3departmentepaitment of biological sciences northern arizona university box 5640 flagstaff AZ 86011

135 136 GREAT BASIN naturalist volume 56 on acceptable food sources once larvae are and2nd instars emerge from diapause feed through on a host tree they search for expanding cur the ath6th instar and pupate in late june or early rent year buds and needles if suitable foliage july adults ecloseecloise within 10 d our laboratory is not available larvae can disperse horizon- population of nondiapausing western spruce tally or vertically within and between tree budworm differs in that there is no overwin crowns and stands but dispersal invariably teringbering stage results in significant losses whether dispersing larvae live or die depends largely on METHODS whether they find hospitable sites brookes et field experiment al 1987 therefore the ability to utilize a broad range of foliage qualities would be to determine larval performance in the advantageous for budworm survival field we selected and tagged 50 douglas fir this study was designed to compare results trees of various sizes and ages on 1 2 june from laboratory and field tests of the null 1993 all tagged trees had abundant foliage in hypotheses that 1 budworm larvae can feed the lower crown sleeve bags made of fine on foliage with a wide range of qualities ie mesh screen were placed over 4 branches on current year versus 1 2 or 3 year old nee- each of the 50 trees and each bagged branch dles without measurable effects on fitness was randomly assigned to a foliage age class and 2 budworm adults do not show any current year I1 year 2 year or 3 year old oviposition preference linked to the age of the needles we removed by hand all needles that foliage they fed on as larvae furthermore we were not of the appropriate age class any wild wanted to determine if conclusions drawn budbudwormsworms present on the bagged branches from laboratory versus field experiments were were also removed similar this is important because many previ- on 4 and 8 june two ath4th or early ath5th instar ous studies conducted with budbudwormsworms and budworm larvae from our laboratory culture other forest defoliatorsdefolia tors have used clipped were placed on foliage inside each bagged foliage without knowing the effects this may branch a total of 400 larvae were used this have on foliar nutrition or host defenses by constituted the parental plP generation we conducting parallel experiments using intact have established that budworm larvae from and excised foliage from the same trees we our laboratory culture have rates of survival were able to evaluate the importance of and reproduction equivalent to wild bud changes in foliar quality that may be associ- worms when reared on douglas fir foliage in ated with bagging larvae on intact branches in the field leyva et al 1995 bags were closed the field versus feeding larvae excised foliage with string or duct tape at each end we exam- in the laboratory ined the bagged branches on 20 june to determine if sufficient foliage remained for comple- STUDY AREA AND ORGANISMS tion of larval development pupae were not observed at this time the study area is located at little springs budworm larvae remained in the field until elevation 2560 m 16 km north of flagstaff about half of them had pupated and then the arizona within the coconino national forest bagged branches were clipped placed inside the site is a high elevation mixed conifer for- large plastic bags and transported to the labo- est with douglas fir as the primary host species ratory pupae were weighed to the nearest 010.1oi and with a recent history of western spruce mg sorted into trays according to treatment budworm infestation foliage age class and sex and then refriger- the western spruce budworm has a univol- ated at IOCloc10 C until we obtained 10 males and tine life cycle adults are present from july to 10 females from the same treatment larvae august with mating typically occurring within that had not pupated were placed in labeled 24 h of eclosion eggs are laid soon after mat- petri dishes lined with moist filter paper dou ing females lay between 25 and 40 eggs per glas fir foliage of the appropriate age class was egg mass brookes et al 1987 eggs hatch in provided for them to feed on until they about 10 d after dispersing to sheltered loca- pupated this foliage was collected at random tions and spinning a hibernaculum larvae molt from tagged douglas fir trees at the study site into and2nd instars and overwinter in spring the foliage was replaced every 2 3 d to ensure 199619961 EFFECTS OF FOLIAGE AGE CLASS ON BUDWORMS 137 freshness petri dishes were checked each mon- foliage used in this experiment was collected day wednesday and friday to remove and from the same 50 trees we used for the field weigh new pupae these were also sorted and experiment four hundred ath4th instar bud refrigerated worms were placed on excised foliage in petri when 10 pairs of male and female pupae dishes lined with moist filter paper 2 larvae were available from a treatment they were per dish fifty petri dishes were used per placed in a brown paper mating bag oviposi- foliage age class treatment current year 1 tion preference tests for both field and labora- year 2 year and 3 year old needles corre- tory experiments were conducted in the labo- spsponding to the 50 trees used in the field ratory brown paper bags provided appropri- experiment needles of the appropriate age ate lighting conditions both for mating which class were left attached to the stem to prevent occurs from 2000 to 2300 h in nature when desiccation of foliage foliage was replaced only safety lights were on in the laboratory at every 2 3 d to ensure freshness petri dishes night and for oviposition which normally were labeled according to the tree number occurs the day following mating when all lights and foliage age class if a single larva or both were on during the day ie bags are not larvae in each petri dish died before pupation opaque bags were checked every other day they were replaced with new larvae from our until 5 or 6 moths emerged then branches of laboratory culture otherwise we used the same freshly clipped douglas fir foliage were added procedures for the laboratory experiment as for oviposition substrate once foliage was for the field experiment added moths were allowed to mate and oviposit for 7 8 d after oviposition occurred douglas RESULTS fir branches removed for were and inspected effects of foliage age class on egg masses F egg masses sorted these were PIP survival and pupal weight according to treatment foliage age class to determine if female moths showed a preference for budworm larvae that consumed current ovipositing on a particular age class of foliage year needles of douglas fir in the field experi- the F egg masses collected were surface ment had higher survival rates from ath4th instar sterilized with formalin and placed into labeled to pupal stage 43 survival compared to lar- cups containing an artificial diet nutritionally vae that fed on 1 year old 2 survival 2 year old 1 survival 3 old 0 similar to douglas fir foliage clancy 1991991ala or year survival aa qc2xa 3 F larvae were reared on the diet until the ath4th needles fig 1a x2 13019130.19 df P 0.001 400 we believe of or early ath5th instar stage after which they were 0001 n that many the larvae bagged on the branches with only placed in labeled petri dishes lined with moist 1 year old needles to feed on escaped from filter paper douglas fir foliage of the same the mesh bag enclosures so it may be more age class that their parents consumed was pro- appropriate to refer to this response as per- vided for them to feed on until they pupated cent larvae accounted for rather than percent this foliage was collected random from at tagged larval survival larvae from older foliage age douglas fir the study trees at site these lar- class treatments were more likely to escape vae placed field were not in the because it was because budworm larvae tend to disperse when too late in the season for conditions suitable suitable food is not available and our bags for budworm development foliage was replaced were not so tightly sealed that larvae could not every other day to ensure freshness F larvae wriggle out through small openings along the were reared on foliage within petri dishes seams or closures at the ends until they pupated pupae were handled in the the age class of foliage ingested had a simi- same manner as in the first generation lar effect on survival from ath4th instar to pupal laboratory experiment stage in the laboratory experiment fig ab1b xax22 594659.46 df 3 P 00010.001 n 727 this study was conducted to determine if approximately 52 of larvae that consumed laboratory experiments using excised foliage current year needles survived survival was would yield results similar to those from field 25 for larvae consuming 1 year old needles experiments using intact foliage the experi- 18 for larvae feeding on 2 year old needles ment was started 24 june 1993 douglas fir and 20 for larvae eating 3 year old needles 138 GREAT BASIN naturalist volume 56

50- A loo100 120 40-

0 loo100 30 E

Q malemaie 20- U a 80 female 10 0 n loo100 n loo100 CL 60 0 gZ current yr 1 y ro 2 yr old 3 yr old CL iliillell W 40 60 n 146 B cliondCM

1 50 20 1 40

30 195 0 0 0 0 n 192 current yr 1 yr rooldoid 2 yr oldoid 3 yr oldoid 20 194 10 foliage age class 0 yr 1 1 current yr oldoid 0 ro 2 yr old 3 yr old I fig 2 mean 2 s or 95 confidence interval P male 0 and female 0 pupal weight for larvae reared foliage age class from the ath4th instar to pupation on foliage of different age t classes in the field experiment ANOVA tests showed that fig 1 percentage of P ath4th instar western spruce bud foliage age class did not affect pupal weight FP 01520.1520 152 worms surviving to the pupal stage when reared on cur and that females weigh more than males FP 00010.0010 ooi001 bars rent 1 2 and 3 year old douglas fir needles for the A without standard errors had a sample size ofofalofnln 1 field experiment and B laboratory experiment x2xa2 tests showed that survival varied among the foliage age classes for both the field P 00010 ooi001 and laboratory P 00010 oolooi001 a dependence between age class of foliage on experiments numbers above the bars indicate sample which the P female was reared and age class sizes iei e number ofbudwormofbudwoim larvae used peipelpefper tieatmenttreatment of foliage on which the moths laid their F egg masses P 00170.0170 017 table 1 this was not a very foliage age class did not have a significant strong test of the hypothesis because there effect on pupal masses for the field experi- were many zero values inm the table which ment F 1971.97 df 241 P 01520.152 fig 2 required that 1 year 2 year and 3 year old this inability to detect differences among age class columns and rows be combined to foliage age classes can be attributed to the meet requirements for minimum expected cell very small sample sizes n 0 2 for 1 year frequencies furthermore many F egg masses old foliage As expected female pupae were were very small areas 3 mm2mma such small heavier than male pupae F 203920.39 df egg masses are nearly always inviable and are 1411.41 P 00010.001 probably aberrant leyva et al 1995 nonethe- there were detectable differences in pupal less the distribution of F egg masses indi- masses among different foliage age classes for cated that moths reared as larvae on current the laboratory experiment F 364736.47 df year or 3 year old foliage laid more of their F 3182 P 00010.001 fig 3 larvae consuming egg masses on current year needles than on current year foliage became much heavier older age classes of needlesofneedles pupae than larvae feeding on 1 year old effects of foliage age class foliage once again female pupae were bigger on F survival and pupal weight than male pupae F 147014.70 df 1182 P 00010.001 only 2 FIF egg masses were produced from the field experiment this precluded analyz- effects of foliage age class on ing data on survivsurviasurvivalaI1 or upalppupal weweightsightsbights for this oviposition preference of P females experiment for the laboratory experiment we sample sizes for the field experiment were found a significant difference in survival from not large enough for data analysis n 2 egg ath4th instar to pupal stage between larvae reared masses but a contingency table analysis of on current year 83383383.383 3 survival versus 3 year data from the laboratory experiment indicated old 30830830.830 8 survival needles xax2 117811.7811 78 df 199611996 EFFECTS OF FOLIAGE AGE CLASS ON BUDWORMS 139

1 70 fig and pupal masses figs 2 3 declined for larvae feeding on I1 year oldoid needles male compared to larvae feeding on current year E 60 E female foliage the same patterns in relation to the effects of foliage age on survival were evident M for both the P and F generations of the labo- 50 ratory experiment it is well established that the nutritional quality of douglas fir needles 40 declines rapidly as current year needles age cn clancy et al 1988 1995 furthermore N clancy et al 1995 point out that 1 30 the general pattern for I1 year or older needles of conifers is typically an extension of the seasonal 20 trends for nutrient concentration changes in cur current yr 1 yrgroldold 2 yr oldoid 3 yr old rent year needles foliage age class needle toughness and fiber content also increase as foliage matures thus making older 3 mean 2 s fig or 95 confidence interval P needles less suitable food for the budworm male 0 and female 0 pupal weight for larvae reared from the ath4th instar to pupation on foliage of different age on the other hand the fact that budworm lar- classes in the laboratory experiment ANOVA tests vae could survive at all when reared on older showed that foliage age class had a significant effect on age classes of needles may indicate that their pupal weight P 0 oolooi001 as did sex P 0.0010 001 0001 0001oolooi nutritional niche is indeed broad as suggested by price et al 1990 and leyva et al 1995 we found less convincing evidence to 1 FP 000060.0006 n 44 sample sizes were 0 support rejection of hypothesis 2 but nonetheless for 1 year and 2 year old needles this result conclude that budworm adults show was consistent with results for the P genera- we may an oviposition preference that is linked to the tion in that survival was higher for larvae age of the foliage they fed on as larvae table reared on current year foliage than on 3 year 1 budworm females that fed as larvae on old needles cur rent year foliage laid more egg masses on cur however IT pupal masses were equivalent rent year needles than on older needles females for pupae from current year and 3 year old reared on 3 year old foliage also laid more egg foliage F 1141.14 df 119 P 02990.299 As masses on current year needles this result is before female larger than male pupae were surprising because the budworm typically ovi pupae 6.01601goi F 601 df 119 P 00240.024 posits on mature foliage brookes et al 1987 price et al 1990 however many egg masses discussion from our experiment were very small and are most likely aberrant leyva et al 1995 thus although budworm s the western spruce we suspect this may not represent normal life history and population dynamics suggestsuggest oviposition behavior for the budworm if we that larvae should be able to utilize a broad remove these very small egg masses from the range of foliage qualities our results confirm data set most egg masses were laid on 3 year previous studies that indicate the budworm is old needles indicating the budworm does not well adapted to feeding on 1 year old or indeed prefer to oviposit on mature foliage older needles talerico 1983 blake and wag- an alternative explanation may be that female ner 1986 thus we must reject hypothesis I1 moths distribute egg masses randomly across and conclude that the budworm cannot feed age classes of needles available current year on foliage with as wide a range in qualities as needles represent a small proportion of total is found in current year versus 1 year old needles present under natural conditions and needles without measurable effects on fitness they may be nearly absent when defoliation is we found that whether budworm larvae were heavy thus our result could be an artifact of feeding on bagged foliage in the field or on providing an atypical distribution of needle excised foliage in the laboratory larval survival age classes for oviposition substrate 140 GREAT BASIN naturalist volume 56

TABLE 1 distribution of F egg masses of the western spruce budworm laid on current 1 2 or 3 year old needles of douglas fir in relation to the age class of the foliage on which the pi moths were rearedalearedareared3 age class of foliage on which PPI female was reared age classi ofr foliager i 1 on which FFI egg masses were laid current year 1 year old 2 year old 3 year old current year 27 0 0 14 I1 searyear old 6 0 0 0 2 year old 5 0 0 1 3 year old 11 0 0 0

dataadata aiaree the ji timbenumber of fjF egg masses fromhrombromfi om the laboratory experiment the distribution of egg masses was examined for row column dependence using a 2 x 2 contingency table current yeaiyealyear veitusversus S I1 year oldaldaid foliage older foliage age classes weiewere combined inin aiderorder to meet requirements for minimum expected cell frequenciesfiequencies witeswatesyates conectedcorrectedconnected x2xa 57255 725225 df 1 P 00170 017 it 64 ITP budwormm larvaeiiilai vae were i earedreared on foliage of different age classes from the ath4th instar to pupation

conclusions from the field versus labora- acknowledgments tory experiments were similar in regard to the effects of foliage age class on larval survival we thank daniel huebner and kenneth and pupal masses this indicates that foliar qual- dodds for assistance in collecting data and ity does not change dramatically when foliage conducting experiments we also thank D is excised at least not over a 2 3 d period leatherman and an anonymous reviewer for nor does intact foliage on bagged branches their comments on an earlier draft of this change markedly in terms of a local or systemic paper our western spruce budworm colony induced response to budworm defoliation was started with egg masses obtained from the it is noteworthy that pupae from the current canadian forest service s pest management year foliage treatment in the field experiment institute in sault ste marie ontario this re- were heavier than equivalent pupae from the search was funded in part by the rocky moun- laboratory experiment figs 2 3 this differ- tain forest and range experiment station ence may well be related to the 15 19 d delay cooperative agreement 28 c3ca 708 between between the start of the field experiment ini- KMC and PWEPWP tiated on 4 and 8 june 1993 and the beginning of the laboratory study initiated on 24 literature CITED june 1993 with the concomitant decline in nutritional quality of the expanding current BLAKE E A AND A R WAGNER 1986 foliage age as a factor in food utilization by the western spruce bud- year foliage alternatively current year needles worm choristoneurachonstoneura occidentoccidentalisocczdentalisoccidentalistalisallsails great basin natu- remaining on bagged branches could have ralist 46 169 172 acted as a local nutrient sink in the absence of BROOKES A H J J COLBERT R G MITCHELLMlTCHELL AND R W competing older needles which were removed STARK EDITORS 1987 western spruce budworm USDA forest service technical bulletin 1694 larvae feeding on bagged branches with thus CLANCY K A 1991a multiple generation biobioassayassay for current year foliage may have benefited from investigating western spruce budworm lepidoptera this improved nutritional quality whereas lar- tortricidae nutritional ecology environmental ento- vae that were fed clipped foliage in the labora- mology 20 1363 1374 have received this 1991b douglas fir nutrients and terpenesterterpeneypenes as tory experiment would not potential factors influencing western spruce budworm nutritional boost defoliation pages 123 133 in Y N Baranchikbaranchikovov W J survival rates were higher overall in the mattson F hainham and T L payne editors forest laboratory experiment than in the field experi- insect guilds patterns of interaction with host trees USDA forest service general technical report NE ment fig 1 we attribute this difference to a 153 combination of factors for example larvae on 1991c western spruce budworm response to dif- bagged branches in the field were exposed to ferent moisture levels in artificial diets forest ecol- some predation since the bags were not per- ogy and management 39 223 235 fect barriers also some budworm larvae un- CLANCY K M A R WAGNER AND P B REICH 1995 herbherbivory pages 125 180 doubtedly escaped from the bags and weather physiologyEcoecophysiology and insect herbivoreivory in W K smith and T M hinckley editors eco could have played a role in the lower survival physiology of coniferous forests academic press of larvae in the field new york 199611996 EFFECTS OF FOLIAGE AGE CLASS ON BUDWORMS 141

CLANCY K M M R WAGNER AND R W TINUS 1988 PRICE P W N COBB T P CRAIG G W FERNANDES J K variation in host foliage nutrient concentrations in ITAMI S MOPPER AND R W PRESZLER 1990 insect i elationrelation to western spruce budworm herbivoreherbherbivoryivory herbivore population dynamics on trees and shrubs canadian journal of forest research 18 530 539 new approaches relevant to latent and eruptive FELLIN D G AND J E DEWEY 1982 western spruce species and life table development pages 1 38 in budworm USDA forest service forest insect and E A bernays editor insect plant interactions vol- disease leaflet 53 ume 2 CRC press boca raton FL FURNISS R L AND V M CAROLIN 1977 western forest TALERICO R L 1983 summary of life history and hosts of insects USDA forest service miscellaneous publi- the spruce budbudwormsworms pages 1 4 in proceedings cation 1339 forest defoliator host interactions a comparison LEYVA K J K M CLANCY AND P W PRICE 1995 perfor- between gypsy moth and spruce budworm USDA mance of wild versus laboratory populations of west- forest service general technical report NE 85 ern spruce budworm lepidoptera tortricidae WULF N W AND R G CATESGATES 1987 site and stand char- feeding on douglas fir foliage environmental ento- acteristicsacteristics pages 89 115 in M H brooks R W mology 24 80 87 campbell J J colbert R G mitchell and R W MATTSON W J AND J M SCRIBER 1987 nutritional stark editors western spruce budworm USDA ecology of insect folivoresfolivores of woody plants water forest service technical bulletin 1694 nitrogen fiber and mineral considerations pages 105 146 inm FE slansky jijr and J rodriguezRodnguez editors received 20 october 1995 nutritional ecology of insects mites spiders and accepted 27 february 1996 related invertebrates john wiley and sons new york cleatgleatgreat basin natuiahstnaturalist 562 C 1996 appp 142 149

trypanoplasmaatrariatrypanoplasma ATRARIA SPSE N KINETOPLASTkinetoplastidaIDA BODONIDAE IN FISHES FROM THE SEVIER RIVER DRAINAGE UTAH

J stephen CranCranneyneyl1 and richard A Heckmann 2

ABSIRACTABSTRACT A total of 181 fishes belonging to 10 species were captured near richfield utah and examined for parasites A new species of hemoflagellate trypanoplasma atrariaadranaatrana sp n was observed in 3 species utah chub gila attana girard ledsideredsidebedside shiner bichardRichardrichardsoniusrichardsomusRichardssoniusomus balteatusbalteatus richardson and speckled dace rhinichthysrhimchthys esculusosculus girard seven otherothel species of fishes examined in the study area were negative for T atrariaadranaatrana sp n the salmonid leech Pispiscicolapescicolacicola salmositica meyer collected in the same area harbored developmental stages of trypanoplasma suggesting a possible leech vector for the hemoflagellate characteristics of trypanoplasma atrariaadranaatrana sp n place it near T salmositica but the new species is twice as large

keykeywordswords trypanoplasma atrariaadranaatrana n sp blood parasites gila atratrariaadranaatranaariaarla fish parasites

trypanoplasma is a biflagellated protozoan copulation while a leech vector was necessary found in the blood of freshwater fishes in the to transfer the flagellate from the blood of one united states it has caused significant mor- fish to another putz 1970 submitted that tality in rainbow trout oncorhynchus mykiss comparative biological studies between simi- waiWalwalbaumbaumlbaumi and king salmon 0 tshawytscha lar morphological types are necessary for a walbaum under hatchery conditions becker correct taxonomic classification use of the and katz 1966 wales and wolf 1995 this genus cryptobiaCryptobia has in most cases emerged genus has also been described from the blood as the popular choice and trypanoplasma is of marine fish strout 1965 another name for generally recognized as a synonym recently the blood biflagellate of salmonsaimonsalmondssalmomdssalmonidsids described lomlornlormlomm and dakovadykova 1992 used trypanoplasma above is Cryptcryptobiaobia there are differing opinions for biflagellated blood inhibiting parasites of on the use of the two genera Cryptcryptobiaobia and fishes in which a leech vector is involved thus trypanoplasma but these differences have been we adopted the classification scheme used by recently clarified by lomlornlormlomm and dakovadykova 1992 lomlornlormlomm and dakovadykova 1992 the genus Cryptcryptobiaobia was first proposed by four species of trypanoplasma from the leidy 1846 for flagellatedbiflagellatedbi protozoaprotozoansns occur- blood of freshwater fishes have been reported ring as parasites in the seminal vesicles of snails in north america mavor 1915 found T borreli chalachmkowchalachnikow 1888 was the first to record in a moribund white sucker catostomus com the parasite in the blood of fishes observing it mersonigersoni lacelelacepeLacepe from lake huron the iden- in freshwater loachescoaches in russia laveran and tification of T borreli was based on similarities mesnil 1901 established the genus tropanotrypano with the species initially described by laveran plasma for a biflagellated blood parasite from and mesnil 1901 katz 1951 recorded C freshwater fishes in france in 1909 crawley trypanoplasma salmositica from silver stated that cryptobiaCryptobia from snails and tropanotrypano salmon 0 kisutch walbaum and C try plasma from fishes were morphologically iden- panopanoplasmaplasma lenchilynchi from cottids in the state of tical and that Cryptcryptobiaobia had taxonomic prior- washington subsequent transmission studies ity in defending the creation of the genus try showed C lenchilynchi to be a synonym of C salmo panopanoplasmaplasma laveran and mesnil 1912 argued siticabitica becker and katz 1965a laird 1961 that morphological similarities were not suffi- described C trypanoplasma gurneyorumgurney orum cient criteria for maintaining a single genus from northern pike esox lucius linnaeus when strong biological differences such as and from 2 salmonidssalmon ids lake whitefish core- method of infection were evident the para- gonus clupeaformisclupeaformis mitchill and lake trout sites in snails were transferred directly during salvelinus namaycush walbaum

utahlutaliI1 division of wildlife resources dudlesDudicsduchesnene UT 84021 2 fpditinentpahentofzo1ogyof zoology brighbiightim yougyoung uiversityuniveisitydiversity fiovopiovopro UT 84602

142 199619961 trypanoplasma ATRARIA BLOOD PARASITE OF FISH 143 A N

AREA I1 UTAH RICHFIELD 90p AA FISH HATCHERY AREA 2

seylerSEVIER RIVER CP 4 6570 BULL CLAIM HILL

AREA 3 V

ANNABELLA CENTRAL

ELSINORE

3 MORE collection sltesSITES I1 km AT approximately EQUAL 0 INTERVALS 0 collection sltesSITES

fig 1 map of the study aleaarea near richfield utah showing collection sites on the sevier river and location of the spring ponds areas 1 2 and 3 near bull claim hill

another species C trypanoplasma cata ence of external flagellates on the gills of carp rachaeractae was described by putz 1972a from cyprinus carpio linnaeus in pennsylvania several cyprincyprinidsids in west virginia this record the use of the scientific name trypanoplasma also included the first comprehensive study of is accurate for these observations lormlommlomlorn and a Cryptcryptobiaobia trypanoplasma species that en- dakovadykova 1992 khan and noble 1972 and compassed comparative morphology mode of khan 1991 recently reported on another transmission natural and experimental hosts species of cryptobiaCryptobia C dahlidagli in viro and in vitro culture histopathology involvement of a vector in transmission of and cryopreservation these criteria and exten- Cryptcryptobiaobia trypanoplasma was postulated sive comparison with T salmositica from the by mavor 1915 katz 1951 observed devel- west coast were used to justify designation of opopmentalmental stages of cryptobiaCryptobia from the gut of T cataractcataractalcataractaeae as a valid species the leech Pispiscicolapescicolacicola salmositica and indicated it an ectoparasitic relationship of trypanoplas as a vector for C salmositica subsequent ma on goldfish carassiusCaras sius auratusauranus linnaeus experiments showed conclusively that the leech maintained in aquaria was recorded by swezy functioned as a vector in the transfer of C 1919 wenrich 1931 also observed the pres salmositica from fish to fish becker and katz 144 GREAT BASIN naturalist volume 56

TABLETABLL 1 prevalence of trypanoplasma sp in fish examined from the main sevier river northern spring ponds and southern spring ponds east of richfield utah fish number positive percent aleaarea species examined infections positive main sevier river cilacliagila copei 10 0 cilacliagila atrariaatwaria 2 0 RichardrichardsoniusrichardsomusRichardssoniusomus balbalteatusteatus 28 0 rhinichthysrhimchthys esculusosculus 1 1 100 coituscottus bairdi 1 0 salmo trutta 2 0 northern spispringing ponds gila atrariaatwaria 20 20 100 RichardrichardsoniusrichardsomusRichardssoniusomus balteatusbalteatus 20 20 100 rhinichthysrhimchthys osculusesculus 20 20 100 oncorhynchus mykiss 10 0 cyprinus carpio 10 0 catostomus ardensaddens 10 0 southern spring ponds gila atrariaatwaria 20 6 30 RichardrichardsoniusrichardsomusRichardssoniusomus balbalteatusteatus 20 0 rhinichthysrhimchthys osculusesculus 5 0 ameiurusAmeturus melas 2 0 totals all areas cilacliagila copei 10 0 gila atrariaatwaria 42 26 62 RichardrichardsoniusrichardsomusRichardssoniusomus balbalteatusteatus 68 20 29 rhinichthysrhimchthys osculusesculus 26 21 81 oncorhynchus mykiss 10 0 cyprinus carpio 10 0 catostomus ardensaddens 10 0 salmo trutta 2 0 ameiurus melas 2 0 coituscottus bairdi 1 0

1965a 1965c burreson 1982 putz 1972b MATERIALS AND METHODS showed a leech cystobranchus virginiousvirginicus to virginicus study area be a vector for T cataractcataractalcataractaeae organisms of the genus Cryptcryptobiaobia and try the primary collection site located approx- panopanoplasmaplasma have been reported as parasites in imately 5 km east of richfield utah was sub- marine and freshwater fishes amanderssalamanderssalamanderisal divided into 3 major areas fig I1 the main frogs heteroheteropodsheteropodapods planariansplanarians siphonophores sevier river area 1 northern spring ponds chaetognathschaetognatha leeches mole crickets lizards area 2 and southern spring ponds area 3 snails and also as free living forms noble the ponds are located east of the sevier river 1968 at the base of bull claim hill the springs are woo and wehnertweimert 1983 and bower and rocky and contain dense stands of watercress margolis 1983 reported that trypanoplasma and other aquatic plants the river is heavily and Cryptcryptobiaobia of many species of fish can be silted and almost dry during the summer fish acquired directly via water and not only by were also examined from source waters of a leeches bower and margolis 1984 and woo fish hatchery in the northern spring area and 1987 also considered trypanoplasma a syn- from 7 stations on the sevier river south of onym of cryptobiaCryptobia a view not helped by the principal study area to determine the local becker and katz 1966 or lomlornlormlomm 1979 prior to range of the hemoflagellate this time collection and examination of fish the species of trypanoplasma described in this article was first observed by mcdaniel in A total of 181 fish representing 5 families 1970 personal communication from utah chub and 10 species were collected and examined gliacilagila atrariaatratwariaaria near richfield utah at that time for blood flagellates trypanoplasma and crapcryp it was considered a species of Cryptcryptobiaobia tobia using the kidney strike technique 199619961 trypanoplasmatkypanoplasma ATRARIAATRAWA BLOOD PARASITE OF FISH 145

TABLE 2 natural hosts vectors and references of trypanoplasma sppapp from freshwater fishes of north americaamelicaamenea natural hosts species vector fish references trypanoplasma atrariaadranaatrana sp n Pispiscicolapescicolacicola salmositicasalmositzca gila attatratrariaadranaatranaariaaflaafia RichardrichardsoniusrichardsomusRichardssoniusomus balbalteatusteatus present study rhinichthysrhimchthys osculusesculus

T cataractcataractalcataractaeae cystobranchus virgimcusvirginicusvirginiousvirginicus rhinichthysrhimchthys cataractcataractalcataractaeae rhinichthysrhimchthys putz 19701970197219701972a1972aa strastratulustulus exoglossum maxilhnguamaxilmaxillingualingua 1972b Campocampostomastoma anoanomalummalum

T salmositica Pispiscicolapescicolacicola salmositica oncorhynchus kisutch cottus katz 1951 wales and rhotheus cottus aleuticus wolf 1995 becker and oncorhynchus mykiss oncorhynchus katz 1965b19661965b 1966 putz tshawytscha salmo trutta catostomus 1972a 1972b becker snysnydenderiderl oncorhynchus keta and katz 1977 oncorhynchus gorgorbuschagorkuschabuscha prosopium wilwilliamwolwoihamsomsoni cottus bairdi cottus gulosusgulosus cottus beldingibeldingi cottusgottus perperplexesperplexusplexus cottus asper rhinichthysrhimchthys cataractcataractalcataractaeae gasterosteus acuaculeatusleatus

T gureneyorum none given coregonus clupeaclupeaformisformis delmusvelmussalvelinussaiSal laird 1961 namaycush esox lucius

T borreli none given catostomus commersonicommetcommersoncommercommersomcommersomsoni mavor1915mavor 1915

putz 1970 hemoflagellates were detected by A smear from the solution was stained follow- characteristic whiplike motions of the flagella ing the fish blood procedure living tropanotrypano examination of stained preparations at higher plasma were observed in wet mounts from magnification confirmed infections and per- infected fish and mortared leeches to deter- mitted morphological studies mine behavioral characteristics collection and identification morphometricsMorpho metrics of leeches stained slides were examined at a magnifi- ectoparasitic leeches of fishes were col- cation of 1000x measurements were recorded lected from the underside of rocks in the 2 for anterior and posterior flagella lengths spring areas and identified using hoffman body length and width kinetoplast length and 1967 specimens were confirmed by dr roy width of the nucleus fifty organisms were W sawyer biology department college of measured and averages compared with exist- charleston south carolina leeches were ing measurements of other described species maintained in the laboratory at 4 C in cov- cryptobiaofcryptobiaof Cryptobia and trypanoplasma ered paper cups where they could be kept in good condition for up to 3 mon RESULTS mounting and staining natural hosts blood was obtained from the caudal pedun- examination of 181 fish at 15 stations re- cle of infected fishes samples of hemhempoeticpoetic vealed trypanoplasma in utah chub cilacliagliacilagila tissue were also taken directly from the kidney atrariaatratwariaaria redsidebedside shiner richardsoniusRichardsonius balteabaldea kidney strike A thin smear was prepared tus and speckled dace rhinichthyes osculusesculusosculus on a glass slide air dried fixed with methyl seven species table 1 appeared to be negative alcohol 100 and stained with giemsagiemza for the blood flagellate utah sucker catosto- humason 1967 mus ardensaddens jordan and gilbert black bull- stained smears from leeches were prepared head ameiurus melas rafinesquerafinesquejRafinesque rainbow by mortaring each leech in a small amount of trout oncorhynchus mykiss brown trout hank s balanced salt solution hoffman 1967 salmosalmo trutta linnaeus carp cyprinus 146 GREAT BASIN naturalist volume 56

3 b 7

5 PM 5 pm

d a

fig 2 trypanoplasmatrypanoplasrna atrariaatwaria sp n from fishesbishes a and a leech vector b c d 1 anterior flagellum 2 blepharoplast 3 kinetoplast 4 nucleus 5 posterior flagellum b both flagella in anterior position c posterior migration of flagellum d commoncornmon stage in leech with short posterior flagellum capio leatherside chub cilacliagilacilagila copei jordan sites reported natural hosts and vectors of and gilbert and mottled sculpin cottuscoituscottus described species of trypanoplasma and crapcryp bairdi girard rainbow trout carp and utah tobia from north america are given in table 2 sucker all came from the northern springs prevalence ponds area 2 while the leatherside chub of trypanoplasma in the richfield utah area brown trout and mottled sculpin were only in the sevier river utah chub and speckled fish infected with trypanoplasma were with dace were abundant in the springs but only 2 1 exception obtained in the 2 spring areas along chub and I1 speckled dace were collected from bull claim hill table 1 one speckled dace the sevier river the 2 black bullhead were was collected where I1 of the northern springs from the southern spring ponds area 3 only emptied into the sevier river in area 1 all redsidebedside shiner was abundant at all collection individuals of the 3 host species were infected 199619961 trypanoplasma ATRARIAATRAWA BLOOD PARASITE OF FISH 147

at area 2 the parasite was present in 30 of utah chub and absent in speckled dace and redsidebedside shiner table 1 microscopic examina-examina tion of kidney thudsfluids from northern spring fishes revealed 3 4 flagellates per field at loox A for the southern springs examination of several fields at the same magnification was necessary to locate a single parasite indicating a much lower level of infection in that area vector the parasitic leech recovered in the study area was identified as Pispiscicolapescicolacicola salmositicasalmositical a common ectoparasite of fish in freshwater streams of the west coast of the united states hoffman 1967 microscopic examination of the mortared leech preparation revealed sev- F eral developmental stages of trypanoplasma which were all morphologically different from 7 J the stage in the fish 2 parasite in fig this cor- 4 relates with observations by other workers in the field lommlomlorn and dakovadykova 1992 Pispiscicolapescicolacicola salmositica was observed from the northern springs ponds and the northernmost portion of the southern area extensive search fig 3 trypanoplasma atrariaadranaatrana sp n note erythrocyte of the remainder of the southern springs and B flagella F nucleus N and body of protozoan sevier river produced no additional specimens 1000x magnification of the leech leech prevalence was high in autumn and continued until peak numbers were observed in the middle of february in by trypanoplasma atrariaadranaatrana sp n under phase late march to july only a small number of microscopy revealed a high degree of poly- leeches were observed morphism and constant whiplike undulatory rainbow trout carp sucker and utah utah movement stages in the leech exhibited a chub were hosts for P salmositica leeches qivering motion with much less distortion of observed were never on redsidebedside shiner or body shape the most common stage visible inm speckled dace the leech had a short posterior flagellum and description of trypanoplasma was less than 12 the overall size of that atrariaatranaadrana sp n observed from the fish host fig ad2d fig 3 discussion average parameters given in micrometers with ranges in parentheses of 50 stained speci- published host records for trypanoplasma mens of trypanoplasma atrariaadranaatrana sp n are as in north america include 25 species of fresh- follows body length 3053030.55 2736273627.3627 36 body width water fishes putz 1972a trypanoplasma salmo 454.54 5 3 7 length of anterior flagellum 29229.229 2 siticabiticasitzca is reported to parasitize 19 species T 23 34 length of posterior flagellum 209 cataractcataractalcataractaeae 4 T gurneyorumgurney orum 3 and T borreli 15 24 nuclear width 272.72 7 2 353.53 5 kinetoplast only a single host species results of this study length 5955.99 45454.54 5 7 type specimens including showed T atrariaadranaatrana in 3 cyprincypnnidscyprinidsids utah chub paratypesparatypes have been deposited USNM hel- redsidebedside shiner and speckled dace minthologicalminthological collection nos 74436 and 74437 the only known vectors of trypanoplasma with additional paraparatypestypes in the junior author s are parasitic leeches two species have been collections morphometncmorphometricMorphometric comparisons with demonstrated as vectors in north america other described species of trypanoplasma Pispiscicolapescicolacicola salmositica as a vector of T salmo from north america are shown in table 3 siticabitica see becker and katz 1965a and 148 GREAT BASIN naturalist volume 56

TABITABLE L 3 Morphomorphometricmoiphometncmetric comparison of trypanoplasma atrariaadranaatrana sp n ranges in parentheses with other species of try panoplasma1panoplasmail described from the blood of north american freshwater fishes all measurements in micrometers total length of anterior length of posterior nuclear kinetoplast species length width flagella flagella width length tryanoplasma atrariaadranaatrana sp n 30530.530 5 27 36 4534 5 3 7 29129.129 12323 34 20920.920 9 15 24 272 7 202 030 353 5 595.95sg 9 45454.54 5 707.07 0

T cataractcataractalcaturactaecataractaeae 17 2 11 14 10 15 26 31

T salmositica 1494 246 1605 896 15 35 458

T gureneyorum 251 67 19 10 none given none given

T borreliborrehborrah 20 25 3 4 none given none given none given none given

usually a leech iraceirhcerheirhet e isis a close relationshipu lationship between the two blood flagellateflagellates cryptobiacryptobtaCryptobia and trypanopkwwtrypanoplasma species of trypanuplasmatrypanopla&m are transmitted by vector

cystobranchus virginicusvirgivirginiousnicus as the vector of T BOWER S M AND L MARGOLIS 1983 direct transmission among ae see putz 1972a salmonid of the haemoflagellate cryptobiaCryptobia salmositica cataractcataractalcataractae the pacific salmon oncorhynchus sppapp canadian jour- leech Pispiscicolapescicolacicola salmositica is probably the nal of zoology 61 1242 1250 hemoflagellate vector in this study no direct 1984 detection of infection and susceptibility of transmission experiments were conducted but different pacific salmon stocks oncorhynchus sppapp leeches were observed parasitizingp4rasitizing fishes at to the haemoflagellatehaernoflagellate CryptCryptocryptobiacryptobwbwobia salmositica journal ofofparasitology70parasitology 70 273 278 the collection sites and trypanoplasma was BURRESON E M 1982 the life cycle of trypanoplasma observed in leech guts the protozoan appears bulbullockibullockylocki zoomastigophorea Kinetoplastkmetoplastidakinetoplastidaidalda journal to undergo developmental changes within the of protozoology 29 72 77 leech with the trailing nagflagellumellumeliumeilum migrating chalachnikow A P 1888 recherches sur les parasites anterior to posterior and forming the undulat- du sang arkhivesarkbivesarchivesArkhivesbives veterinary science st petersburg I1 65 membrane 2 size of the flagel- ing fig the COPE 0 B 1958 incidence of external parasites on cut- late in the leech was about 13 to 12 that of throat trout in yellowstone lake proceedings of the the parasite in the fish host becker and katz utah academy of science arts and letters 35 igsa165a reported P salmositica as endemic to 95 100 the pacific coast of north america cope CRAWLEY H 1909 studies on blood and blood parasites II11 the priority of Cryptcryptobiaobia leidy 1846 over try 1958 and heckmann 1971 identified panopanoplasmaplasma laveran and mesnil 1901 USU S depart- salmonid leeches from cutthroat trout in yel- ment of agriculture bulletin of the bureau of ani- lowstonelowstone lake direct transmission studies mal industry 119 16 20 would clarify the role of the leech relative to HECKMANN R A 1971 parasites of cutthroat trout from yellowstone lake wyoming progressive fish cul fish T atwaria infections with atraria turistauristtunst3333 103 106 HOFFMAN G L 1967 parasites of north american fresh- literature CITED water fishes university of california press berkeley and los angeles 486 appp BECKER C D AND M KATZ 1977 flagellate parasites of HUMASON G L 1967 animal tissue techniques freeman fish pages 357 416 in J P kreier editor parasitic and company san francisco 596 appp protozoa volume 1 academic press new york KATZ M 1951 two new hernohemoflagellateshernoflagellatesflagellates genus Cryptcryptobiaobia 1965a transmission of the flagellate Cryptcryptobiaobia from some western washington teleoststeleosts journal of salmositica katz 1951 by a rhynchobdelhdrhynchobdellidrhynchobdellida vector parasitology 37 245 250 journal of parasitology 51 95 99 KHAN R A 1991 further observations on cryptobiaCryptobia dahlidagli 1965b infections of the hemoflagellate Cryptcryptobiaobia mastigophorea Kinetoplastkmetoplastidakinetoplastidaidalda parasitizing marine salmositica katz 1951 in freshwater teleoststeleosts of the fish journal of protozoology 38 326 329 pacific coast transactions of the american fisheries KHAN R A AND E R NOBLENORLE 1972 taxonomy preva- society 94 327 333 lence and specificity of Cryptcryptobiaobia dahlidagli mobius 1965c distribution ecology and biology of the mastigophora bodonidaeBodon idae in lumpfish cyclopteruscyclopterous salmonid leech Pispiscicolapescicolacicola salmositica rhynchobdel- lumpus journal of the fisheriesFishenes research board of lae piscicolidaepiscicohdae journal of the fisheries research canada 29 1291 1294 board of canada 22 1175 1195 LAIRD M 1961 parasites from northern canada II11 1966 host relationships of Cryptcryptobiaobia salmositica haematozoa of fishes canadian journal of zoology plotPiotprotozoaozoa mastigophora in a western washington 3954139 541 548 hatchery stream transactions of the american fish LAVERAN A AND F MESNIL 1901 sur les flaflagellesgelles Aa mem- erieserleselieselles society 95 196 202 brane ondulante des soissonspoispoissonssons genres trypanosoma 199619961 trypanoplasma ATRARIA BLOOD PARASITE OF FISH 149

frubyaruby et trypanoplasma n gen transactions of the fishes of west virginia proceedings of the french academy of science 133 670 675 helminthological society of washington 39 18 22 1912 trypanosomes et trypanosomiasistrypanosomiases and2nd edi- 1972b biological studies on the bemohemofldgellatesbemoflagellatesflagellates tion masson et cie editeursEditeurs parispans 999 appp Cryptcryptobiaobia cataractcataractalcataractaeae and Cryptcryptobiaobia salmositica LEIDY FE 1846 descriptions of a new genus and species of technical papers bureau of sport fisheries and entozoa proceedings of the national academy of wildlife 63 1 25 science philadelphia 3 100 101 stroutFROUT R G 1965 A new hemoflagellate genus crypto LOM J 1979 biology of the trypanosomes and tropanotrypano bia from marine fishes of northern new england plasmsplasma of fish pages 269 337 in W H R lumsden journal of parasitology 51 654 659 and D A evans editors biology of the kinetoplaskinetoplast SWEZYWEZYnezy 0 1919 the occurrence of trypanoplasma as an tida volume 2 academic press new yorkyolk ectoparasite transactions of the american micro- LOM J AND I1 DYKOVA 1992 protozoan parasites of fishes scoscopicalpical society 38 20 24 developments in aquaculture and fisheries science WALESALES J H AND H WOLF 1955 three protozoan dis- volume 26 elsevier publishers amstersdam 315 appp eases of trout in california california fish and MAVOR J W 1915 on the occurrence of a trypanoplasm game 41 183 187 probably trypanoplasma borreli laveran et mesnil WENRICHENRICH D H 19319311 A trypanoplasm on the gills of carp in the blood of the common sucker catostomus com from the schuylkill river journal of parasitology 18 mersoniimersomimersoniinil journal of parasitology 2 1 6 133 MCDANIEL D W 1970 personal communication USU S woo00 P T K 1987 cryptobiaCrypt obia and cryptocryptobiosisbiosis in fishes department of interior fisheries Springspnngvillespringvilleville utah advances in parasitology 26 199 237 NOBLE E R 1968 the flagellate Cryptcryptobiaobia in two woo00 P T K AND WEHNERT S D 1983 direct transmis- species of deep sea fishes from the eastern pacific sionslon of a hemoflagellate cryptobiaCryptobia salmositica journal of parasitology 54 720 724 Kinetoplastkmetoplastidakinetoplastidaidalda BoBodondoninama between rainbow trout PUTZ R E 1970 biological studies on the hemoflagel under laboratory conditions journal of protozoology lates Kinetoplastkinetoplastidaida cryptobiidae Cryptcryptobiaobia catalaccatarac 3033430 334 337 tae sp n and cryptobiaCryptobia salmositica katz 1951 unpublished doctoral dissertation fordham university received I1 september 1995 new york 98 appp accepted 19 january 1996 1972a Cryptcryptobiaobia cataractcataractalcataractaeae spnspdpn n Kinetoplastkinetoplastidaida cryptobiidae a hemoflagellate of some cyprinid gleatgreat basin naturalist 562 C 1996 appp 150 156 geographical REVIEW OF THE historical AND CURRENT STATUS OF OSPREYS PANDION HALIAETUS IN UTAH

clark S monsoni

ABSIRALI small numbers ofofospreysospreys pandion haliaetushaliahaliaeetusetus are known to have nested historically in utah A precise baseline figure is unavailable but the 19th century osprey population in utah probably consisted of at least 15 breeding pahspairspans scattered in 4 geographic regionslegions human persecution is believed to have caused the abandonment of nesting ter- riritories along the wasatch front and in the western uinta mountains by 1900 and 1960 respectively osprey populations in the southern plateaus and green river areas however began increasing in the late 1970s several recent nesting attempts and numerous summer sightings at nontraditional and abandoned historical sites in utah suggest the osprey is also expanding its langeiangerange in utah high productivity for local pairs and long range dispersal from more northerly osprey populations aiealeare discussed as sources for the current surge in utahs osprey population which now consists of approximately 35 breeding pairs

kenkeifkey words osprey pandion haliaetushaliahaliaeetusetus raptor flaming gorge reservoir dispersal

the osprey pandion hahaetushaliahailahaliaetushaliaeetusetus is one of the their range in utah this paper reviews histor- most widely distributed species of raptorsraptores dur- ical osprey breeding territories in utah sub- ing the breeding period the extent of its cos- sequent population declines and current momopolitanpolitan range is exceeded by only 2 other osprey population and range expansion in raptorsraptores the peregrine falcon falco peregri- utah nus cade 1982 del hoyo et al 1994 and barn owl entotytoTatoyto alba marti 1985 eckert and geographic HISTORY OF THE karalus 1987 despite the osprey s broad OSPREYOSPREYIN IN UTAH geographic distribution local populations occur in fragmented and low densities in much of nesting ospreys have been reported from the species range bent 1937 palmer 1988 del 4 geographical areas of utah fig I1 the hoyo et al 1994 this scenario holds true for wasatch front uinta mountains southern pla- most of the intermountain region of the west- teaus and green river table 1 accounts of ern united states henny 1986 johnsgard early ornithologists naturanaturalistslists and egg col- 1990 in utah osprey distribution has been lectors indicate the osprey was a regular sum- paipal ticultitularlyticularlyparticularlyarlyariy limited recently however several mer resident and breeder in utah allenalienailen 1872 personal summer observations of ospreys over found them along the great salt lake marshes 140 km from known breeding pairs prompted west of ogden and henshaw 1874 saw them an investigation into the possible occurrence of at utah lake near provo neither discussed ospreys at other nontraditional utah localities nest observations in these areas but R G bee A survey of individuals from the USU S forest unpublished ornithological notes mentioned service utah division of wildlife resources that ospreys formerly nested along the shores utah state parks and other persons familiar and tributatributariesries of utah lake fig 1 table 1 with osprey ecology was conducted during region A 1994 95 the survey revealed many osprey other records were for the uinta mountains sightings and several nesting attempts between fig 1 table 1 region B J D daynes unpub- I1 june and 15 august at numerous lakes res- lished ornithological notes described the ervoirservoirs and rivers from nearly every legionregionfegion of repeated use of an osprey nest from 1915 to the state since 1990 these sightings represent 1938 on the weber river 20 km east of oak- the first widespread effort by ospreys to expand ley summit county also hayward 1931

epiiilinciitdepartment of geogiapliyofgegraphy brigham yognoungyoungyb g UnNunnerityuniveisityunverityerity brovpiovoprov UT 84602552684602 5526

150 199619961 OSPREY STATUS IN UTAH 151

0 kilometers 80

7 1 D

fig 1 known historical distribution of ospreys nesting in utah A wasatchwas itch front B western uintibintiuinta mountainsmount linstins C southern plateaus D green river

surveyed 8 july 21 august 1930 birds in the and hutchings 1942 specifically cite mirror western cintasuintas where summit duchesne and and trial lakes as having osprey nests twomey wasatch counties converge while not giving 1942 382 visited an occupied nest between actual locations he said a few ospreys nested 16 and 20 july 1932 at the north end of mirror in the mirror and trioltryol sic lakes region bee lake wasatch county this nest was actually 152 GREAT BASIN naturalist volume 56

TABLE 1 nesting populations of ospreys in utah STATUS region historical pairspan s current pairs historical events A wasatch front unknown I1 although numerous records of ospreys nesting in utah exist these birds have apparently B uinta Mounmountinsmountmsmountainstins 5 8 0 undergone 2 separate declines the ist decline C southern plateaus 2 4 8 11 involved ospreys nesting along the wasatch front fig 1 during winter 1848491848 49 depreda- D green river 6 8 20 25 tions upon livestock poultry and grain led to a much publicized contest to kill the basterswasters and destroyers arrington 1958 51 hundreds of mammalian predators and thousands of raptorsraptores in duchesne county and perhaps the same were killed arrington 1958 ospreys would nest hayward et al 1976 referred to when have been on their southern wintering grounds they listed wasatch county as a former nest- during this assault on local predators but the ing area R G bee unpublished ornithological incident suggests that early pioneers in utah notes also cited single pairs at fish scout and treated all carnivorescarnivores and birds of prey with lily lakes in the western uintascintas on 23 may contempt other similar hunts followed and 40 1945 bee recorded that a game warden in years later the utah legislature implemented a duchesne informed him of 2 pairs at moon law awarding bounties for the killing of preda- lake and another pair at an unidentified uinta tors rawley 1985 rewards were available for lake several species of fish eating birds including other early observers of osprey nests in ospreys the destruction that this bounty in- utah include wolfe and cottam hayward et flicted upon fish eating birds in the name of 11 al 1976 who along with bee and hutchings conservatioiconservation was significant and is vividly 1942 saw ospreys nest at fish lake not the described by pritchett et al 1981 uinta mountains lake with the same name the attitudes of early residents toward pred- sevier county beginning in 1928 fig 1 table ators coupled with laws encouraging their 1 region QC on 18 july 1936 R G bee un- destruction may have led to the osprey s published ornithological notes visited the fish extirpation from the wasatch front fig 1 lake nest A local rancher told him ospreys table 1 region A around the turn of the cen- had used that particular nesting site for at tury in 1935 R G bee unpublished ornitho- least 20 years logical notes speculated that human persecu- caused the abandonment of osprey nests behle et al 1958 noted a pair of ospreys tion near utah bee did not record when these in southwestern utah at navajo lake kane lake ospreys disappeared but his manner of re- county on 17 and 18 june 1950 fig 1 table flection on their absence suggests the loss 1 region C particular territory and an Q this occurred well before his 1935 notation nearby panguitch additional site at lake the and2nd period of osprey decline occurred garfield county has been used regularly since in the western uinta mountains fig 1 the s eyre paul behle discovery and 1973 salt uinta mountain nests that daynes bee hay- lake tribune 13 august 1978 walters 1981 ward and twomey reported were observed anonymous 1989 apparently after the 1950s and green before but not ospreys also nested along the river 1960s when osprey colonies along the eastern northeastern utah fig 1 table 1 region D seaboard were decimated by organoorganochlorinechlorine on 23 and 24 july 1959 C M white and C compounds palmer 1988 poole 1989 although bosley white and behle 1960 located 2 nests the impact of synthetic agricultural biocidesbiocides along this river in horseshoe canyon daggett upon ospreys in utah is unknown ospreys in county both nests contained 2 young an other areas of the western united states were additional nest was discovered on the green generally less affected by environmental con- river in uintah county by M horton in june taminantstaminants than eastern populations poole 1974 behle 1981 white 1969 suggested the 1989 white 1994 total population of ospreys nesting along the another possible reason for the decline of green river probably consisted of 6 8 pairs ospreys in the uinta mountains is indiscrimi 199619961 OSPREY STATUS IN UTAH 153 nate shooting resulting from a hostile attitude hatchery wasatch county fig 2aaa in 1995 a by local residents bee unpublished ornitho- and2nd pair built a nest 2 km away at deer creek logical notes twomey 1942 many ospreys reservoir on a 5 m high artificial platform were formerly shot at northern utah fish erected for ospreys fig ab2b deer creek hatcheries during spring migrations white reservoir and the adjacent midway fish hatch- 1969 hayward et al 1976 and some of these ery have been frequented by ospreys during casualties could have been local breedersbleedersbreeders spring migrations for many years behle and hayward et al 1976 66 recorded the perry 1975 additional osprey nesting attempts osprey was formerly a sparse but regular in 1995 include I1 pair at Jorjordanelledanelle reservoir summer resident in utah now greatly reduced wasatch county fig 2qaq2cac and another pair in numbers and considered to be rare and near highland utah county fig ad2d incu- endangered they preface their discussion of bation behavior at the latter site was observed birds in utah by stating they have included all for approximately 2 wk before strong winds records concerning rare species in the state destroyed the nest this site was possibly the however they cited ospreys only in the west- first osprey nest along the wasatch front in ern cintasuintas and records for fish lake in sevier 80 100 yr county they did not include information on the origin of ospreys colonizing new waters the I1 or 2 pairs nesting in the navajo lake in utah is currently unknown but their reluc- panguitch lake area southern utah or the tance to disperse more than 125 km from their pairs at flaming gorge reservoir northeast- natal sites is well documented henny 1986 ern utah in the same year 1976 that hay- poole 1989 reproduction for nests at flam- ward et al 1976 66 described the osprey as ing gorge reservoir is generally high craw- greatly reduced more ospreys 6 pairs nested ley and white 1989 and considering the at flaming gorge reservoir wagner 1977 osprey s pronounced philopatryphilopatry one might salt lake tribune 13 august 1978 than had expect that ospreys at new locations in utah been recorded in any particular year in the derive from this local population while high western uinta mountains productivity has augmented the osprey population on flaming gorge the frequency with current events which ospreys are being witnessed in utah is flaming gorge dam on the green river too great to be the sole result of dispersal from was completed in 1964 and created a narrow that reservoir moreover if flaming gorge were 150 km long reservoir on the utah wyoming the primary source of ospreys pioneering new border the osprey population here remained waters in utah one would expect lakes and relatively stable until the late 1970s and 1980s rivers near that reservoir to be the initial areas when an increase was noted behle 1981 of range expansion this has not been the case crawley and white 1989 found 21 pairs and A more plausible source of ospreys attempt- 1 trio of ospreys at flaming gorge in 1989 of ing to colonize nontraditional and abandoned these 15 pairs succeeded in fledging 37 young historical waters in utah is from spring migrants osprey numbers at fish lake in sevier stopping short of their natal territories farther county increased from 2 pairs in 1989 anony- north osprey populations in idaho and wyo- mous 1989 to 6 in 1993 B lowry US forest ming number in the hundreds of pairs henny service personal communication addition- 1986 poole 1989 and osprey counts made at ally I1 or 2 pairs now nest 3 km away at john- several migration points in the west have bur- son valley reservoir PE wagner personal com- geoned since 1983 hoffman et al 1992 fur- municationmunication other current osprey nest sites thermorethermore migrating subadult ospreys are known at traditional waters in utah include 2 pairs to linger sometimes and even remain at pro- in the panguitch lake navajo lake area of ductive foraging sites south of their traditional southern utah anonymous 1989 breeding grounds swenson 1981 poole 1989 in 1990 a pair of ospreys nested at tropic these lingering individuals may represent reservoir garfield county sorensen 1990 young adults without an established history of this site is 20 km east of region C fig 1 table breeding elsewhere 1 and should be regarded as a geographical if more northerly populations constitute the extension of that area in 1994 a pair of ospreys primary source of ospreys currently pioneer- constructed a nest near the midway fish ing nontraditional waters in utah this long 154 GREAT BASIN naturalist volume 56

A A 7

J 60

ismiePATIN 0060 1 N

fig 2 A osprey nest midway fish hatchery B osprey nesting platform and nest deer creek reservoir C osprey nest Jorjordanelledanelle reservoir D osprey nest and incubating adult near highland utah 199611996 OSPREY STATUS IN UTAH 155 distance dispersal is a recent phenomenon and DAYNES J D 1975 choice experiences in ornithology possibly indicates a saturated breeding popu- unpublished ornithological notes monte L bean museum brigham young univeiuniver provo lation in the northern intermountain west A universitysitssity UT DEL HOYO J A ELLIOTT AND J SARGATAL EDITORSLDITORS 1994 current quantitative evaluation of osprey pop- handbook of the birds of the world volume 2 new ulatulationsions in idaho and wyoming and extensive world vultures to guineaguineafowlfowl lynx editionedicioncdicionediedl eioncionelon banding efforts in these states could help barcelona spain 638 appp determine if this speculation is correct until ECKERTECKERTAA WANDKW AND K E KARALUS 1987 thetheowlsofnoithowlsowis of north amenea crown publishers new york such a project is undertaken the origin of america inc 278 appp ospreys EYRE L AND D PAUL 1973 raptorsraptores of utah utah divi- presently colonizing new waters in sionslon of wildlife resources publication 73773 7 76 appp utah is open to conjecture HAYWARD C L 19319311 A preliminary list of the birds of the subalpine and alpine zones of the uintah sic moun- acknowledgments tains proceedings of the utah academy of science 81518 151 152 HAYWARD C L C COTTAM A M WOODBURY AND H H I1 the following individuals thank who pro- FROST 1976 birds of utah great basin naturalist vided recent sighting information of ospreys memoirs 1 229 appp in utah howard brinkerhoff dwight bunnel HENNY C J 1986 osprey pandion haliahaifahaliaetushahaetushaliaeetusetus technical steve cranny steve fielding burt lowry report EL 86 5 section 431 US aimyalmyarmy corps of kent keller justin neighbor dennis shirley engineers wildlife resources management manual george washington DC 36 appp simmons joann stoddard kenneth HENSHAW H W 1874 annotated list of birds of utah tuttle phil wagner frank welch and leslie annals of the new york lyceum of natural history welch I1 also thank clayton white and jimmie 111lii11 1 14 parrish who reviewed this manuscript hoffmansHOFFMANhoffhopfHoprMANSS WWWRW R deraconandjDERAGON AND J C BEDNARZ 1992 patterns and recent trends in counts of migrant hawks in western northnoi th america unpublished manu literature CITED scriptscnpt hawkwatchhawkwatcbHawk Watch international albuquerque NM JOHNSGARD P A 1990 hawks eagles and falcons of north ALLEN J A 1872 notes of an ornithological reconnais- america smithsonianSmith soman institution press washington sance of portions of kansas colorado wyoming DC 403 appp and utah bulletin of the museum of comparative MARTI C D 1985 longtermlong term ecological studies on barn zoology 3 113 183 owls in the great basin utah birds journal of the ANONYMOUS 1989 utah division of wildlife resources utah ornithological society 1 1 4 nongame bulletin volume 2 no 2 5 appp PALMER R S EDITOR 1988 handbook of north ameri- ARRINGTON L J 1958 great basin kingdom an economic can birds volume 4 diurnal raptorsrapraptorestors paitpaltpart 1 yale history of the latter day saints 1830 1900 harvard university press new haven CT 433 appp university press cambridge MA 534 appp POOLE A F 1989 ospreys a natural and unnatural his- BEE R G unpublished ornithological notes 1932 1959 tory cambridge university press cambridge UK monte L bean museum brigham young university 246 appp provo UT PRITCHETT C L H H FROST AND W W TANNER 1981 BEE R G AND J HUTCHINGSHUTCHINCS 1942 breeding records of terrestrialteioeioel restnalresteal vertebrates in the environs of utah lake utahbndsutah birds great basin naturalist 3 61 90 great basin naturalist memoirs 5 128 168 BEHLEWHBEHLE W H 1981 the birds of northeastern utah utah RAWLEY E V 1985 early records of wildlife in utah museum of natural history university of utah salt utah division of wildlife resources publication 86- lake city 136 appp 2 102 appp BEHLE W H AND M L PERRY 1975 utahbndsutah birds check- SORENSEN E 1990 utah summer bird report june july list seasonal and ecological occurrence charts and 1990 utah birds journal of the utah ornithologi- guides to bird finding paragon press salt lake city cal society 6 33 42 UT 142 appp SWENSON J E 1981 status of the osprey in southeastern BEHLE W H J B BUSHMAN AND C M greenhalgh montana before and after construction ofofieservonsreservoirs 1958 birds of the kanabcanab area and adjacent high westeinwestern birds 12 47 51 plateaus of southern utah university of utah bio- TWOMEY A C 1942 the birds of the uintaumtaomta basin utah logical series 11 1 92 annals of the carnegieCaigaigal negienegle museum 28 341 490 BENT A C 1937 life histories of north american birds WAGNER P 1977 report to the bureau of land manage- offreyofpreyof presprey part 1 smithsonian institution USU S national ment and USU S forest service on raptorsraptores inhabiting museum bulletin 170 409 appp brown s park and the flaming gorge national CADE T J 1982 the falcons of the world london edit- recreation area report issued by the utah state ions limited london UK 192 appp division of wildlifeofwildlife resources vernal UT 77 appp CRAWLEY J AND C M WHITE 1989 interim leportreport pan- WALTERS R E EDITOR 1981 utah bird latilatilonglong distribu- dion haliaetushaliaettishaitahaliahaliaeetusettisetus on flaming gorge 1989 1990 breeding tion utah division of wildlife resources publica- season report issued by the utah state division of tion 811581 15 84 appp wildlife resources and USU S forest service vernal WHITE C M 1969 population trends in utah raptorsraptores UT 99ppappappp pages 359 362 in J J hickey editor peregrinePeregnne falcon 156 GREAT BASIN naturalist volume 56

populations their biology and decline university of studies of the flora and fauna of flaming gorge wisconsin press madison reservoir basin utah and wyoming university of 1994 population trends and current status of utah anthropological papers 48 selected western raptorsraptores studies in avian biology 1516115 igiigl161 172 received 30 march 1995 WHITE C M AND W H BEHLE 1960 birds of flaming accepted 26 february 1996 gorge reservoir basin pages 185 208 min ecological great basin naturalist 562 0 1996 appp 157 161

EFFECTS OF TURBIDITY ON FEEDING RATES OF LAHONTAN CUTTHROAT TROUT oncorhynchus CLARKI HENSHAWI AND LAHONTAN REDSIDE SHINER richardsoniusRICHARD SONIUS EGRECIUSEGREGIUS

gary L Vinyardvinyardlvinyard11 and andy C yuanlyuan1yuang

ABSTRACT the spawning population of lahontan cutthroat trout oncorhynchus clarki henhenshawihenshaweshawi in summit lake nevada has reportedly declined since the early 1970s coincident with the appearance lahontanoflahontanof ledsideredsidereeside shiner RichardrichardsoniusrichardsomusRichardssoniusomus egregius in the lake we investigated the relative predatory abilities of these 2 fish species foraging on live daphnia magna in turbidity conditions commonly observed in summit lake experiments were performed under controlled light and temperature conditions in separate trials we fed trout and shiner lofalof3I1 of 3 size classes ofofdoadD magna 171 7 mm 222 2 mm and 303 0 mm at 6 levels of turbidity ranging from 353 5 to 25 NTU feeding rates for both species varied inversely with turbidity for all prey sizes feeding rates of shiner were greater than trout at all turbidity levels in low turbidity 5 NTU shiner consumed approximately 3 more prey during 2 h feeding trials however at high turbidity levels the difference in feeding rates between species was proportionally higher 10 at high turbidity levels 20 NTU trout predation latesrates were relatively insensitive to prey size however shiner continued to consume more larger prey at the highest turbidity levels these results indicate that lahontan redsidebedside shiner may be superior to lahontan cutthroat trout as zooplankton predators at high turbidity levels and may explain the recent success of shiner in summit lake

key words daphnia lahontan cutthroat trout oncorhynchus clarki henhenshawihenshaweshawi lahontan redsidebedside shiner richardso niusmus egregius planktivory predation size selectivity turbidity

the lahontan cutthroat trout oncorhynchus cowan and blake 1989 valeska 1989 other clarki henhenshawihenshaweshawi is an inland subspecies endemic lacustrine populations are either maintained to the physiographic lahontan basin in northern by artificial stocking or are subject to higher nevada eastern california and southern ore- levels of harvest and disturbance conservation gon these trout were once widespread through- of this population is compelling and it has out the basins of pleistocene lake lahontan been identified as important for recovery of USFWS 1995 currently they occupy 1 the subspecies USFWS 1995 of their former lacustrine range and 11 of cutthroat trout spawning runs at summit their former stream habitat within the native lake have generally declined since the late range USFWS 1995 listed as endangered in 1970s cowan and blake 1989 collection of 1970 the fish was subsequently reclassified as roe during the 1960s and 1970s and excessive threatened in 1975 this facilitated manage- loss of spawning habitat in mahogany creek ment and permitted regulated angling USFWS from livestock overgrazing cowan and blake 1995 1989 vinyard and winzeler 1993 have been summit lake is located in the summit lake blamed however coinciding with the decline paiute indian reservation in northwestern in trout lahontan redsidebedside shiner richardso humboldt county nevada 41n41 N latitude nius egregius also increased in abundance in 119w longitude at an elevation of 1828 m the lake suggesting a competition effect formed by a landslide about 20000 years ago redsidebedside shiner are native to the great summit lake is relatively shallow maximum basin but they do not occur naturally in sum- depth 12 m and has historically been subject mit lake origins of the present shiner popu- to high turbidity levels during summer months lation in the lake are unknown but they have from suspended algae and silt larivers 1962 been used frequently as live bait lahontan it contains the most secure remaining lacus- redsidebedside shiner feed on drift in streams and are trine population of lahontan cutthroat trout zooplanktivorous in lakes vinyard and winzeler and no other salmonsalmonidsids occur in the basin 1993 laboratory observations suggest they

department of biology university of nevada reno nevada 89557001589557 0015

157 158 GREAT BASIN naturalist volume 56

may also prey on larval trout vinyard and by both species lighting was provided by a winzeler 1993 analysis of stomach contents bank of three 56 watt fluorescent tubes con- suggests that lahontan cutthroat trout and trolled by an automatic timer iolloltio111101iol14d10l 14d light lahontan redsidebedside shinershmel probably consume intensity at the water surface averaged 93 aletuejue similar foods both in summit lake and in m 2 S 1 an airstineairairstonestone in the center of each of mahogany creek the primary spawning tribu- four 38381L aquaria provided aeration and kept tary for trout from summit lake vinyard and turbidity in suspension turbidity nephelo- winzeler 1993 both species consume drift in metric turbidity units NTU was measured the stream and mostly amphipodsamphipoda min summit with an HF instruments model DRT 15 tur- lake cowan and blake 1989 in contrast simi- bidimeterbi six turbidity levels 35353.53 5 6 10 20520 larly large lahontan cutthroat trout inm pyramid 22 and 25 NTU were produced using sus- lake are piscivorous USFWS 1995 because pensions of bentonite bentonite concentra- most fish species depend on vision to locate tions eglmgl were significantly correlated with pleyprey hobson 1979 guthrie 1986 it is possi- measured turbidity NTU 25832.5832 583 0162olgg0.1620 162 B ble that high turbidity in summit lake limits r2ra 0990.99ogg0 99 this material is nontoxic and the visibility of prey and impedes the ability of remains in suspension for long periods trout to catch redsidebedside shiner and other large feeding rates were determined for fish ex- prey posed to single sized groups of daphnia magna our experiments compared the relationships at each turbidity level laboratory reared D of feeding rate turbidity and prey size for magna were sorted into 3 size groups using a lahontan cutthroat trout and lahontan red dissecting microscope 171.71 7 mm 222.22 2 mm and side shiner with the primary focus being to 303.03 0 mm top of head to base of tail spine 03 examine the relative performance of both mm before each feeding trial a single fish species under various turbidity levels was placed into each experimental tank and allowed to acclimate for 24 h A group of 200 METHODSMLTHODS AND MATERIALS daphnia were introduced into the tank and the fish allowed to feed for 2 h fish were then lahontan redsidebedside shiner weiewelewere captured from removed and the water and remaining preyprcy mahogany creek humboldt county nevada siphoned through a 363 micron mesh net prey and transported to the university of nevada retained on the net were counted to deter- lahontan cutthroat trout from the current mine consumption rates this procedure was pyramid lake stock were acquired from the repeated for each of the 3 prey size classes lahontan national fish hatchery gardner and 6 turbidity levels with 4 fish from each ville nevada although the historical origins species yielding a total of 144 feeding trials of the existing pyramid lake stock are mixed fish used in the feeding trials ranged from 70 summit lake fish were heavily planted into nunmm to 93 mm SL analysis of variance and lin- pyramid lake for a number of years and they ear regression were used to assess the effects likely constitute the dominant component of of fish species prey size and turbidity level the population USFWS 1995 fish were on predation rates housed in 19igl191 L tanks and acclimated to local watelwater conditions for at least 3 wk prior to RESULTS experiments experiments were conducted in a secluded an analysis of overall predation rates for both section of a greenhouse at the university of fish species consuming all prey sizes figs la nevada the experimental protocol was simi-simisiml ib indicates that feeding rates varied inversely lar to that employed by vinyard and winzeler with turbidity multiple regression F 1894 1993 and li et al 1985 visual isolation of P 00010.0010 ooi001 and between fish species F experimental tanks was ensured by opaque 28428.428884 4 P 00010.0010 ooi001 and that larger prey gener- black polyethylene sheeting 10 mil 25m252.595gs ra ally were consumed at greater rates F 38338.338 3 high which enclosed all sides of the experi- P 00010.0010 ooi001 significant results were observed mental area and controlled external light for both the species NTU and speciesdaph temperatures ranged between 12c12 C and 17cIVC nia size interaction terms indicating that the 2 during the experiments and diel variation fish species differ in their responses to these 2 never exceeded 4cac a range easily tolerated variables lahontan redsidebedside shiner consumed 199619961 TURBIDITY EFFECTS ON FISH FEEDING 159 significantly more prey than lahontan cut- our results demonstrate that turbidity throat trout at the lowest turbidity level 35353.5 reduces predation rates for all prey sizes for NTU approximately 90 of all prey were both lahontan redsidebedside shiner and lahontan consumed by both fish species however even cutthroat trout larger prey were generally small increases in turbidity reduced predation consumed with greater frequency although rates this decrease in predation with turbid- this frequency varies with turbidity and fish ity was strongly linear and there was no indi- species the effect of prey size was most con- cation of a minimum value having been reached sistent for lahontan redsidebedside shiner these fish by 25 NTU at that turbidity level predation consumed more large 30303.0 mm prey at all tur- rates declined by approximately 80 for trout bidity levels than did lahontan cutthroat trout fig la and by 60 to 80 for shiner fig figs la ib in contrast prey size had little ib depending on prey size predation rates effect on the relative numbers of prey of each for trout were significantly affected by prey size consumed by trout at turbidity levels of size and turbidity multiple regression F 20 NTU or above fig la 2672.67967 P 00090.009 for prey size F 35135.1 P redsidebedside shiner also consumed more prey of 00010.001 for turbidity similar results were all 3 sizes combined over all turbidity levels observed for shiner multiple regression F for all prey sizes combined shiner consumed 6546.54 P 00010.001 for prey size F 271527.15 P approximately 3 more prey than lahontan 00010.001 for turbidity cutthroat trout at low turbidity levels and at higher turbidity levels differences in approximately 10 more at high levels figs performance of the 2 fish species became most la ib angradi and griffith 1990 found pre- apparent at turbidity levels of 20 NTU or more dation by rainbow trout 0 mykiss to be more prey of all sizes were consumed at virtually selective for large prey in clear water whereas equal rates by lahontan cutthroat trout fig selectivity was reduced in elevated turbidity la in contrast lahontan redsidebedside shiner showed similar effects on prey selection under reduced increasing predation on 3 mm prey relative to visibility conditions have been observed in the smaller sizes at high turbidity levels fig bluegill sunfish lepomis macrochirusmacrochires under ib and shiner showed the greatest differ- lowlightlow light conditions bluegill sunfish consumed ences in predation rates between prey of dif- fewer zooplankton but proportionally more turbidity levels ferent size at the highest large individuals miner and stein 1993 exhibited the lahontan cutthroat trout oppo- neither trout nor shiner have been shown differences preda- site trend with greater in explicitly to possess adaptations that might en- between prey of different sizes at tion rates hance their effectiveness as foragersforagers in turbid low turbidity levels waters however fish that feed nocturnally discussion such as walleye stizostedion vitreum may perform equally well in either clear or turbid foraging behavior and efficiency are affected waters vandenbyllaardt et al 1991 walleye by local visibility many workers have demon- have higher densities of retinal cells and also strated reduced effectiveness by visual preda- develop scotopic vision earlier in life in tors at elevated turbidity vinyard and obrien comparison to salmonsalmonidsids vandenbyllaardt 1976 li et al 1985 barrett et al 1992 gregory et al 1991 borgstrom et al 1992 hurberburber and northcote 1993 sigler et al 1984 found and rylander 1992 such species specific fac- that chronic high turbidity impedes growth tors may contribute to differences in visual and increases mortality of steelhead 0 mykiss performance and coho salmon 0 kisutch evidence sug- behavioral responses of fish to turbidity may gests that high turbidity or low light intensity also affect their feeding abilities or rates in reduces predator selectivity because relative laboratory experiments golden shiner noheminotemi differences in prey detection distance for dif- gonus crysoleucas showed increased flight ferent sizes of prey are reduced vinyard and responses with increased turbidity chiasson obrien 1976 gregory and northcote 1993 1993 juvenile chinook salmon apparently gregory and northcote 1993 observed log experienced reduced predation from piscivo- linear declines in reactive distance with rous birds and fishes at elevated turbidity lev- increased turbidity in chinook salmon 0 els gregory 1993 during our experiments tshawytscha redsidebedside shiner were observed to search faster 160 GREAT BASIN naturalist volume 56

loo100 mm so Z uj 22mm a 60 Z 171.7 mm 040LU W 4 LU LAHONTAN CUTTHROAT TROUT 20 oncorhynchus clarki henhenshawlhenshaweshawl

01 1 1 1 1 0 6 10 is15 20 25 30 TURBIDITY NTU

loo100 303.0 mm so

LU 660 Z 171.7 mm LU 40 222.2 mm LAHONTAN REDSIDE SHINER 20 Richardsonrichardsonluslus reglusregiusegregluseg

1 1 1 01 1 0 5 10 is15 20 25 30 TURBIDITY NTU

fig I1 mean percent prey consumed in relation to turbidity upper panel a shows results from feeding trials with lahontan cutthroat trout oncorhynchus karetnaretclarki henhenshawihenshaweshawi and lower panel b shows results from lahontan redsideledsidereeside shiner RichardrichardsoniusrichardsomusRichardssoniusomus egregius four fish of each species were exposed to prey of a single size for 2 hb feeding trials daphnia magna prey sizes are as indicated vertical bars indicate I1 standard deviation and more widely at higher turbidity elevated high turbidity in summit lake may decrease turbidity may have provided greater visual reactive distance and search volume unequally isolation and promoted greater mobility by for shiner and trout this may differentially predators as suggested by confer et al 1978 reduce the probability of successful prey cap- and gradall and swenson 1982 increased ture and could produce altered prey selection activity may have compensated for reduced patterns under different turbidity conditions visual effectiveness resulting in larger search although our results are generally similar to volumes for shiner than for trout in a study of those shown for other fishes vinyard and brook trout salvelinus fontinalis and creek obrien 1976 berg and northcote 1985 li chub semotilusSemotilus atromaculatus gradall and et al 1985 we document highly significant swenson 1982 found creek chub to be less differences between potentially competing affected by turbidity than brook trout they fish species because lahontan cutthroat trout suggested such differential effects may explain and lahontan redsidebedside shiner consume the same local disparities in fish density prey in summit lake competition for food 199619961 TURBIDITY EFFECTS ON FISH FEEDING 161 may exist our results suggest that in elevated GREGORY R S 1993 effects of turbidity on the piepredatordatordamor turbidity conditions lahontan redsidebedside shiner avoidance behavior of juvenile chinook salmon may be a better competitor for food than oncorhynchus tshawytscha canadian journal of fisheries and aquatic sciences 50 241 246 lahontan cutthroat trout A factor contribut- GREGORY R S AND T G NORTHCOTE 1993 surface ing to the success of lahontan redsidebedside shiner planktonic and benthic foraging by juvenile chi- in summit lake may be that their predation nook salmon oncorhynchus tshawytscha in turbid rates are higher than those of cutthroat laboratory conditions canadian journal of fisheries trout and at elevated turbidity levels aquatic sciences 50 233 240 GUTHRIE D M 1986 role of vision in fish behavior pages 75 113 in T J pitcher editor the behavior of acknowledgments teleost fishes croomgroom helm london HOBSON E S 1979 interactions between piscivorous we thank larry marchant of the lahontan fishes and their prey pages 231 242 in R H stoud and H clepper editors predator prey systems in national fish hatchery and alice winzeler for fisheries management sport fishing institute wash- providing fish and louis christensen for ington DC assistance in setting up the experimental appa- HUBER R AND M K RYLANDER 1992 quantitative his- ratus we thank R S gregory and 2 anony- totological study of the optic nerve in species of min- nows cyprinidae teleostei inhabiting clear and mous reviewers for helpful suggestions for this turbid water brain behavior and evolution 40 manuscript the university of nevada depart- 250 255 ment of biology undergraduate thesis commit- LARIVERS I1 1962 fishes and fisheries of nevada nevada tee facilitated completion of this project state fish and game commission reno 782 appp li K T J K WETTERER AND N G HAIRSTON JR 1985 fish size visual resolution and prey selectivity literature CITED ecology 66 1729 1735 MINER J G AND R A STEIN 1993 interactive influences ANGRADI T R AND J S GRIFFITH 1990 diel feeding of turbidity and light on larval bluegill leporaslepomislepormsLeporms chronology and diet selection of rainbow trout macrochiresmacrochirus foraging canadian journal of fisheries oncorhynchus mykiss in the henry s fork of the and aquatic sciences 50 781 788 snake river idaho canadian journal of fisheries SIGLER F W AND J W SIGLER 1987 fishes of the great and aquatic sciences 47 199 209 basin a natural history university of nevada press BARRETT J C G D GROSSMAN AND J ROSENFELD 1992 reno 425 appp turbidity induced changes in reactive distance of SIGLER J W T C BJORNN AND F H EVEREST 1984 effects rainbow trout transactions of the american fish- of chronic turbidity on density and growth of steel eries society 121 437 443 heads and coho salmon transactions of the ameri- BERG L AND T G NORTHCOTE 1985 changes in territo- can fisheries society 113 142 150 rial gill flaring and feeding behavior in juvenile USU S FISH AND WILDLIFE SERVICE 1995 lahonLabonlahontantan cut- coho salmon oncorhynchus kisutch following short throat trout oncorhynchus clarki henhenshawihenshaweshawi recov- term pulses of suspended sediment canadian jour- ery plan portland OR nal offisheries and aquatic sciences 42 1410 1417 VALESKA J P 1989 summit lake lacustrine study tech- BORGSTROM R A BRABRAND AND J T SOLHEIM 1992 niques unpublished manuscript summit lake paiute effects of siltation on resource utilization and dynam- tribe winnemucca NV iesles of brown salmo ics allopatric trout trutta in a reser- vandenbyllaardt L F J WARD C R BRAKEVELT AND voir environmental biology of fishes 34 247 255 D B MCINTYRE 1991 relationships between tur- CHIASSON A 1993 effect of suspended sediments the on bidity piscipiscivoryvory and development of the i etinaretina in ninestineninespineninemnespmespine stickleback fungitiuspungitiusPungiFungitius pungipungitiustius and golden juvenile walleyeswalwalleyedleyes transactions of the american shiner notemigonusnotermgonus chrysoleucas in a current of fisheries society 120 382 390 varying velocity environmental biology of fishes VINYARD G L AND W J OBRIEN 1976 effects of light 3728337 283 295 and turbidity on the reactive distance of bluegill CONFER J L G L HOWICK M H CORZETTECOKZETTE S L KAMER lepomisLep orms macrochiresmacrochirus canadian journal of fish- S fitzgibbon AND R landesberg 1978 visual eries and aquatic sciences 33 2845 2849 predation by planktivores oikosbikos 31 27 37 VINYARD G L AND A L WINZELERWINZFLER 1993 results of COWAN W AND R BLAKE 1989 fisheries management investigations at summit lake report to summit services contract cth50913089 annual report lake paiute tribe 62 appp report to summit lake paiute tribe 31 appp GRADALL K S AND W A SWENSON 1982 responses of received 12 june 1995 brook trout and creek chubs to turbidity transac- accepted 19 january 1996 tions of the american fisheries society 111 392 395 great basin naturalist 562 0 1996 appp 162 166

pogonomyrmex OWYHEEI NEST SITE DENSITY AND SIZE ON A MINIMALLY IMPACTED SITE IN CENTRAL OREGON

peter T soul6lsouleasoule1 and paul A knappeknapp2

alwrracrab&iraci little is known about the basic characteristics of the western harvester ant pogonomyrmex owyheei in the absence of anthropogenic disturbances we examined the role of P owyheei as an agent of disturbance in an area of semiarid vegetation in central oregon known as the horse ridge research natural area HRRNA that has been largely freeh ee of livestock grazing and other significant anthropogenic influences for over 23 yr we determined density and size characteristics of nest sites and estimated total area cleared by P owyheei activities on HRRNA from random sampling of twentytwentyfivefive 0040 04 ha plots we found a mean nest densitystandarddensity standard enerroror of 161 6 olg0160.160 16 nests0nests004nests 00404 ha mean area cleared peipelper nest site was 484.84 8 m2ma which results in an estimated barren area of 46080 m2ma on the 240 haba HRRNA comparing our findings to others on P owyheei and P occidentoccidentahsoccidentalisoccidentalistalisallsails we found nest density and mean cleared area to be in the middle range of reported observations under a variety of land use influences the literature suggests that moderate disturbance may increase nest site density but little relationship exists between disturbance history and mean size of nest sites

key words pogonomyrmex owyheei western harvester ants nest density nest size vegetation clearing

western harvester ants are a major compo- of P owyheei nest sites and 2 to estimate the nent of and rangeland ecosystems in the united total area denuded by clearing and foraging states because of the combined effects of seed activities of P owyheei within a largely undis- predation seed dispersal and vegetation re- turbed semiarid ecosystem moval harvester ants are keystone species meaning their effects on vegetation structure STUDY AREA and dynamics exceed expectations given their density and biomass holldobler and wilson the horse ridge research natural area 1990 616 the most visible impact of har- HRRNA is a 240 ha exclosmeexclosure 31 km south- vester ant activities is vegetation clearing east of bend oregon managed by the prine around their nest sites although the size of ville district bureau of land management the cleared area or disc varies pogonomyrmexpogonomynnex BLM the natural area was established in harvester ants have the capacity to cut annual 1967 and a surrounding fence was completed plants surrounding their nest sites at rates of in 1974 the exclosure ranges from 1250 to over 200 million plantshayrplants hayr clark and 1430 m elevation over rolling topography of comanor 1975 while much of the plant bio- columbia basaltsbasales anonymous 1972 direct mass cut is not consumed by the ants it human impacts on the site are minimal as reduces the total volume available for con- there is only occasional use by hunters and sumption by livestock and other glazersgrazers willard naturanaturalistslists and fire suppression is not active and crowell 1965 range managers have halvorson 1991 R halvorson personal com- viewed pogonomyrmex as pests that need to be munimunicationcation 1995 the fence has kept the area controlled giving the ant both economic and free of livestock grazing since 1974 but before ecological importance in and rangelands wight its establishment the area apparently received and nichols 1966 cole 1968 minimal domestic animal grazing pressure because of the paucity of undisturbed areas because of a lack of a permanent water source in the semiarid west little is known about the to attract animals anonymous 1972 and the basic characteristic of FP owyheei nest sites in distance from well traveled public roads the absence of anthropogenic disturbances baldwin 1974 additionally the abundance the primary objectives of this study are 1 to on HRRNA of threadthreadleavedleaved sedge cateycareycarexfilicarex jibjtb determine the density and size characteristics folianoilajohajoba a species that has been shown to decline

11xpartrnentdlpditmtiitotgcoyiof geograpliyandiphy mclmci planning appalachianAppakichunLichun stitestate university boone NC 28607 21epartrnentapaixpai tin nt of giogigcogigeographyiphy atoigeorgiagtoi fiifai i ststateitette uiuvetuniversitysity atlanta GA 30303

162 199611996 P OWYHEEI NEST DENSITY AND SIZE 163 because of overgrazing in the central oregon METHODS sagebrush steppe and the absence of cheat grass bromus tectoriumtectorumtectorum suggest a minimally in roughly the center of HRRNA a 196 ha disturbed site anonymous 1972 personal ob- permanent grid was established by gashwiler servation 1995 1977 for use in an ecological study in 1972 vegetation on HRRNA is classified as the stations on the 12 X 12 grid are marked by re- western juniperjuniperbigbig sagebrushthreadleaved bar stakes and spaced 40240.2 in apart using this sedge community juniperusiunilunifuniperus occidentoccidentalisoccidentalisloccidentalismallsalisails grid we randomly selected 25 stations and artemisia tridentatacarextridentatalcarex filifilifoliafolia franklin established 004 ha circular plots from the rebar and dyrness 1988 less common but present marked center points for a total sample area of species are bluebunchbluebunch wheatgrasswheatgrass agropyron I1 ha we tallied and measured each active P spicatumspicatum idaho fescue festuca idahoensisidahoenslsl owyheei nest site within each plot we placed junjunegrassegrass koeleriaKoeleria cristacristatacricrlcristatdcristanastatatd and horsebrushhorsebrush line transects over the center of each nest site tetradymia glagiaglabrateglabratdglabratabrata anonymous 1972 and measured the cleared disc area in north HRRNA climate is dominated by winter south and east west directions the edge of the precipitation over half the annual 31 cm falls each disc was determined by intersection perennial with the N S E as snow mean temperatures at bend range of any NS or EWW transect lines from 06c060.6og in january to 177c17717.7 in july karl et al 1990 RESULTS soils on our study plots are entirely within the stookmoor wesbutte complex soil series there were 40 active P owyheei nest sites USDANRCSUSDA NRCS in press this soil series is in our ihaI1 ha sample we found nest sites on 23 found on approximately 85 of HRRNA A of the 25 circular plots and the maximum typical soil profile is represented by a surface number of nest sites was 3 per 0040.04 ha mean ash and loamy layer of mixed material approxi- nest densitystandarddensdensityitystandardstandardjandard error was 161.6lgig olg0160.16 mately 15 cm thick and a pale brown sandy nests004nests 004 ha characteristics of the cleared 46 loam subsoil cm deep overlying bedrock discs are shown in table 1 assuming a circu- percentage of organic matter in the topsoil is lar shape the mean area cleared per nest site 1 2 0.505os 2 subsoil and 05 in the USDA- is 484.8 m2ma factoring in the nest density results NRCS in press in an estimated barren area of 192 nha or besides P owyheei there is disturbance 1921.92 of the total land area of the permanent pressure on HRRNA from grazing activities of grid if the influences of P owyheei are consis- herbherbivoresivores and granivoresgranivores such as rocky moun- tent throughout the 240 ha HRRNA then ant tain mule deer odocoileus hemionus hemionus foraging and plant cutting surrounding a total badger taxidea taxus and cottontail rabbits of 9600 nest sites should leave approximately sylvilagusSylvilagus nunatallinutallitalli gashwiler 1972 personal 46080 m2ma of barren land on the 240 ha site observation 1995 BLM records on HRRNA report no outbreaks of intense herbivoreherbberbherbivoryberbivoryivorsivory or discussion episodes of pathogens causing severe plant losses in the last 20 years R halvorson per- the premise of this article is to provide sonal communication 1995 information on P owyheei nest site density

TABLE 1 characteristics of R owyheeiowyheez nest sites on HRRNA standard discs mean median maximum minimum deviation N cm cm cm cm cm

NSN S diameter 2411241.1241 1 2075207202 5 740 60 1444144.4144 4 40

EWE W diameter 2546 2200 670 68 1568 40 164 GREAT BASIN naturalist volume 56

TABLETABLL 2 pogonomyrmex owyheeiowyheel and P occidentoccidentahsoccidentalisoccidentalistalisallsails nest site densities and mean size of nest site and estimated bar- reni en aiarea due to FP owyheeiowyheel and P occidentoccidentalisoccidentalistalis activities reported in the literature estimated pogonomynnexpogonomyrmex nest site nest site barrenban en study site dominant souisoulsourcecc state species densityhadensity ha size in m2ma area disturbance vegetation sharpshelishellsheti paneipanelpanciand barr 1960 idaho owyheelowyheeia 40 08 60 misused atriplex nuttalnuttnuttallnnuttalldallnaituld 1 depleted halogeton glomeratusglomeratus sbarpandsharsheilshellSbar pindpundpand bailballbalibarr 1960 idaho owyheelowyheeia 9 13 37 vigorous stand atriplex nuttnuttallnnuttalkiaikiallnalki sharpardsharpandshellSharshoil pand barrbelli 1960 idaho owyheelowyheeia 12 nrbarbnr11 nr not discussed anplexatiplexatriplex conferticonfertifoliafolia willaidwillard and clowellcrowell 1965 oioregonegon owyheeiowyheel 49 74 225 11 17 not discussed bromus tectoriumtectorumtectorum wight and nichols 1966 wyomingWyorning occidentalisdoccidentalis1occidentalism nr 657 nr lightly grazed atriplex nuttalnuttalliinuttallffnuttallialiiitiittlffiff bogelrogerss and lavigne 1974 colorado occidentoccidentalisoccidentalistalis 23 12 03 ungrazed for buchloe dactyloidesdactyloides 1 30 years bouteloua gracilis rogeisetalRogeisrogers etalet al 1972 colorado occidentoccidentalisoccidentalistalis 28 07 nr lightly grazed buchloe dactyloidesdactyloides bouteloua gracilis rogersrogeisctalet al 1972 colorado occidentoccidentalisoccidentalistalis 31 04 nr moderate grazingglazing buchloe dactyloidesdactyloides bouteloua gracilis

Rogeisrogersrogeisetaletetalal 1972 coloradocoloichloi ado occidentoccidentalisoccidentalistalis 3 06 002 heavy grazing buchloe dactyloidesdactyloides 1 bouteloua gracilis ciarclarkandClarkandand comanor 1975 nevada occidentoccidentalisoccidentalistalis 30304343 242 4 15915 9 nr varied lightly artemisia tntritrldentatedentata grazed recent agropyron desertorumdesertorum burnsbums sneva 1979 oioregonegon owyheeiowyheel 32 93 30 grazed pasture artemisia tntritrldentata 1 no intensity agropyron spicaspicatumtum 1 specified stipa thurthurberianathurbenanathurberbenanebenanalanaiana sneva 1979 olegonoregon owyheeiowyheet 80 09 07 lightly grazed artemisia tntritrldentatatridentatatridentate brush control agropyron spicaspicatumtum 1 10 yr prior to stipa thurberianathurbenanathurthurberthunbenanaiana study killed 95 bromus tectectommtectoriumtectorumteotorum ofplantsofplants sneva 1979 oregon owyheeiowyheel 57 15 08 lightly grazed artemisia tntridentata 1 brushbi ush control agropyron spicaspicatumtum 1 22 yr prior to study stipa thurberianathurbenanathurthurberbenanaianalana 1 killed 95 of plants bromus tectoriumtectorumtectorum coffin and lauenrothLau emoth 1988 colorado occidentoccidentalisoccidentalistalis 25 14 nr moderately grazed bouteloua gracilis coffin and lauenrothlowlomLewemoth 1990 Coloiacoloradodo occidentoccidentalisoccidentalistalis 31 12 nr lightly grazed bouteloua gracilis Nowanowaknowaketalketalketaiet al 1990 idaho owyheeiowyheel nr 35 nr no grazing or artemisia tntritrldentata 1 fire in 30 yr oryzopsis hymenoideshyme noides nowaknowakctalet al 1990 idaho owyheeiowyheet nr 53 nr burned 5 yr artemisia tndentata 1 prior to sample oryzopsis hymenoideshyme noides then ungrazed ilidentifiedaslckick utidcd is occidtntalisolcidcntlilv buthut in the known ungerangei efowofofowoujifhtuyheei noibnotabnotnot reportreportedcd alicallcailall referencesi feifelacitci eulesenlesLULLS to gigrazingtang eielefelreferi to gigrazingizing of ciatleciagleittlelttieuttie 01or othelother livestock P iwifhcctotvyheei wasw is considncdlonsicllicd to belielyetye partp ittlit of P occidentocctdtntuhsoccidentalisoccidentalistallsailsaliS until 1950 and cleared disc size in an undisturbed area for example rogers and lavigne 1974 995 much of the information on areas cleared by found an increase in nest site density under pogonomynnexpogonomyrmex harvester ants relates to study light and moderate grazing but sharply re- sites with varying degrees of disturbance his- duced densities under heavy grazing findings tory however few studies examine the role of of sharp and barr 1960 and sneva 1979 also P owyheei and P occidentoccidentalisoccidentalistalis as agents of suggest increases in nest site density are asso- plant removal in undisturbed environments ciated with disturbance table 2 across the in our study we briefly compare results of range of P owyheei and P occidentoccidentalisoccidentalistalis nest plant removal in undisturbed areas with those site densities are likely controlled by a suite of results presented elsewhere factors soils vegetation composition climate our nest site density of 40ha is in the disturbance history acting synergistically approximate middle range of reported obser- increases in nest site density in grazed areas vations under a variety of land use influences probably result from alterations of the dynam- table 2 disturbance may serve to increase the ics of competition between plant species that nest site densities at any given site up to a point in turn modify seed density distributions 199619961 P OWYHEEI NEST DENSITY AND SIZE 165

holldobler and wilson 1990 on their study harvesterharvesteister ants pogonomynnexpogonomyrmex occidentoccidentalisoccidentalistalis environ- entomology 4 site in southern arizona for example david- mental 52 56 COFFIN D P AND W K LAUENROTH 1988 the effects of al son et 1984 found that harvester ant popu- disturbance size and frequency on a shortshortgrassgrass plant lations began to decrease approximately 2 yr community ecology 69 1609 1617 after rodent populations were intentionally 1990 vegetation associated with nest sites of reduced davidson et al 1984 1780 con- western harvester ants pogonomyrmex occidentoccidentalisoccidentalistalis cresson in a semiarid grassland american midland cluded that rodent removal led to a differen- naturalist 123 226 235 tial increase in large seeded annuals because COLE A C JR 1968 pogonomyrmex harvesterhalvesterhaivesterlvester ants a of the cessation of granivoregranivorygranivory and this in turn study of the genus in north america the university precipitated the competitive displacement of of tennessee press knoxville 222 appp DAVIDSON D W R S INOUYE AND J H BROWN 1984 small seeded that were the ant s pri- species granivorygranivoreGranivory in a desert ecosystem experimental evievlevi- mary food source dence for indirect facilitation of ants by rodents although other studies have used larger ecology 65 1780 1786 sample sizes to determine nest density eg DEMERS M N 1993 roadside ditches as corridors for range of the western harvester ant pogo coffin and lauenroth 198819881 used a ha expansion 25 nomyrmex occidentoccidentalisoccidentalistalis cresson landscape ecology sample we believe our nest site density is a 8 93 102 reasonable estimate for HRRNA because 1 FRANKLIN J F AND C T DYRNESS 1988 natural vegeta- the study is in regard to soils tion of oregon and washington oregon state uni- site consistent press and vegetation and has only minor topo- versity corvallis 452 appp GASHWILER J S 1972 list of birds mammals and plants graphic variability 2 our standard error per found on horse ridge letter 1227 to paul W sample for nest density is small suggesting lit- arrasmisthArrasmisth PrPrinevillemeville OR BLM tle variability within our study area and 3 1977 bird populations in four vegetational types research from studies on other pogonomyrmex inm central oregon special scientific report wildlife no 205 united states department of agriculture species has shown that soil texture can affect fish and wildlife service washington DC nesting location ege g johnson 1992 demers HALVORSON R 1991 natural ignition fire in horse ridge 1993 and that a uniform dispersion of ant ACECRNA concerns and challenges correspon- colonies develops regardless of spatial scale dence 916 to ADM resource services PrPrinevillemeville OR BLM examined and 1995 wiernasz cole there holldobler B AND E 0 WILSON 1990 the ants the appears to be little relationship between dis- belknap press cambridge MA 732 appp turturbancebance history and mean size of nest sites JOHNSON R A 1992 soil texture as an influence on the table 2 sneva 1979 has speculated that distribution of the desert seed harvester ants pogon messor dei oecologia variability omynnexomyrmex rugosus and perganpergandeipergander while there may be great in nest 8911889 118 124 site density and disc area the potential avail- KARL T R C N WILLIAMS JR FE T QUINLAN AND T A able forage per nest site generally remains BODEN 1990 united states historical climatology consistent suggesting that vegetation cover network HCN serial temperature and precipitation data carbon dioxide information analysis and species composition can affect disc size center oak ridge TN 379 appp soil characteristics also impact disc size with NOWAK R S C L NOWAK T DEROCHER N COLE AND a tendency for colonies to expand horizontally M A JONES 1990 prevalence of orysopisOry sopis hymehymenoidesnoides in shallow soils sneva 1979 therefore disc near harvester ant mounds indirect facilitation by ants oikosbikosoikos5858 190 198 size may be largely linked to the amount of roROGERSERS L E AND R J LAVIGNE 1974 environmental vegetation cover plant species composition effects of western harvester ants in the shortshortgrassgrass and soil depth and less influenced by distur- plains ecosystem environmental entomology 3 bance than is nest density 994 997 ROGERS L E R J LAVIGNE AND J L MILLER 1972 bioenergeticsBioenergetics of the western harvester ant in the literature CITED shortshortgrassgrass plains ecosystem environmental entomology 1 763 768 ANONYMOUS 1972 horse ridge research natural area SHARP L E AND W FE BARR 1960 preliminary investiga- federal research natural areas in oregon and tions of harvester ants on southern idaho range- washington pacific northwest forest and range lands journal of range management 13 131 134 experiment station portland OR SNEVA F A 1979 the western harvester ants their den- BALDWIN E M 1974 evaluation of horse ridge natural sity and hill size in relation to herbaceous productiv- area Dedeschutesschutes county region for eligibility for ity and big sagebrush cover journal of range man- registered natural landmark designation supple- agement 32 46 47 mental report university of oregon eugene USDA NATURAL RESOURCES conservation SERVICE in CLARK W H AND P L COMANOR 1975 removal of press upper Dedeschutesschutes river area oregon soil sur- annual plants from the desert ecosystem by western vey USDA NRCS washington DC 166 GREAT BASIN naturalist volume 56 wiernaszWILRNASZ D C AND B J gollgoleCOLECOLL 1995 spatial distribu- WILLARD J R AND H H CROWELL 1965 biological tion of pogonomynnexpogonomyrmex occidentoccidentalisoccidentahsoccidentalistalis lecruitmentrecruitment mor- activities of the harvester ant pogonomynnexpogonomyrmex owyheei tality and dispersionoveroverdispersion journal of animal ecology in central oregon journal of economic entomology 6451964 sig519 527 5848458 484 489 wicilljwigirr J RRANDJAND J T NICHOLS 1966 effects of harvester ants on production of a saltbush community journal received 13 november 1995 of range management 19 69 71 accepted 28 february 1996 great basin naturalist 562 C 1996 appp 167 171 FIELD measurements OF alkalinity FROM LAKES IN THE UINTA MOUNTAINS UTAH 1956 1991

dennis D austini

ABSTRACT data collected from alpine lakes in the uinta mountains during fishery surveys by the utah division of wildlife resources indicate alkalinity has decreased in some drainages since the mid 1950s implications for continued monitoring as well as environmental and recreational values are discussed

key words alkalinity acid precipitation alpine lakes water quality utah

alpine lakes in the uinta mountains have rationstittitrationsnitrations were made at increments of 686.8 the lowest total alkalinity of all surface waters eglmgl per drop in all drainages except during in utah EPA 1982 the low alkalinity is due period 2 in rock creek duchesne and provo to the precambrian rock geologic origin com- river drainages and during period 3 in rock posed primarily of metamorphic quartzite creek burnt creek and sheep carter creek phyllite and diamictite because of low alkalin- drainages when the increments were 17117.1 ity these lakes are sensitive to acid precipita- eglmgl per drop the effects of 3 weaknesses tion which may affect longtermlong term water quality in the available data the lack of data sets fish and other aquatic organisms the utah from all drainages during all 3 periods the 3 division of wildlife resources has measured levels of sensitivity in the alkalinity measure- alkalinity in many of these lakes since 1956 ments and the differences in sample sizes the purpose of this paper is to document the are unknown and suggest interpretive caution changes in alkalinity between 1956 and 1991 of the results the significance level was set at in the uinta mountains by drainage FP 005oos0.05 for the 3 comparisons of statistical testing METHODS to test for changes over all drainages mean alkalinity among drainages was compared water from lakes in the uinta mountains between periods using ANOVA for unequal utah was sampled and measured for alkalinity sample sizes sokal and rohlf 1981 from 1956 to 1991 by the utah division of to test for changes in alkalinity within drain- wildlife resources in conjunction with the ages data were compared between periods T fisheries surveys data were collected during tests of the mean were used when data from 2 summers on selected lakes within 16 of the 18 periods were available and ANOVAs for un- major drainages fig 1 and during 3 desig- equal sample sizes were used when 3 periods nated sampling periods mid 1950s early 1960s of data were available period 1 1970s early 1980s period 2 and to test for changes in alkalinity within mid 1980s early 1990s period 3 all alkalin- drainages for the same sampled lakes I1 com- ity data were collected in the field using col- pared data between periods T tests for paired orimetricorimetric methods and converted to eglmgl in comparisons were used when 2 periods of data period 1 tests were made using methylpurplemethylpurple were available and ANOVAs for equal sample indicator and titrating with 0020.02 N sulfuric sizes when 3 periods of data were available acid alkalinity titrations were made at stepwise increments of 505.0so eglmgl per drop in periods RESULTS 2 and 3 tests were made with hach hach company PO box 389 loveland CO 80539 mean alkalinity among drainages signifi- water ecology kits model AL 36b alkalinity cantly decreased FP 0050.05 between all 3

epaitmentdepartment of range science utah state university logan UT 84322523084322 5230 present address 43 south 700 east hyrum UT 84319

167 168 GREAT BASIN naturalist volume 56

saltIAIT LAKE CITY

US 80 NORTH US 189 SOUTH

U S 40 U S 30 HEBER

KAMAS

4 UTAH PROVO it i WYOMING EVANSTON WEBER riverariverk

DUCHESNE 1 U 1 50

TABIONA ROCK CREEK beagBEAR giverRIVER

eakeLAKE FORK BLACKS FOR

yellowstone giversRIVERS ITHS FORK

SWIFT CREEK HENRYS FORK MOUNTAIN VIEW DUCHES E larewelarmwe i DRY gultchGUILCH P eaverEAVEP CREEK 1 I TA RIV E Rs LONETREE

BURNT FORK whiterocks RIVER rork

whiterocks US HIGHWAY 30 EAST ROOSEVEL SHEEP CREEK

C ARTER ashliiashlie CREEK CREEK LAPOINT MANILA RED CLOUD LOOP ROAD

VERNAL FLAMING GORGE RESERVOIR U 44

DUTCH lohnJOHN US HIGHWAY 40 EAST

NORTH

fig 1 general location and major drainages in the uinta mountains utah 199619961 UINTA LAKES alkalinity 169 periods table 1 from 33 eglmgl in period 1 to tation pough and wilson 1977 EPA 1979 23 eglmgl in period 2 to 17 eglmgl in period 3 haines 1981b kretser et al 1983 and result standard deviations for all means listed in in a general reduction in diversitybiodiversitybio fryer table 1 are available from the author 1980 alkalinity within individual drainages sig- alkalinity and ph are directly related in nificantlynificantly decreased in the duchesne river maintaining aquatic ecosystems and as alka- and provo river drainages between all 3 peri- linity decreases lakes become increasingly sus- ods alkalinity decreased between periods 1 or cepceptibletible to acidification haines 1981a acidi- 2 and period 3 in the rock creek weber river ficationfi rates were reported by dillian et al and whiterocks river drainages no change in 1987 for 2 canadian lakes as 2 ueqlyr be- alkalinity was found between periods 1 2 and tween 1979 and 1985 with a 3 fold decrease in 3 in the bear river drainage similarly no alkalinity accompanied by a 020.2 ph decrease change in alkalinity between periods 2 and 3 decreases in alkalinity have been reported was found from beaver creek blacks fork in colorado in the colorado rockies 64 lakes smiths fork and henrys fork drainages due were compared between 1938 1960 and 1979 to lack of data no additional comparisons could with a mean decrease between periods 1938 be made 1960 vs 1979 of 17 alkalinity lewis 1982 changes in alkalinity within drainages where in the mt zirkel wilderness area turk and data from the same lakes were available be- campbell 1987 reported an approximate loss tween 2 periods were variable alkalinity did of buffering capacity of 10 in most lakes not change in the rock creek drainage between they surveyed periods I1 and 3 or in the weber river drain- data from this study indicate a 50 de- age between periods I1 and 2 alkalinity also crease in alkalinity since the 1950s with the showed no change in the rock creek bear rate of decrease about 050.5 mglyr in the uinta river blacks fork smiths fork or whiterocks mountains at this rate of decrease studies river drainages between periods 2 and 3 extended for only a few years would likely however alkalinity decreased in the duchesne show no change in alkalinity river and provo river drainages between contrary to our results 2 previous studies periods I1 and 2 2 and 3 and 1 and 3 alkalin- conducted in utah indicated no effects of acidi- ity also decreased in the weber river and ficationfication in a snowbeltsnowsnowmeltmelt study of the wasatch bear river drainages between periods I1 and 3 mountains messer et al 1982 reported a but increased in the henrys fork and beaver mean snowbeltsnowsnowmeltmelt ph of 6176.17git and concluded that creek drainages between periods 2 and 3 no enough buffering capacity was retained in the additional comparisons could be made snowsnowpackpack to neutralize acid equivalents from alkalinity in the duchesne river and provo air pollution in a report from the utah techni- river drainages where the same lakes were cal advisory committee on acid deposition sampled during all 3 periods was significantly ellis 1986 concluded that although lakes and P 0050.05oos different between all 3 periods for streams in the uinta mountains are very sen- both drainages mean alkalinity values using sitive to acidification no evidence was found the combined data from these 2 drainages that demonstrated acidification had occurred decreased from 37 eglmgl in period I1 to 22 in the lack of acidification was based primarily period 2 to 6 in period 3 no comparisons on data collected in the mirror lake water- over the 3 periods could be made from the shed during 1983 1986 both studies sug- other drainages gested windblown particulates from the great salt lake desert were sufficient to buffer acid discussion deposition decreased alkalinity from alpine lakes sam- negative effects of acid precipitation on pled by the utah division of wildlife resources aquatic ecosystems have been well documented in the uinta mountains over 35 years indi- particularly in europe and eastern north cated a slow decline in alkalinity particularly america haines 1981a198 la acid precipitation in the provo river and duchesne river drain- can have negative impacts on water chemistry ages unaltered this decline may eventually and quality algae bacteria invertebrates result in deterioration of the aquatic ecosys- amphibians fish waterfowl and aquatic vege tem and subsequently recreational values 170 GREAT BASIN naturalist volume 56

TABLE 1 mean total alkalinity eglmgl by drainage from alpine lakes in the uinta mountains utah

combined data from all sampled lakes phase n 18 period 1 period 2 period 3 drainage year n eglmgl year n eglmgl year n eglmgl rock creek 1956 9 3111311 1 1973 3 34 1983 54 alb2lb21b duchesneduehesne river 1956 30 351 1979 7 2411 1985 34 8cac provo river 1956 23 361 1979 20 23b 1986 54 8cac weber river 1956 27 353511 1983 3 22ab22all 1987 16 121 beaibealbear river 1956 5 30 1982 26 17 1989 30 21 blacks fork ndend3 1982 22 26 1989 21 25 smiths fork ND 1983 24 16 1990 20 15 henrys forkfolkfoik ND 1984 22 15 1990 21 24 beaver creek ND 1984 23 20 1991 17 29 burntbiubinblu nt fork ND ND 1984 11 17 sheepcartersbeepcarterSheep Carter ND ND 1984 32 19 creeks ashley cicreekeek ND ND 1988 21 13 whiterocks river ND 1976 2 343411 1985 43 14b uinta river ND ND ND dry gulch ND ND 1987 12 14 yellowstone river ND ND 1986 24 14 lake fork river ND ND ND swift creek ND ND 1987 17 14 a b TotalMean 94 33331 152 23 427 177cac lnunibcrsunibus witwith different letters across rows weiewelewere significantly different P 005 2ndcNDC no datacl it i frornbrornhiomblom conanoncommonconnnon 1 ikeslakes availableavail thle and3ndND no alkalinityilkllimtyddtidata collected additional sampling is essential to monitor HAINES T A 1981a acidic precipitation and its conse- and document alkalinity and potential acidifi- quences for aquatic ecosystems a review transactions of the american fisheries society 110 cation of uinta mountain lakes 669 707

1 1981b waterfowl and their habitat threatened by acknowledgments acid rain pages 177 190 in ath4th international waterfowl symposium KRETSER A COLQUHOUN AND M H PFEIFFER this report was funded in part by the utah W J R 1983 acid rainram and the adirondack sportsportfisheryfishery state division of wildlife resources pittman conservationist 37 22 29 robertson federal aid project W 105 R spe- LEWIS W M JR 1982 changes in ph and buffering cial thanks is given to jerry D weichman for capacity of lakes in the colorado rockies limnology his careful reviews of this manuscript and oceanography 27 167 172 MESSER J J L SLEZAK AND C I1 LIFE 1982 potential for acid snowbeltsnowsnowmeltmelt in the wasatch mountains water literature CITED quality series UWRLQ 8206 water research laboratory utah state university logan DILLIAN D J R A REID AND E DE GROSBOISGHOSBOIS 1987 POUCHPOUGH F H AND R E WILSON 1977 acid precipitation the laterate of acidification of aquatic ecosystems in and reproduction success of ambystoma salaman ontario canada nature 329 45 49 ders water air and soil pollution 7 307 316 ELLIS M T 1986 acid deposition inm utah utah depart SOKAL R R AND FE J ROHLEROHLF 1981 biometry 2ndand edi- ment of health salt lake city tion W H freeman and company new york EPA 1979 research summary acid rain USU S environ TURK J TTANDDAND D H CAMPBELL 1987 estimates of acid- mental protection agency publication 6008790286008goos 79 028 ification of lakes in the mt zirkel wilderness area washington DC colorado water resources research 23 175717611757 1761 EPA 1982 total alkalinity of surface waters US environmental protection agency publication 600dgood 82 received 22 march 1995 333 corvallis OR accepted 30 december 1995 FRYERFRYLR G 1980 acidity and species diversity in fresh water crustacean faunalfaunas freshwater biology 10 41 45 199619961 ulntauentaUINTA LAKES alkalinity 171

TABLE 1 continued

data from trtheie same 1lakes all data from the same cikeslakes samsampledpledpied dunnduonduring 2 periods sampled duringduardunrig 3 periodspenidsenods

1 2 1 3 2 3 1 2 3 n eglmgl eglmgl n eglmgl eglmgl n eglmgl eglmgl n eglmgl eglmgl eglmgl ndc2 8 30 24 3 34 17 NDC 6 35 23b 21 36s36a ab5b 6 231 661 6 333 23b 66cac 14 373 2111 14 38383 ab7b 15 223 ab5b 10 414pap 211 6 2 33 7 8 333 ioblobloyjob103 NDC NDC NDC 5 303011 1313b 17 20 20 NDC NDC NDC 18 24 26 NDC NDC NDC 13 11 12 NDC NDC NDC 14 123 ith NDC NDC NDC 17 222211 2911 NDC NDC NDC NDC NDC NDC NDC NDC NDC

NDC NDC NDC NDC NDC NDC 2 34 14 NDC NDC NDC NDC NDC NDC NDC NDC NDC NDC NDC NDC NDC NDC NDC NDC NDC NDC NDC NDC NDC 3711 221 g0ga 22 353511 1711711 56 33s33a 1211211 88 23 17 16 371 22 6 great basin naturalist 562 C 1996 appp 172 176

DENSITY BIOMASS AND DIVERSITY OF grasshoppers orthoptera ACRIDIDAE IN A california NATIVE GRASSLAND

eric E Porterporterl2porterlyporter1212 richard A redall and H elizabeth brakertbraker3

ABSABSTRACITRAur A native california perennial grassland was sampled for grasshopper populations the grassland is managed foiformol the preservationpresel vation of the native perennial bunchbunchgrassgrass cassellanassellaNassella pulchrapulchera hitch grasshopper density biomass diseidiversitydivei sitssity and richness were measuimeaskimeasureded from july 1993 to october 1994 average density of all grasshoppers was 2302 30 hoppersm2 066ogg0660.660 66 s for 1994 june through august overall forage consumed for 1994 was 140 kghaagha suggesting that grasshopper populations exist at economically damaging levels grasshoppers do not appear in the grasslands until late springspong after annual grasses have set seed biomass of grasshoppers peaks in july when adults are predominant both grasshopper density and biomass were higher in 1993 than in 1994 and a total of 5 species were found throughout the study melanoplus sanguinsanguinipessanguimpesipeslpes fabricus dominated the acndidacridicacridid communities and accounted for more than 95 of the individuals

ketikeifkey words cassellanassellaNassella pulchpulchrapulcherara melanoplus sanguinsanguinipessanguimpesipeslpes california native grassland density diversity grasshopper herbivoreherbherbivoryivory acrididae

Californicaliforniadss native perennial bunchbunchgrassbuncligrassgrass grasses in Californicaliforniadss native grasslands grass- communities have been reduced to less than hopper herbivoreherbherbivoryivory is presumed to be greatest 1 of their original range heady 1977 with in summer months when annual grasses already much of this loss attributable directly to the have set their seed and prior to germination in development of agricultural and urban areas the fall joern 1989 therefore only perennial huenneke 1989 additionally most undevel- grasses and summer forbs are susceptible to oped patches of native grasslands have con- damage by grasshopper herbivoreherbherbivoryivory furthermore verted to grasslands dominated by annual many grasshopper species exhibit preference grasses native to the mediterranean region for perennial grasses in the field capinera and jackson 1985 factors leading to the success sechrist 1982 joern 1989 suggests that this of these mediterranean species are not com- phenology based selective damage could re- heavy pletely understood however grazing duce the competitive ability of native perennial implicated as factor pressure has been a major grasses against naturalized annuals favors these more ruderal annual that species there are few data available to support or burcham 1957 in their pristine state before refute joern s 1989 hypothesis beyond basic the arrival of european settlers california s surveys of grasshoppers throughout the state grasslands had light grazing pressure wagner strohecker et al 1968 no population or 1989 removal of major anthropogenic distur- commacommunitycommunityanity levellevei studies are available for cali- bances such as grazing and fire does not lead to the recovery of native perennial grasslands fornia s grasshoppers in california native peren- white 1967 keeley 1981 most investigators nial grasslands eg population density species now agree that the annual grass species should abundance and biomass estimations the ob- be considered naturalized and a return to the jective of this study was to describe the grass- pristine disturbance pattern will not lead to hopper community found in a representative reestablishment of native grasslands heady remnant stand of native perennial grassland 1977 over a period of 2 seasons these data will joern 1989 suggests that through differen- provide information necessary to understand tial herbivoreherbherbivoryivory upon the perennial grasses rela- the role of grasshoppers in cailCalicaliforniaforniass grass- tive to annuals grasshoppers may have con- lands and should lead to more informed deci- tritributed to the establishment of exotic annual sions for grassland conservation managers

department of entomology university of california at riverside riverside CA 92521 please address all correspondence to this author department of biology occidental college los angeles CA 90041

172 199619961 grasshoppers OF A california GRASSLAND 173

METHODS AND MATERIALS and august 1994 grasshoppers were identified to and weighed the study area species to nearest mg identifications provided the proportion of the study was conducted in the santa rosa adults p in each sample given total density plateau ecological reserve SRPER located d adult density dadg was calculated with the 10 km west of murrieta california the site is following formula actively managed by the nature conservancy for the restoration and preservation of its rare da d pa habitats the reserve covers 2800 ha and contains about 1200 ha of native perennial grass- species diversity was measured using the lands amongst oak woodlands coastal sage shannon weiner index pielou 1977 feeding scrub and chaparral six sites were established category designations follow capinera and within the perennial grasslands these sites sechrist 1982 and otte 1981 graminivorous were burned in june of 1992 as a management forbivorous or mixed identification of nymphal practice to retard annual grass establishment stages is difficult and damaging feeding does grazing has been excluded from all sites since not occur until the ath4th instar onsager 1984 at least 1990 R wells SRPER reserve man- therefore where possible adult grasshopper ager personal communication data are analyzed separately from total grass- purple needle grass nassellacassellaNas sella pulchrapulchera A hopper data hitch is the most abundant native grass in the reserve common exotic annual grasses RESULTS include slender wild oats avena barbata link and red brome bromus laevipeslaevipes rubens labillcabillLabill species of grasshoppers collected are listed common forbs include annual barsagebursage ambro- with subfamily and known feeding prefer- sia acanthicarpa hook doveweed eremo ences table 1 average grasshopper density carpus setisetigerussetigerousgerus H and filaretfilaree erodium for the 1994 season june august was 2302.30 cicutarium lher lathrop and thorne 1985 grasshoppersm2 table 1 density measurement began too late in the season to estimate grasshopper sampling an average for 1993 A total 198 GHD were six transects were arbitrarily placed through- determined for 1994 leading to an estimated out the perennial grassland areas representing 140 aghakgha of forage consumed density esti- maximum topographic and vegetational het- mates of zero grasshoppersm2 were found from erogeneity each transect measured 200 m november 1993 through may 1994 density long by 20 m wide grasshopper density was peaked in june for 1994 at 292.9 grasshoppers determined with twenty 025 m2ma hoops m2ma this peak in density was dominated by onsager and henry 1977 thompson 1987 immature stages fig IA density measures hoops were placed along each transect at 10 were higher in 1993 than in 1994 for all paired m intervals density was determined monthly sample dates in july august and october f beginning in july 1993 grasshopper days 4694.69 df 1 20 P 00410.041 biomass peaked GHD and forage consumption estimates in july when most grasshoppers were in the were determined following onsager 1984 adult stage fig ab1b biomass days for 1994 GHD is a measure of total grasshoppers found totaled 13213.2 g dm2dmd table 1 peak biomass per m2ma for a given year forage consumption is august was higher in 1993 than in 1994 t an estimate of the yearly forage consumption 2432.43 P 00360.036 of grasshoppers based on estimated daily con- the shannon weiner diversity index includ- sumption 065ogs0650.65 times body weight and GHD ing adults and nymphs combined averaged biomass days were calculated using the same 01400.140 over the 5 collection dates fig 2aaa the formula for GHD replacing grasshoppersm2 peak in adult diversity in june 1994 repre- with gm2gme when possible density 05os050.5 grass sents only 6 individuals of 2 species combined hoppersm2 100 individual grasshoppers adult and nymph species richness averaged were collected from each site and frozen 343.4 for the sampled dates highest species immediately these collections were taken richness was found in august in total 5 grass- directly following density counts and were hopper species were found in these sites for made in august and october 1993 june july the collection dates 174 GREAT BASIN naturalist volume 56

rableTABLEFABLE 1 species of grasshoppers collected in 1993 94 on the SRPER with known feeding types capineraCapmera and sechrist 1982 otte 1981 and appearance composition GHD biomass days average density and weight as calculated foiroiforboroor the sampling period GHD average average sub feeding grasshopper biomass density adult species familylfamilyamilya typetipelltypell appearance icomposition daysm2 dayse macm2c veightweight mg camnulacarnnulaCarncampulanula pelpellucidalucida scudder 0 G may aug 02 036 0076 001 72 melanoplus andus scudder M NA jun aug 03 050 0031 001 90 Medanmelanoplusoplus sanguinsanguinipessangwmpesipeslpes fabifabricusleusicus M M may oct 975 19347 12754 224 132 MermermeriaMerrmermerianmerrnenamerlamerianena bivittatebivittata servilleSeiville G G may oct 16 318 0375 003 81 Psolessa texana scudder G G jun oct 05 089 0121 001 61 tohalTOTALTCHAL 198 13240 230 131

010 ocdipoclinocdipodinae IL M melmeimehimplinaeinoplin e G gompholeringomphocerinae ie 11cC grassgi iss M mixed NA iiottvailablenot ivul ibie lidiidcg111110 ilmtomasswomasslimalm iss irysdays anindunddaverageivelIVLI cige densityd sity toimoimot 1994 seasonse son only

discussion biomass only at grasshopper densities equivalent to 845 GHD or greater we found only grasshopper populations inm the SRPER 198 GHD in 1994 suggesting that densities appear in late may or early june grasshopper during these years may not greatly affect grass- biomass peaks and the most severe herbivoreherbherbivoryivory land plant community dynamics according to occurs in july and august by this time quinn 1993 annual grasses have already died and their the climate of Californicaliforniadss grasslands makes seeds are buried and protected from above- comparison with other north american stud- ground herbivoreherbherbivoryivory savelle and heady 1970 ies difficult most comparable studies examine grasshopper densities decrease dramatically talltallgrassgrass mixed grass and shortshortgrassgrass prairies after august and few are present by october east of the sierra nevada the dry hot summers annual grasses are triggered to germinate that characterize Californicaliforniadss mediterranean after the first fall rains heady 1958 by the climate severely limit growth risser et al time these rains arrive grasshopper densities 1981 regrowth following summer herbivoreherbherbivoryivory 0 the annual are near therefore both mature is similarly limited clearly only perennial seedlings grass- grasses and their escape serious grasses are susceptible to herbivoreherbherbivoryivory at this herbivoiherbivoreherbivoryivoryy hopper herbivors time and the actual effect of such herbivoreherbherbivoryivory in one melanoplus sanguinsanguinipessanguimpeslpesipes species california grasslands is undocumented there- accounted for over 95 of grasshoppers found fore specific studies on the effect of grass- SRPER table 1 this commonly in species hopper herbherbivoryivory on native california grass- damages crops and rangelands throughout herbivore lands must be conducted to predict the level north america hewitt 1977 intense out- and of infestation if any that may favor breaks arearc common and can remove up to 92 type perennial species of aboveground vegetation nerney 1966 hil- annual grasses over native Californiass grasshopper fauna is rich com- bert and logan 1981 melanoplus sanguinsanguinipessanguimpesipeslpes cailcalifornia to other regions with is classified as a mixed feeder and may prefer pared north american is half of grasses or forbs depending upon the area sam- over 120 species joern 1989 about considered pled if M sanguinsanguinipessanguimpesipes feeds extensively on these grasshopper species are has the grasses within the SRPER given its phenol- rangeland species southern california ogy it will damage perennial grasses more richest grasshopper fauna of any region in cal- than annual grasses overall forage consump- ifornia strohecker 1968 still richness for tion was 140 aghakgha in 1994 which is an eco- the SRPER was very low compared to other nonomically damaging level according to onsager studies of north american grasslands few 1984 however using M sanguinsanguinipessanguimpeslpesipes in a studies report diversity indexes but richness shortgrassshortgrass prairie community quinn et al has been measured for other native north 1993 found significant reductions in grass american grasslands richness is nearly always 199619961 grasshoppers OF A california GRASSLAND 175

a A A 5 T 45 05 nymphs adults 6E 4 0a 045 a adults and 0C 35- 0 adults nymphs 42 04 14 3 035 QS0 25 0a 1 03 0 2 025 ii U 155 02 La 1- CY 015 05 01 i H i B 0 j 0 05 s s s s s s s s s s s s s s s 0 I1 CO ol01 cstc M CO m to f CO OS U 60 i0 0 0 0 0 0 0 0 0 0 0 0 0 c c 600goo B

500 B

ei nymphs 400 nadults 5 1 N adults 300 I 4 aduits and nymphs 0 E 200

00 1 too100 CL U 1 I 2 0 0 I 6 cp 10 1 I1 i fig 1 A grasshopper density for all sampling dates 0 and B biomass for dates collected a of proportion 0 00oo adults vs nymphs unknown for july 1993

fig 2 diversity A and species richness B for all dates in which collections were made higher than the value of 5 species determined here eg joern 1982 evans 1988 we feel it is likely that the grasshopper community is timothy famefainepainepame anthony joern and an anony- particularly species depauperate due to isola- mous reviewer are also greatly appreciated tion of the SRPER grasslands macarthur and we also thank robin wells and cedra shapiro wilson 1967 frequent burning of the grass- of the nature conservancy for their assis- lands may also help explain the low diversity tance as well as the use of the reserve this found in the SRPER on the other hand work was partially supported by academic burned sites contain more even species com- senate grants to R A redak positions than unburned grasslands and contain species not found in unburned sites porter literature CITED 1995 therefore we feel that to preserve the BURCHAM L T 1957 california rangeland california diversity of not only grasshoppers but presum- division of forestry sacramento ably many arthropods birds plants and other CAPINERA J L AND T S SECHRIST 1982 grasshoppers taxa it may be necessary to preserve larger acrididae of colorado identification biology and tracts of native grasslands furthermore the management colorado state universityuniveiuniver sity experiment station fort collins bulletin 584s effects of grasshopper herbherbivory in these grass-grass herbivoreivory EVANS E W 1988 community dynamics ofprairieof prairie grasslands must be accounted for in a well rounded hoppers subjected to periodic fire predictable tra- conservation effort jecjectoriestories or random walks in timetimeatime9 oikosbikos 52 283 292 HEADY H FE 1958 vegetational changes in the annual acknowledgments grassland type ecology 39 402 416 1977 valley grassland pages 491 574 in M G barbour and J major editors terrestrial vegetatiovegetationn we thank james bethke brian cabrera of california wiley interscience new york miriam cooperband kim hammond mari- HEWITT G B 1977 review of forage losses caused by anne van laarhoven carl matthies and james rangeland grasshoppers USDA miscellaneous publi- 1348 24 appp nichols for the cation participating in grasshopper HILBERT D W AND J A LOGAN 1981 A review of the collections helpful reviews by james bethke population biology of the migratory grasshopper 176 GREAT BASIN naturalist volume 56

melanoplus sanguinsanguinipessangwnipesipeslpes colorado state university tera acrididae in experimental plots acridaacarida 6 experiment station fort collins bulletin 577s 231 237 huennekeHULNNEKL L F 1989 distributions and regional pat- OTTE D 1981 the north american grasshoppers volumes terns of california grasslands pages 1 12 in L F I1 and 2 harvard university press heuenneke and H mooney editors grassland struc- PIELOU E C 1977 mathematical ecology wiley new ture and function kluwer academic publishers york and london dolddoidrechtdordrechtDoidDordrecht PORTER E E 1995 the grasshoppers of a california JACKSON L E 1985 ecological origins of cailcaliforniaCali forniass medi- native grassland a description of the community and terraneantenanean grasses journal of biogeography 12 its ecological importance unpublished master s the- 349 361 sis university of california riverside JOERNJOLRN A 1982 distributions densities and relative abun- QUINN M A ET AL 1993 effect of grasshopper orthop- dances of grasshoppers orthoptera acrididae in a tera acrididae density and plant composition on sandhillssandhills prairie prairie naturalist 14 37 45 growth and destruction of grasses environmental 1989 insect herbivoreherbherbivoryivory in the transition to califor- entomology 22 993 1002 nia annual grasslands did grasshoppers deliver the RISSER P G ET AL 1981 the true prairie ecosystem coup de grass pages 117 134 in L F huenneke hutchinson ross publishing co Stroudsstroudsburgburg PA and H mooney editors grassland structure and 557 appp function kluwer academic publishers dordrechtDordrecht SAVELLE G D AND H F HEADY 1970 mediterranean KEELEYKLELEY J E 1981 reproductive cycles and fire regimes annual species their responses to defoliation procla- pages 231 277 in H A mooney et al editors pro- mations oftheodtheof the lith international grassland congress ceceedings of the conference fire regimes and eco- 548 551 system propertiespropel ties USDA forest service general strohecker H FETALE ET AL 1968 the grasshoppers of cal- technical report WO 26 ifornia orthoptera acndoideaacridoidea bulletin of the cali- lalLArLATHROPHROP E W AND R F THORNE 1985 A flora of the fornia insect survey no 10 university of california santa rosa plateau southern california botanists press special publication 1 THOMPSON D C 1987 sampling rangeland grasshoppers macarthur R H AND E 0 WILSON 1967 the theory pages 219 233 in J L capineraCapmera editor integrated of island biogeography princeton university press pest management on rangelands a shortshortgrassgrass prairie princeton NJ perspective westviewWestview press NERNEYNLRNLY N J 1966 interrelated effects of grasshoppers WAGNER F H 1989 Graglazersgrazerszers past and present pages and management practices on shortshortgrassshoitgrassgrass rangeland 151 162 in L F huenneke and H mooney editors USDA Agnagriculturalcultural research service special report grassland structure and function kluwer academic z192 1 14 Publishpublisherseisels dordrechtDordrecht ONSAGER J A 1984 A method for estimating economic WHITE K L 1967 native bunchbunchgrassgrass stipa pulchra injury levels for control of rangeland grasshoppers on hastings reservation california ecology 48 with mamalathionlathion and carcarbarylcarbamylbaryl journal of range man- 949 955 agement 37 200 203 ONSAGERONSAGLR J A AND J E HENRY 1977 A method for esti- received 10 july 1995 mating the density of rangeland grasshoppers orthoportbopnorthop accepted 19 october 1995 great basin naturalist 562 0 1996 appp 177 179

SUMMER NOCTURNAL ROOST SITES OF BLUE GROUSE IN northeastern OREGON

kenneth J popperl eric C Pelrenpelrenl2peirenpelren1212 and john A CrawforcrawfordlCrawfordcrawford1grawforddl1

key words blue grouse dendragapus obscurusobscures nocturnal oregon roost

avian habitat studies frequently focus on blue grouse during summer we identified 20 diurnal habitat use because of ease of observa- independent nocturnal roost sites our objec- tion and high levels of activity associated with tive here is to describe these roost sites breeding and foraging nocturnal habitat use may be critical for all birds but has received STUDY AREA AND METHODS far less attention thus there is a need to better understand nocturnal habitat use espe- the study area is located in northeastern cially by crepuscular and diurnal birds and oregon 30 km north of enterprise in the wal factors that may contribute to this use lowaiowa whitman national forest in wallowagallowa blue grouse dendragapus obscurusobscures are county elevation ranges from 900 to 1500 in associated primarily with true fir abies sppapp with ridge slopes as great as 35 north facing and douglas fir pseudotsuga menziemenziesiimenziesiasii forests slopes are dominated by stands of douglas fir in mountainous regions of western north amer- and ponderosa pine pinus ponderoponderosaponderosdponderososd and ica johnsgard 1983 breeding season habitat common shrubs are mallow ninebark physo- associations often include nonforested and carpus malvaceusmalvamalvaceousceus snowberry symphoricar- shrub or steppe regions these birds are diur- pos albus and big huckleberry vaccinium nal with increased activity in the morning and membranaceum bunchgrassBunchgrass meadows pre- evening hours pekins et al 1991 determined dominantly bluebunchbluebunch wheatwheatgrassgrass agropyron that both diurnal and nocturnal winter boostsroosts spicatumspicatum and idaho fescue festuca idahoenidahoan of blue grouse were located in conifers blue sis occur on south facing slopes cattle graze grouse shifted from eating conifer needles in parts of the area during summer months winter to ground layer vegetation in summer resulting in variable grass cover and fall in northeastern oregon crawford et grouse were captured in walkinwalk in traps and al 1986 blue grouse summer habitat studies fittedbitted with poncho or necklace mounted radio have dealt with diurnal activities mussehl transmitters 15 to 18 g advanced telemetry 1963 bendell and elliot 1966 zwickel 1975 systems inc isantibisanti MN and telemetry sys- but nocturnal observations are minimal john- tems inc kequonmequon WI from june through son 1929 witnessed a brood fly into a tree august 1993 radio equipped juvenile birds apparently to roost overnight and blackford were t 500 g capable of flight and I1 mon of 1958 1963 observed 3 adult males flying age each radio equipped bird was located at into roost trees in spring where they pre- night once between 5 july and 3 august 1993 sumably stayed overnight blackford 1963 in addition to radio telemetry a spotlight was also observed a male displaying on the ground used to verify the location of the bird the exact approximately I1 h after dark zwickel 1992 roost site was identified by the presence of fresh suggested that ground roosting may occur fecal droppings when 2 or more grouse were particularly on breeding ranges where trees observed roosting together 10 in apart only are unavailable or before chicks are able to fly I1 roost site was counted for use in analyses to in the course of monitoring radio equipped ensure independence of locations

department of fisheries and wildlife nash hall 104 oregon state university corvallis OR 97331 address all coicorrespondcoreor respondencecorrespondericeerleeerice to eric C pelrenpeirenfellpelfpeif en

177 178 GREAT BASIN naturalist volume 56

RESULTS AND discussion greater use of ground cover forage and inver- tetebrates corresponded with observed diurnal twentyfivetwenty five radio equipped blue grouse and nocturnal use of ground habitat by blue and 38 birds without radios were located at 20 grouse in summer blackford 1963 suggested independent nocturnal roost sites table 1 that selection of roosting sites may result from the radio equipped birds consisted of 12 foliage preference and feeding habits motion adults and 13 juveniles sexes and ages of the sensors on grouse transmitters indicated that other birds were unknown all roost sites were some birds continued foraging on moonlit on the ground males usually roosted alone nights which implied that benefits of feeding whereas hens and juveniles frequently roosted outweighed energy loss associated with move- together sixteen of 20 independent boostsroosts ment or increased risk of predation including birds of all sex and age groups were pekins et al 1991 suggested blue grouse in grass of a relatively consistent height the selection of conifers as boostsroosts in winter may be others were in forbs n 2 and shrubs n based primarily on thermal properties of the birds 2 twenty three of 25 radio equipped sites higher temperatures during summer within 50 of potentially useful were in roost make thermal considerations less relevant to trees an adult female and a juvenile female survival than during winter the lowest tem- roosted 75 and 100 inm from trees respectively peraperatureture we noted at a nocturnal roost site was both easy flight distances for grouse adult ac well above the lower critical temperature males usually roosted closer to than 4c trees pekins other birds of 10cloc to 15c 1988 during daytime radio equipped birds hines 1986 found that 96 of juvenile and adult blue grouse mortalitiesmortalities were the were seldom located inm trees 1 of 614 grouse observations july august 1991 through 1993 result of predation in winter blue in E pelren unpublished data however almost trees may be less conspicuous or available to those the ground bergerud all birds flushed during the day landed in predators than on gratson 1988 andd pekins 1988 trees and conifer needles were found in crops and an observed of birds taken from the study area in august snow roosting only occasionally after heavy and september 1981 and 1982 crawford et al snowsnowstormsstorms however lack of snow and in- 1986 crawford et al also found plants such creased presence of grasses forbs and shrubs as prickly lettuce lactuca sernoldrriola yellow in summer along with cryptic coloration of salsify tragopogon dubicsdudubiusbius wild buckwheat blue grouse provide ground layer camou- eriogonum sppapp and snowberry sylphonsymphonsymphorisymphoria flage superior to that available in winter food carpos albus as well as short homed grasshop- availability may outweigh any increased risk of pers acrididae in at least 30 of 145 blue predation and account for use of nocturnal grouse crops in this area douglas fir needles ground boostsroosts by blue grouse in summer where were found in only 16 of the crops this selection of ground boostsroosts occurs

tailleTABLE 1 characteristics of20of 20 blue grouse nocturnal roost sites northeastern oregon july august 1993 adult male adult female juvenile male juvenile female no of roost sites 6 6 38 5 no of other birds 1 161 7 99eae plant cover at roost grass 4 6 38 3 forb 1 0 0 1 shrub 1 0 0 1 plant height m at roost median 0500.50 0450.45 0500.50 1 0750.75 range 0250.25 1201.20 025 1001.00loo 030oso0.30QSO 075a0 0300.30 1301.30 distance m to potential roost tree median 45 37537.5 50011 200 range 101.0iolo 40040.0 15015.0 75075.0tso 303.0 750175.01 50 1000100.0

ilrieludincludes diditafoidatatafoifor 5 ladio equipped juvenile males that weiewere with radioladioradlo equipped adult or juvenileorjuvenile females idoefidoeidocs not include 2radioaradio2 i adioadlo equipped juvenile malesindies that weiewelewere with radioiddioradlo equipped adult femalelemale ii does not include 3 ladioradioradlo equipped juvenile males that weiewelewere with radio equipped juvenile females 199611996 NOTES 179

acknowledgments HINESINES J E 1986 recruitment of young in a declining population of blue grouse unpublished dissertation this research was conducted as part of a university of albertaofalberta edmonton 256 appp grouse JOHNSGARDIHNSGARD P A 1983 the grouse of the world univer- blue winter ecology study funded by sity of nebraska press lincoln 413 appp the US forest service and oregon depart- JOHNSONIHNSON R A 1929 summer notes on the sooty grouse ment of fish and wildlife we thank R L of mount rainier auk 46 291 293 grouse jarvis for assistance during the development MUSSEHLUSSEHL T W 1963 blue brood cover selection oregon and land use implications journal of wildlife man- of this paper this is agricultural agement 27 547 555 experiment station technical paper 10673 PEKINSekinsdkinsSKINS P J 1988 winter ecological energetics of blue grouse unpublished dissertation utah state univer- literature CITED sity logan 141 appp PEKINSekins P J FE G LINDZEY AND J A GESSAMAN 1991 physical characteristics of blue grouse use BENDELL J F AND P W ELLIOT 1966 habitat selection winter trees and roost sites great basin naturalist 51 in blue grouse condor 68 431 466 244 248 BERGERUD A T AND M W GRATSON 1988 survival and ZWICKELOTCKEL FE C 1975 nesting parameters of blue grouse breeding strategies of grouse pages 473 575 in A T and relevance to populations 77 bergerud and M W gratson editors adaptive strate- their condor and population ecology of northern grouse 423 430 gies grouse poole university of minnesota press minneapolis 1992 blue in A P stettenheim and FE gill editors birds of north america no BLACKFORD J L 1958 Ternterritorialityaerntonality and breeding behav- the 15 academy of natural sciences philadelphia iorlor of a population of blue grouse in montana con- the dor 60 145 158 the american ornithologists union washington 1963 further observations on the breeding be- DC 28 appp havior of a blue grouse population in montana 4 august 1995 condor 65 485 513 received accepted 23 october 1995 CRAWFORD J A W V DYKE S M MEYERS AND T F HAENSLY 1986 fall diet of blue grouse in oregon great basin naturalist 46 123 127 gleatgreat basin naturalist 562 0 1996 appp 180 182 oochoristica SCELOPORI CESTODA linstowiidae IN A GRASSLAND population OF THE BUNCH GRASS LIZARD sceloporus SCAIARISSCALARIS phrynosomatidae FROM ARIZONA

stephen R coldcoidgoldberglgoldbeig1goldbergeGoldberglbergi charles R bursey2 chris T McAllister 3 hobart M smith4smitha and quynh A truongltruong1

neilkey words sceloporus scalaris bunch grass lizard phrynosomatidaephrynosomatzdae omboristicaoochonstica sceloponscelopori cestoda arizona

the bunch grass lizard sceloporus scalaris identified using a glycerol wet mount repre- wiegmann 1828 is known from the Huahuachucachuca sentativesentative cescustodescestodestodes were stained with hema dragoon santa rita and chiricahua moun- toxylintoxylon and mounted in balsam for further ex- tains of arizona the animas mountains of new amination voucher specimens were deposited mexico and in the sierra madre occidental in the US national parasite collection and sierra del nido of mexico usually above beltsville maryland 20705 USNPC 85053 1830 m but a few isolated valley populations terminology use is in accordance with margo- occur as low as 1200 m stebbins 1985 to our lis et al 1982 knowledge the only report of helminthshelminthes of this only I1 helminth was found the cestode species was a study of a high elevation 2438 oochoristica scelopori voge and fox 1950 2560 m chiricahua mountain population of prevalence of infection was 10 5 of 51 sceloporus scalaris slevinislevins by goldberg and mean intensity 121.2 0450.45 s range 1 2 bursey 19921992aa the purpose of our note is to in the only other investigation of helminthshelminthes report on a helminthological examination of a of S alarisscalarsscscalaris goldberg and bursey 1992a low elevation ca 1524 m grassland popula- reported finding tetrathyridia of the cestode tion of S scalaris slevinislevins smith 1937 from ari- mesocestoides sp prevalence 8 and larvae zona and to compare our findings with those of the nematode physaloptera sp prevalence of goldberg and bursey 19921992aa 3 that study was done on a coniferous for- we examined 51 S scalaris slevinislevins mean est high elevation population approximately snout vent length 51 343.4 mm s range 2500 m in the chiricahua mountains whereas 40 55 mm collected mostly by hand a few the current study considered a low elevation by dust shot on the sonoita plain elevation ca population ca 1524 m on the sonoita plain 1524 rn 3139n 11132w in the vicinity of located ca 126 kmkra SE of the chiricahua moun- elgin santa cruz county arizona specimens tains study site although both populations were deposited in the university of colorado harbored mutually exclusive helminth faunalfaunas museum of natural history boulder colorado additional work on larger S scalaris samples as UCM 572595728257259 57282 572845728657284 57286 57289- from these sites will be required to determine 57292 572955729857295 57298 573005730557300 57305 573075731057307 57310 the constancy of these differences 573135731657313 57316 573185731957318 57319 UCM 573185731957318 57319 oochoristica scelopori is a common cestode were collected 20 august 1989 others were of north american lizards and has been found collected 12 19 july 1990 in 14 other north american phrynosomaphrynosomatidtid the abdomen was opened and the esopha- lizards table 1 in addition amrein 1951 gus stomach and small and large intestines and telford 1964 reported finding 0 iceloscelo were removed from the carcass each organ pori in the xantusiids xantusiaXantusia henhenshawihenshaweshawi X was slit longitudinally and examined under a riversianariversiana and X vigilis measurements of dissecting microscope the liver and body various structures of these cescustodescestodestodes were strik- cavity were also examined each helminth was ingly different from the measurements as given

I1 departnientdepaitnient ofot biologyofbiology whittier college whittier CA 90608 address correspondence to this author 21cpaltulentlptiitincnt of biologyofbiology Peonpennpennsylvaniasylvania state university shenangoShenango valley campus sharon PA 16146 dlpaitiiicnt3department of biology texistextstexas wesleyanWesl eyancyan university 1201 wesleyan fort worth TX 76105153676105 1536 4epocepoPO biology University of Coloiacoloradodo bulderboulderBoul derdei CO 80309033480309 0334

180 199611996 NOTES 181

TABLE 1 definitive hosts of oochoristicaoochonstica sceloponscelopori in north america host locality prevalencePievalence reference crotaphytus colcoicollanscollariscollaraslaris california 10011loo100 11 telford 1970 Gamgambellagambeliabelia wishwislizeniiwishzennzenn california 402540 25 telford 1970 sceloporus claren arizona 5120sigo5 120 Goldbergoldbergetalgoldberggetalet al 1994 S graciosusgraciosus california not given voge and fox 1950 california 1077110 ttl771 telford 1970 idaho 222922 29 waitz 1961 idaho 111181 1118 lyon 1986 utah 51225 122 pearce and tanner 1973 S farrojarrojarroviijarrovndilvii arizona 10 47489 goldberg and bursey 1990 arizona 31313 131 goldberg and bursey 1992b arizona 5153025 15302 Goldbergoldberggoldbergetalgetalet al 1995a S magister arizona 3 walker and mathias 1973 texas 61176 117 Goldbergoldberggoldbergetalgetalet al 1995b S occidentoccidentalisoccidentalistalis california 20136520 1365 voge and fox 1950 california 23 27116 telford 1970 idaho 1121911 219 lyon 1986 oregon 33206033 9060gogo2060 white and knapp 1979 utah 91119 iliiii111 pearce and tanner 1973 S olivaolivaceusolivaceousceus texas 32613 261 Goldbergoldberggoldbergetalgetalet al 1995b S orcuthorcuttorcutti california 22167422 1674 goldberg and bursey 1991 S poinsettii texas 3031030 310 Goldbergoldberggoldbergetalgetalet al 1993 S scalaris arizona 1055110 ssi551 this paper S undulatesundulatus arizona 63486 348 goldbeigetalgoldberg et al 1994 uma inorinornataincornatanata california 71157 ils115 telford 1970 U notatanatata california 42102442 1024 telford 1970 Urourosaurussaurus graciosusgraciosus california 62346 234 telford 1970

in the original description of scelopori by GOLDBERG S R AND C R BURSEY 1990 gastrointestinal 0 yarrow lizard sceloporus voge and fox 1950 amrein 1951 reported helminthshelminthes of the spiny jar rounrovn farrofafrojarrofarjarjarroviijarrovitrouitrovitulivii cope american midland naturalist the average length of 25 mature cescestodescustodestodes from 124360124 360 365 X henhenshawihenshaweshawi and X vigilis to be 158215.8215 82 mm the 1991 intestinal helminthshelminthes of the granite spiny cescustodescestodestodes from X riverriversianarwersianasianaslana measured 33 37 lizard sceloporus orcuttiorcuttsorcutti journal of wildlife dis- mm telford 1964 indicated his cestode spec- eases 27 355 357 helminthes the grass lizard from xantusxantusndxantusiidndiidildlid lizards were less than 45 1992a helminthshelmidths of bunch imens sceloporus scalaris slevinslevimslevinislevins iguanidae journal of the mm both amrein and telford identified these helminthological society of washingtonofwashington 59 130 131 cescestodescustodestodes as 0 scelopori bursey and goldberg 1992b prevalence of the nematode spauligodon 1992 found Amreamreirsamreinsirs measurements of ces gigangiganticusticus oxyundaOxyoxyuridaoxyuridaeunda pharyngodomdaepharyngodonidae in neonatal todes from X henhenshawihenshaweshawi and X vigilis to approx- yarrow s spiny lizards sceloporus fattofartojarrojarroviijarrovnvii sauriasauna journal of parasitology 78 539 541 the measurements of bezzibezyi whereas iguanidaeiguamdae imate 0 GOLDBERG S R C R BURSEY AND R L BEZY 1995a telford s measurements of cescestodescustodestodes from X helminthshelminthes of isolated montane populations ofyarrow s fluerfiverriverriversianarwersianasianaslana approximated measurements of 0 spiny lizard sceloporus jarrojarroviijarrovtidilvii phrynosomatidae islandensis and suggested that X henhenshawihenshaweshawi X southwestern naturalist 40 330 333 and X vigilis be removed from the GOLDBERG S R C R BURSEY AND C T mcallister riversianarwersianariversianaslana gastrointestinal helminthes of leaving only phrynoso 1995b helminths of nine species host list of 0 scelopori sceloporus lizards phrynosomatidae from texas matidmacid lizards as hosts for 0 sceloporiseelopori journal of the helminthological society of washington 62 188 196 literature CITED GOLDBERG S R C R BURSEY AND R TAWIL 1993 gas- trotrointestinal helminthshelminthes of the crevice spiny lizard AMREIN Y U 1951 the intestinal entozoa of the night sceloporus poinsettpoinsettiipoinsettiapomsettnii phrynosomatidae journal of lizards of california and their mode of transmission the helminthological society of washington 60 unpublished doctoral dissertation university of cali- 263 265 fornia los angeles 162 appp 1994 gastrointestinal helminthshelminthes of sceloporus BURSEY C R AND S R GOLDBERG 1992 oochonsticaoochoristica lizards from arizona journal of the helminthologi- islandensis n sp cestoda linstowiidaelmstowndae from the cal society of washington 61 73 83 island night lizard xantusiaXantusia riversianariver siana sauriasauna xan LYON R E 1986 helminth parasites of six lizard species tusitusiidaeidae transactions of the american microscopi- from southern idaho proceedings oftheodtheof the helminthol- cal society 111 302 313 ogical society of washington 53 291 293 182 GREAT BASIN naturalist volume 56

MAHCOLISMARGOLIS L G W escilESCIIESCH J C HOLMES A M KURIS AND VOGE M AND W fox 1950 A new anoplocephalidanoplocephalic ces- CG A SCHAD 1982 the use of ecological terms in tode oochoristicaoochonstica sceloponscelopori n sp from the pacific parasitology report of an ad hoc committee of the fence lizard sceloporus occidentoccidentalisoccidentalistalis occidentoccioccidentalsoccidentaleoccidentalisdentalsalisallsails american society of parasitologists journal ofofparaokparapara- transactions of the american microscopical society sitology68sitology 68 131 133 6923669 236 242 PLARCPEARCE L R C AND W W TANNER 1973 helminthshelminthes of WAITZ J A 1961 parasites of idaho reptiles journal of sceloporus lizards in the great basin and upper parasitology 47 51 colorado plateau of utah great basin naturalist 33 WALKER K A AND D V MATTHIAS 1973 helminthshelminthes of 1 18 some northern arizona lizards proceedings of the SIEBBINSSTEBBINS R C 1985 A field guide to western reptiles helminthological society ofwashington 40 168 169 and amphibians houghton mifflin company boston WHITE II11 R L AND S E KNAPP 1979 helminth para- 336 appp sites of scelopormesceloporine iguanidae lizards from central TLLIORDTELFORD S R 1964 A comparative study of endopara oregon proceedings of the helminthological soci- sitismsinism among some southern california lizard popu- ety of washington 46 270 272 lations unpublished doctoral dissertation university of california los angeles 260 appp received 25 july 1995 1970 A comparative study of endoendoparasitismparasitism accepted 31 october 1995 among some southern california lizard populations american midland naturalist 83 516 554 great basin naturalist 562 V 1996 appp 183 185

POCKET GOPHERS DAMAGE SALTCEDAR TAMARIX ramosissima ROOTS

sara J ManningMannmanninglmanning1manningaingl1 brian L CasgasCashorecashorelcashoralcashore1horel1 and joseph M Szewczakszewczak22

key words saltsaltcedarcedar tamarix ramosissima pocket gopher thomomys bottaibottae tamarisk owens valley invasive plant exotic plant

saltcedarSaltcedar tamarix ramosissima ledealedeb during the winter of 1995 when foliage tamaricaceae is an invasive exotic woody was absent from saltcedarsaltcedar we observed that a shrub native to asia baum 1978 hickman few plants within a young even aged stand 1993 that has colonized extensive areas were dead some of the plants were leaning throughout the western united states robin- over supported by neighboring plants upon son 1965 brotherson and winkel 1986 inspection we observed that dead plant tap- saitcedarsaltcedarSaltSaItcedar possesses many characteristics that roots had been gnawed apart approximately 10 render it a nuisance plant brotherson and cm beneath the soil surface teeth marks were winkel 1986 and because it has been viewed clearly visible on the tapered stumps in addi- as a threat to native vegetation communities tion prolific gopher tunneling was evident researchers have examined its ecology car- within and around the saltcedarsaltcedar stand and man and brotherson 1982 brotherson and excavated dirt mounds were located near the winkel 1986 shafroth et al 1995 water con- dead saltcedarsaltcedar examination of growth rings of sumption robinson 1958 van hylckama 1970 plants within the stand showed the saltcedarsaltcedar davenport et al 1982 bureau of reclamation plants to be 7 years old in 1995 1992 and cost of control efforts brotherson in early april 1995 when saltcedarsaltcedar was just and field 1987 neill 1990 barrows 1993 it beginning to break bud we revisited the site is known to inhibit flows in creeks and springs to quantify the extent of animal damage and to robinson 1965 rowlands 1990 thus its capture and identify the species tunneling at spread has been detrimental not only to native the site we examined plants by working from vegetation but also to native wetland and one end of the stand toward the center every aquatic fauna neill 1983 saltcedarsaltcedar plant in approximately 12 of the although efforts are under way in the united stand was sampled for a total of 545 plants states to develop biocontrolbiocontrol agents using height was measured and then plants were insects that occur on saltcedarsaltcedar in its native tugged to detect the degree of below ground range deloach 1990 to date there have damage if tugged plants freely exited the soil been no reports of native herbherbivoresivores insects and had no attached live roots the damage or diseases causing saltcedarsaltcedar mortality herein was scored as fatal all of these plants appeared we report the first known mortality caused by dead no sproutingreresprouting was evident and each native mammals on saltcedarsaltcedar had a chewed taproot stump the diameter of our discovery occurred in owens valley which was measured and recorded if tugged california water has been exported from plants could be pulled from the ground easily owens valley located in the rain shadow but still hadbad live literalslateralslaterals above the chewed created by the sierra nevada range directly to taproot they were noted as sustaining severe its west since 1913 alteration of natural damage in these instances diameter of the water flows created conditions favorable to the largest chewed root was measured typically spread of saltcedarsaltcedar cashore 1985 babb these plants had many dead but a few living 1987 branches if tugged plants felt loose but could

ilnyonyo county water department 163 may street bishop CA 93514 beepeep springs college dyer NV 89010

183 184 GREAT BASIN naturalist volume 56 not be easily pulled from the soil they were TABLE 1 extent of gopher damage within a stand of saltsaltesaitecedar plants the owens valley scoledscored as sustaining minimum damage if salteedarsaltcedaredar in tugged plants were tightly rooted in the soil gopher of aagavg ht we assumed no root damage the majority of damage plants total cm s branches on plants in both these categories none 420 770 1289 333 appeared alive minimum 58 106 1203 305 results of gopher damage are listed in severe 29 53 1162 326 fatal 38 70 1134 253 table 1 nearly 23 of the plants sampled had experienced some degree of gopher damage all total 545 1000 1262 328 of these 707.07 0 were dead as a result of gophers 535.35 3 had been severely affected and 10610.610log 6 damage so the had been minimally affected the diameter of valley again we found gopher phenomenon is isolated to this single stand gopher chewed roots ranged from 11 mm to not in general the influence of fossorial ani- 55 mm and averaged 27727.727 7 mm mals on plant communities has received rela- gopher damage appeared to affect plant tively little research attention andersen 1987 height analysis of revealed significant variance although gophers may kill or slow the growth height differences between plants in the 4 in of saltcedarsaltcedar their longtermlong term effects on stand categories of damage F 44634.4634 463 FP 00040.0040 004 size and vigor or on saltcedarsaltcedar establishment 3 saltcedarsaltcedar plants not dam- df however in the meadow remain unknown other re- only slightly in aged by gophers tended to be searchers have found that pocket gophers damage taller than plants sustaining gopher cause significant woody plant mortality in a table 1 suggesting that gopher damage had variety of plant communities crouch 1971 been relatively recent marsh and steele 1992 cox and hunt 1994 the study aleaarea was searched for evidence ferguson and adams 1994 and huntly and of active gopher mounds early in the evening inouye 1988 and cantor and whitman 1989 7 active mounds were excavated and sherman reported that tree encroachment into mead- live traps baited with seeds and fresh plant ows was significantly slowed when gophers material were placed at the tunnel level these were present in meadows however given the traps were then covered with soil using local vigorous growth of saltcedarsaltcedar in general materials to prevent caveeave ins at hethe trap en- gopher damage may merely thin the stand trance trapping was done under the provision allowing the remaining individuals to continue of a scientific collector s permit issued by the unabated california department of fish and game traps were checked the following morning shortly literature CITED after sunrise from the 7 traps set in active gopher tunnels ANDERSEN D C 1987 below ground herbivoreherbherbivoryivory in natural communities a emphasizing fossorial 1 valley pocket gopher thomomys bottaibottae review I animals quarterly review of biology 62 261 286 s ingles 1965 was captured two other traptraps BABB D E 1987 report on the saltcedarsaltcedar control study were found packed with soil presumably by unpublished report prepared for the inyolosInyoLos ange- gophers the 4 remaining traps showed no les technical group inyo county water department obvious sign of gopher activity bishop CA BARROWS control 11 a story the first reported evidence C W 1993 tamarisk II success these data are restoration and management notes 11 35 38 inducing of a native species thomomys bottaibottae bmBAUM B R 1978 the genus tamarixtamanytomany israel academy of mortality in the exotic tamarix ramosissima science and humanities 209 appp the proximity of a saltcedarsaltcedar stand to gopher brotherson J D AND D FIELD 1987 tamarix impacts of a successful weed rangelands 9 110 112 habitat may increase its susceptibility to gopher brotherson J D AND V WINKEL 1986 habitat rela- damage at our site gopher mounds appeared tiontionshipsships of saltcedarsaltcedar tamarix ramosissima in cen- more extensive in the alkali meadow immedi- tral utah great basin naturalist 46 535 541 ately adjacent to the saltcedarsaltcedar stand than in BUREAU OF reclamation 1992 vegetation management the stand itself we subsequently made obser- study lower colorado river phase I1 report lower colorado region boulder city NV 103 appp at other even aged stands of saltcedarsaltcedar vations CANTOR L F AND T G WHITMAN 1989 importance of that occur adjacent to alkali meadows at other belowbelowgroundground herbivoreherbherbivoryivory pocket gophers may limit locations in owens valley and in deep springs aspen to rock outcrop refugia ecology 70 962 970 199611996 NOTES 185

CARMAN J G AND J D brothersonBROTH ersonEKSON 1982 comparison approaches to animal damage management in pacific of sites infested and not infested with saltcedarsaltcedar northwest forests USDA forest service general tamarittamanttamarix pentandrapenpentandriatandra and russian olive eleagnusEleelaeagnusagnus technical report PNW GTR 287 angustifoliaangustifoha weed science 30 360 364 NEILL W M 1983 the tamarisk invasion of desert ripar- CASHORE B 1985 saltcedarSaltcedar in the owens valley unpub- ianlan areas desert protective council educational lished report inyo county water department bulletin 83483 4 bishop CA 32 appp 1990 control of tamarisk by cut stump herbicide cox G W AND J HUNT 1994 pocket gopher herbivoreherbherbivoryivory treatments pages 91 98 in M R kunzmann R R and mortality of ocotillo on stream terrace bajada johnson and P S bennett editors tamarisk control and hillside sites in the colorado desert southern in southwestern united states proceedings of the california southwestern naturalist 39 364 370 tamarisk conference 2 3 september 1987 special CROUCH G L 1971 susceptibility of ponderosa jeffrey report 9 university of arizona cooperative national and lodgepole pines to pocket gophers northwest park resources studies unit tucson science 45 252 256 ROBINSON T W 1958 phreatophytes USU S geological sur- DAVENPORT D C P E MARTIN AND R M HAGAN 1982 vey water supply paper 1423 84 appp evapotranspiration from riparian vegetation water 1965 intiintroductionanti eduction spread and aerial extent of relations and irrecoverable losses for saltcedarsaltcedar jour- salteedarsaltcedarsaltsaitsaltesaiteedarcedar tamanyTatamarixmanxmany in the western states USU S geol- nal of soil and water conservation 37 233 236 ogical survey professional paper 491 A DELOACH C J 1990 prospects for biological control of ROWLANDS P G 1990 history and treatment of the saltcedarsaltcedar tamarix sppapp in riparian habitats of the saltcedarsaltcedar problem in death valley national monu- southwestern united states pages 307 314 in E S ment pages 46 56 in M R kunzmann R R john- delfosse editor proceedings of the ath7th interna- son and P S bennett editors tamarisk control in tional symposium on biological control of weeds southwestern united states proceedings of the 6 11 march 1988 rome italy tamarisk conference 2 3 september 1987 special FERGUSON D E AND D L ADAMS 1994 effects of report 9 university ofarizonaofarizona cooperative national pocket gophers bracken fern and western cone- park resources studies unit tucson flower on survival and growth of planted conifers SHAFROTH P B J M FRIEDMAN AND L S ISCHINGER northwest science 68 241 249 1995 effects of salinity on establishment of populus HICKMAN J C EDITOR 1993 the jepson manual higher fremonfremontiifremontiatii cottonwood and tamarix ramosissima plants of california university of california press saitsaltsaltcedaisaltcedarcedar in southwestern united states great basin berkeley 1400 appp naturalist 55 58 65 HUNTLY N AND R INOUYE 1988 pocket gophers in VAN HYLCKAMA TTEE A 1970 water use by salt cedar ecosystems patterns and mechanisms bioscience water resources research 6 728 735 3878638 786 793 INGLES L G 1965 mammals of the pacific states stan- received 11 june 1995 ford university press stanford CA 508 appp accepted 19 january 1996 MARSH R E AND R W STEELE 1992 pocket gophers pages 205 230 in H C black editor silvicultural great basin naturalist 562 0 1996 appp 186 187

SALTCEDAR TAMARIX ramosissima AN UNCOMMON HOST FOR DESERT MISTLETOE phoradendron californicumCALIFORNICUM

sandra L haighlhaighthaigh1

ketikeifkey words phoradendron californicumcalifornicum tamarix ramosissima mistletoe saltsaltcedarcedar host parasite

the genus tamarixtamant saltcedarsaltcedar contains 16 october 1995 voucher specimens from I1 approximately 54 species of phreatophytic parasite and host are deposited in the depart- plants whose origins are in europe asia and ment of biological sciences herbarium uni- africa several members of the genus were versity of nevada las vegas accession num- introduced into the united states in the early ber38971ber 38971 1800s mainly as ornamental plants approxi- the host tree was growing in a canyon mately 8 species have since escaped cultiva- approximately 2 in from a small flowing stream tion and have become naturalized to varying on quartz monzonite derived soil the first degrees baum 1967 tamarix ramosissima mistletoe clump measured 33 cm long X 32 cm ledealedeb has become established in liparianriparian areas high X 14 cm wide and was growing on the throughout the west and southwest where it southwest side of a branch 212.1gi91 in above the has proven to be an aggressive invader that ground the branch to which the mistletoe eventually displaces native vegetation was attached measured 525.2 cm in diameter desert mistletoe phoradendron califorcahformcumcalifornicumnicum and 16216.2 cm in circumference the length of nutt is a native parasitic plant that grows on the branch from trunk to point of mistletoe several species of riparian plant hosts its range attachment was 212.1glgi in the trunk base of the includes southern nevada southwestern utah 5 m high saltcedarsaltcedar measured 8 cincm in diameter southeastern california southwestern arizona and 29 cm in circumference which would and northern baja california sonora and indicate an age of approximately 24 yr based sinaloa benson and darrow 1981 previously on average value of california and arizona published information on hosts for desert sites as reported by smith 1989 the and2nd mistletoe include blumer 1910 shreve and mistletoe also faced southwest and was located wiggins 1964 walters 1976 daniel and on the main trunk of the tree 9.9 in above the butterwick 1992 and overton 1992 none ground it was a newly sprouted plant that con- of whom mentions T ramosissima holland et sisted of only 12 stems the longest of which al 1977 and benson and darrow 1981 state measured 4 cm both mistletoesmistle toes and the host that saltcedarsaltcedar and the introduced tamatamarisksrisks tree appeared to be healthy actively growing are possible hosts while munz and keck 1965 specimens the parasites were young plants and mcdougall 1973 list tamarix but men- and were a more vivid green than other mis tion no particular species cohan et al 1978 tletoes in the area sex of the mistletoesmistletoes could state that P califorcalifornicumcahfornicumnicum does not occur in not be determined saltcedarsaltcedar this paper describes 2 occurrences other hosts for P californicumcalifornicum at this site of P califorcalifornicumcahformcumnicum on T ramosissima in south- include catclaw acacia acacia greggiigregggreggioii honey ern nevada mesquite prosopis glandulosaglandulose and creosote I1 found the ist parasite and host specimen bush larrea dentatatritridentatatridentate although many other on 27 june 1995 at hiko springs in clark tamarix trees occur here none have been in- county nevada approximately 11 kmkin west of fected by mistletoe desert mistletoe is usually laughlin along state highway 163 3894000 spread from host to host by birds which ingest N 711650 E at an elevation of 605 inm fig 1 the seeds and later defecate them onto a branch A and2nd specimen was found on this host tree on two bird species that occur frequently at this

idepartmentepaitmcnt of biologicalofbiological sciences 4505 maryland parkway box 454004 las vegas NV 89154400489154 4004

186 199611996 NOTES 187

WA literature CITED BAUM B R 1967 introduced and naturalized tamatamarisksrisks in the united states and canada tamaricaceae baileyabaileysbaileyal515 19 25 BENSON L AND R A DARROW 1981 trees and shrubs of the southwestern deserts university ofarizonaof arizona press X tucson 416 appp BLUMER J C 1910 mistletoe in the southwest plant X in world 13 240 246 COHAN D R B W ANDERSON AND R D OHMART 1978 avian population responses to salt cedar along the lower colorado river pages 371 382 in R R johnson and J F mccormick editors strategies for pro- 4 tection and management of floodplainfloodplain wetlands and other riparian ecosystems USDA forest service ilk general technical report WO 12 DANIEL T F AND M L butterwick 1992 flora of south mountains of south central arizona desert plants z 103 99 119 L HOLLAND J S R K GRATER AND D H huntzinger ZP 1977 flowering plants of the lake mead region 0 0 southwest parks and monuments association popular series no 23 49 appp 77 MCDOUGALL W B 1973 seed plants of northern arizona the museum of northern arizona flagstaff 41aa41.41 594 HHPP fi ll11 appp 1 M I MUNZ P A AND D D KECK 1965 A california flora university of california press berkeley 1681 appp OVERTON J M 1992 host specialization in desert mistle- fig 1 parasite phoradendron califorcalifornicumcaliformcumnicum growing on toe phoradendron califorcalifornicumcaliformcumnicum bulletin of the eco- host plant tamarix ramosissima logical society ofamericaof america 73 293 SHREVE F AND I1 L WIGCINSWIGGINS 1964 vegetation and flora of the sonoran desert volume I1 stanford university press palo alto CA 840 appp site and have been seen feeding on mistletoe SMITH S D 1989 the ecology of saltcedarsaltcedar tamarix chi and perching in saltcedarsaltcedar are the phainopepla bensisnensis in death valley national monument and phainopepla nitensmtens and northern mocking- lake mead national recreation area an assessment bitens of techniques and for salteedarsaltcedarsaltsaitsaltesaitecedar control in polyglottospolyglottous monitoring edar in bird mimus polyglottos personal observation the park system contribution CPSUUNLV 04103 national park serviceuniversityService University of nevada las acknowledgments vegas 65 appp WALTERS J W 1976 A guide to mistletoesmistletoes of arizona and new mexico USDA southwestern I1 wish to thank wesley niles for help with forest service region and management 7 identification of and review of the forest insect disease specimens review PP manuscript and delbert wiens who provided information on mistletoe hosts this project received 6 november 1995 was funded by a research grant provided by accepted 4 march 1996 the harry reid center for environmental studies las vegas nevada great basin naturalist 562 0 1996 appp 188 189

BOOK REVIEW

wild plants of the pueblo province explor- and yet it is so easily applied and can be ing ancient and enduring uses william observed in the field when adequate informa- W dunmire and gail D tierney foreword tion is provided the main focus of the text is by gary paul nabhan museum of new the center section that includes photographs mexico press santa fe NM 1995 290 appp and descriptions of 73 plants line drawings 199519.9519 95 softtacksoftbacksoftback accompany each plant treated the technical descriptions are somewhat brief but the illus- this book immediately appears field wor- trationstrations provide enough detail that field iden- thy and feels good in the hands and that s tification can be made easily in most cases simply judging the book by its cover I1 once perhaps one of the most valuable sections is opened there is much to praise about this text the annotated plant list included at the end of the authors have succeeded in putting together the book in an easy to read format a great deal a wonderfully interesting and well written field of information is concisely summarized for over guide for the lay person as well as a useful ref- 300 plants the chart is subdivided into 7 gen- erence for serious students and professionals eral categories of plant use ie food and bev- interested in ethnobotany of the southwest erage medicine construction etc with infor- within 9 chapters of text an illustrated section mation given on how each plant is used by involving about 73 plants and an extensive specific pueblos the chart is well referenced chart summarizing plant uses the reader learns and includes original citations for every use of the ecology representative flora ethnobotany A brief yet well organized analysis of the and cultural history of the pueblo province changes in plant utilization that occurred with the original intent of the book was to provide the spanish colonization in the southwest is a guide to commonly seen plants of bandelier provided in chapter 3 the authors take a very national monument and the pajarito plateau complex history and present it in the context in central new mexico and a discussion of the of plant ecology it provides an informative plants prehistoric and recent uses the authors view of the ecological consequences of the have surpassed this goal collision of cultures contemporary culture the 9 chapters reveal a cohesive and inter- plant use and ecological modification are also esting history of the people plants and land included in this text two chapters provide itself ample information provides the reader insightful information on current cultural and insight as to how these elements interact and ecological issues throughout the text and what the consequences of those interactions reflected in the annotated plant list as well have been and continue to be it is easy not the authors have attempted to treat religious only to move through the spatial and geo- and ceremonial plant uses with appropriate graphical regions but to enjoy a voyage in respect an added benefit of the book is the time as well and feel as if you were there line authors personal association with individuals drawings photographs and maps lend addi- in different pueblo tribes their sense of respect tional interest to the text although there is a and honor for these cultures is felt throughout great deal of information given about vegeta- the book tive zones human history and other topics our only complaint relating to this text is the authors have retained the importance of the lack of references citing specific informa- plants by referencing particular species wher- tion it is quite difficult to identify references ever appropriate the chapter on indicator for much of the information included within species is particularly interesting and useful the text with the exception of the chapter dis- this is a subject that few field guides address cussing indicator species A bibliography with

188 199611996 BOOK REVIEW 189

145 references is included at the end of the as a reference for plants and their uses by cul- book but it is difficult to relate these refer- tures of the southwest and in a travel file as it ences to particular chapters and specific infor- gives suggestions for specific hikes located in mation this omission weakens the usefulness the pueblo province for anyone interested in of the book as a reference for serious students plant ecology taxonomy ethnobotany cultural it may be that the authors consciously omitted anthropology or simply those with a general citations in an effort to allow the text to flow love for the southwest this book is highly rec- more easily but it is a constant frustration omommendedmended it is well written informative and when one is interested in identifying sources aesthetically delightful A list of suggested reading is included at the end of each chapter but no reference is given reneerenge van buren to original sources that support specific facts kimberly hamblin hart in the preface the authors do mention many department of botany and range science sources that contribute in a general way brigham young university overall this book is one that should be provo UT 84602 included in a field book box on the bookcase university OF NEVADA RENO department OF anthropology HISTORIC preservation biological RESOURCES RESEARCH CENTER DIVI- SION OF continuing EDUCATION

the university of nevada reno offers continuing education training courses in heritage resources management courses are designed for professionals in cultural and natural heritage management positions in the public and private sectors the program is conducted in cooperation with the advisory council on historic preservation the bureau of land management the national park service and the US forest service the following information is offered on one of the upcoming courses

ECOSYSTEM management

303130 31 may 1996 reno nevada 900 am 400 pm fee250Fee 250 registration deadline 2 may 1996

instructor peter F brussard phd is chairman of the biology department at the university of nevada reno and director of the nevada diversitybiodiversityBio initiative housed in the biological resources research center brussard is a founding mem- ber and past president of the society of conservation biology and recognized as a leading authority in conservation biology and ecology

ecosystem management is only recently beginning to be understood and used this course will address the scientific basis for ecosystem management as well as the steps for managing areas so that biological diversity and ecosystem services remain conserved while human needs are also met ecosystem management focuses on systems as a whole rather than simply on the parts and involves the public in setting management goals it represents a shift from linear comprehensive management to adaptive management

for further information phone 1 70702784404617027844046702 27784 8444046 or FAX 1 70702784480117027844801702 27784 8444801 to register call 180023389291 800 233 8929 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 richard 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 A cover letter accompanying the man- first author for citations having more than two uscript must include phone numbers of the author authors submitting the manuscript and FAX number and acknowledgments under a centered main emailE mail address when applicable the letter must headingbeading include special publication numbers when also provide information describing the extent to appropriate which data text or illustrations have been used in literature CITED also under a centered main other papers or books that are published in press heading lists references alphabetically in the fol- submitted or soon to be submitted elsewhere lowing formats authors should adhere to the following guidelines manuscripts not so prepared maybemay be returned for mack G D and L D flake 19198080 habitat rela- revision tiontionshipsships of waterfowl broods on south dakota manuscript preparation in general the great stock ponds journal of wildlife management basin naturalist follows recommendations in 4469544 695 700 CBE scientific style and format the manual for sousa W PE 1985 disturbance and patch dynamics publishers authors editors and ath6th edition on rocky intertidal shores pages 101 124 in council of biology editors 11 south lasalle inc S T A pickett and PE S white eds the ecolo- street 1400 chicago IL 60603 USA PHONE suite gy of natural disturbance and patch dynamics FAX we 3122010101312 201 0101 3122010214312 201 0214 do however academic press new york differ in our treatment of entries in literature cited coulson R N and J A witter 1984 forest ento- vol 51 no 2 of this journal authors may consult mology ecology and management john wiley for specific instructions on format these instruc- and sons inc new york 669 appp tions guidelines FOR manuscripts SUBMITTED TO THE GREAT BASIN naturalist are printed at the TABLES are double spaced on separate sheets and back of the issue also check the most recent issue designed to fit the width of either a 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text acknowledgments literature cited appendices A CHARGE of 50 per page is made for articles tables figure legends figures published the rate for individual subscribers will TITLE PAGE includes an informative title no longer be 35 per page however manuscripts with com- than 15 words names and addresses of authorsanthors a plex tables andor numerous half tones will be running head of fewer than 40 letters and spaces assessed an additional charge reprints may be pur- footnotes to indicate change of addressofaddress and author chased at the time of publication an order form is to whom correspondence should be addressed if sent with the proofs other than the first author FINAL CHECK abstractstatesABSTRACT states the purpose methods results cover letter explaining any duplication of and conclusions of the research it is followed by information and providing phone numbers 6 12 key worwordsds listed in order of decreasing FAX number and emailE mail address importance to be used for indexing 3 copies of the manuscript and WordwordperfectPerfect TEXT has centered main headings printed in all diskette capital letters second level headings are centered conformity with instructions in upper and lowercase letters third level head- photocopies of illustrations ings begin paragraphs issn0017ISSN 001736140017 3614 GREAT BASIN naturalist vovolvoi 56 no 2 april 19951996 CONTENTS articles selecting wilderness areas to conserve utah s biological diversity diane W davidson william D newmark jack W sites jr dennis K shiozawa eric A rickart kimball T harper and robert B keiter 95 nutrient distribution in quercus gambeliigambelligambelii stands in central utah A R tiedemann and W PE clary 119 comparsioncompursionComparsion of two roadside survey procedures for dwarf mistletoesmistletoes on the sawtooth national forest idaho robert L mathiasen james T hoffman john C guyon and linda L wadleigh 129 effects of douglas fir foliage age class on western spruce budworm oviposition choice and larval performance kimberly A dodds karen M clancy kathryn J leyva david greenberg and peter W price 135 trypanoplasma atrariaatwaria sp n Kinetoplastkmetoplastidakinetoplastidaidalda bodonidaeBodon idae in fishes from the sevier river drainage utah J stephen cranney and richard A heckmann 142 geographical review of the historical and current status of ospreys pandion haliahaliaetushaliaeetusetus in utah clark S monson 150 effects of turbidity on feeding rates of lahontan cutthroat trout oncorhynchus clarki henhenshawihenshaweshawi and lahontan redsidebedside shiner richardsoniusRichardsonius egregius gary L vinvinyardyard and andy C yuan 157 pogonomynnexpogonomyrmex owyheei nest site density and size on a minimally impacted site in central oregon peter T Ssoulesouie and paul A knappcnappxnapp 162 field measurements of alkalinity from lakes in the uinta mountains utah 1956 1991 dennis D austin 167 density biomass and diversity of grasshoppers orthoptera acrididae in a california native grassland eric E porter richard A redak and H elizabeth braker 1721 72 notes summer nocturnal roost sites of blue grouse in northeastern oregon kenneth J popper eric C pelren and john A crawford 177 oochoristica scelopori cestoda linstowiidae in a grassland population of the bunch grass lizard sceloporus scalaris phrynosomatidae from arizona stephen R goldberg charles R bursey chris T mcallister hobart M smith and quynh A truong 180 pocket gophers damage saltcedarsaltcedar tamarix ramosissima roots sara J manning brian L cashore and joseph M SszewczakZ 183 saltcedarsaitcedarSaltSaItcedar tamarix ramosissima an uncommon host for desert mistletoe phoradendron califorcalifornicumcalifomicumnicum sandra L haigh 186 book review wild plants of the pueblo province exploring ancient and enduring uses william W dunmire and gail D tierney reneerenae van buren and kimberly hamblin hart 188