Full Issue, Vol. 50, No. 3

Volume 50 | Number 3 Article 15

10-31-1990 Full Issue, Vol. 50, No. 3

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VOLUME 50 NON 3 OCTOBER 1990

BRIGHAM YOUNG university GREAT BASIN naturalist editor

JAMES R BARNES 290 MLBM brigham young university provo utah 84602

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MICHAEL A BOWERS JEANNEJEANNECC CHAMBERS JEFFREYJEFFREYRR JOHANSEN blandy experimental farm USDA forest service research department of biology university ofvirginia 860 north 12th east john carroll university box 175 logan utah 84322800084322 8000 cleveland ohio 44118 boyce virginia 22620 paulgPAULCPAUL C MARSH BRIAN A MAURER JIMMIEJIMMIERR PARRISH center for environmental studies department ofzoology department ofzoology arizona state university brigham young university brigham young university tempe arizona 85287 provo utah 84602 provo utah 84602

other associate editors are in the process of being selected

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VOLUME 50 31 OCTOBER 1990 no 3

SPROUTING AND SEEDLING establishment IN PLAINS SILVER SAGEBRUSH ARTEMISAARTEMISIA CANA PURSH SSP CANA

C L wambolt T P waltonwaiton and R S white 2

ABSTRACT the importance and nature of vegetative productionreproductionle was compared with seedling establishment in plains silver sagebrush arveArleartemishartemismarlemisiaArtemmisiamislaism cana pursh sspasp canacanucang sixty three percent of plants excavated originated flomfromhiom rhizomesrhizomes sites that experienced habitat disturbance did not have a significantly different number of plants originating from vegetative reproduction than did undisturbed sites parent rhirhizomeszomes wereere significantly older than taproots which were significantly older than aboveground stems rhizome dystesystesystemsillsliis were spread 333 3 times that of plant height seventyse entaentv nine percent of rhizomatous daughter plants were 100 cmem or less from parent plants up to 52 sprouts were found on one rhizome seedling establishment was greatest during wet growing seasons and vegetative reproduction was greatest during dndry yearsvears

sagebrush artemisia L taxa are among folfoiformol each taxon individually the most perni- the most important plants on langrangelandselands ciouselous weeds generally grow from under- of the western united states beetle 1977 ground lootsroots rhirhizomeszomes and buds cook plains silver sagebrush artemisia cana 1983 thus these are important traits to un- pursh sspasp cana is a major consideration in derderstandstand in successful rangeland taxa para- the management of rangelands in the north- doxically vegetative reproduction in sage ern great plains this is due to the taxon s brush taxa has not been previously studied in competitive nature with livestock forages and detail despite the importance of these taxa its importance as a habitat component for beetle 1977 mcarthur and plummer 1978 several wildlife species together with the and their obvious reproductive success har- other two subspecies of silver sagebrush vey 1981 understanding sagebrush repro- mountain silver sagebrush A cana sspasp cisvisoisols ductive success would provide insight into cidulabidula osterhout and bolander silver sage- plant population dynamics throughout west- brush A cana sspasp bolanboianbolandenderiderl gray ward ern north america mott 1979 our objective this complex is encountered on millions of was to assess the importance and nature of hectares min 13 western states and 2 canadian vegetative reproduction sprouting versus provinces harvey 1981 seedling establishment in plains silver sage- the literature young and evans 1972 brush bostock and benton 1979 went 1979 pro vides contradictory evidence as to whether studSTLDSTUDY SITES description seed or vegetative reproduction is more im porlantportant for survival of plants displaying both six study sites were selected in drainages habits in andaridarld and semiarid environments of the tongue and yellowstone rivers near accordingly this will have to be determined miles city in southeastern montana the sites

deparinidepartnidep trtmentt ofaniinalof vannalvnnnal indaridarld rangeR nireolge sciencess montana slatestate enierLnier sitgit bozenianbozegianBOzeniannidiinidai montana 5971739717591 au 2u2lsdaSDA agricultural research semetservice fort keogh livestockLn estock andd rangrange riresearchsearell station mllescltmiles cit monimontanamont tili1111 59301

201 202 clwamboltetalC L WAMBOLT ET AL volume 50 were considered typical in climatic and each excavated plant aging of sagebrush was edaphic relationships of plains silver sage- feasible despite the difficulty created by com- brush habitats in the northern great plains mon stem splitting and layering ferguson harvey 1981 soils at each study site are 1964 texturally heterogeneous but are largely a mo- to determine whether differences existed saic of loamscoams with silty clay loamscoams predomi- in the number ofsprouts to seedlings over the nant three sites yellowstone river frigid six study sites we conducted a paired stu- ustic torrifluvents lower black springs dent s t test FP 025025.025 A chi square analysis mixed calcareous frigid ustic torrifluvents P 0505.05 was performed to learn if fire or and lower flood fine montmorillonitic bor ice action on three of the sites was significant ollic camborthids had experienced fire or ice in determining the ratio of sprouts to seed- scraping and shearing in the preceding five lings compared with three undisturbed sites years no evidence of such disturbance was analysis of variance ANOVA was used for found at the remaining study locations lig- comparison of age and growth means among nite creek camborthidCamborthid torrifluvents paddy plant parts duncan s multiple range test pro- faye fine montmorillonitic borollicBorollic cam tected by a prior F test was used for compar- borthborthidsids and moon creek fine montmoril ing treatment means leniticlonitic borollicBorollic natriargids all sites have isolated plant excavations received periodic cattle grazing the area large well has an average annual precipitation of 340 two established plants at the the mm with peak precipitation received in may lower flood site were subject of a com- in plete plants and june root excavation were subjec- tively selected based upon two criteria 1 the plant had to be relatively isolated from METHODS other large plains silver sagebrush plants to mini- transect excavations mize major competitive influences and 2 there had to be an abundance of small plains each study site a plains silver sagebrush at silver sagebrush plants surrounding the po- plant 16 to 40 cm height was in located at each tential parent plant the two plants selected 5 interval along 25 m transects 4 estab- ni for excavation were slightly more than 1 of this in lished plants size were selected be- in height A 5 m area around each of the rhizomatous cause connections to parent large plants was excavated so that all roots plants if present were still readily apparent including rhirhizomeszomes of all smaller plants were the 20 plants located at each site were exca- exposed the size of each plant and the vated to determine whether they had sexual root distribution from the excavations were or asexual origins mapped to differentiate seedlings and roots were carefully excavated by hand so sprouts although we did not analyze the that fragile rhizome connections remained in- data statistically our direct observation of tact to determine if plants were of indepen- the root networks facilitated interpretation dent origin or connected to another plant of the transect data rhizomesRhizomes were generally found in the top 10 cm of soil while taproots were excavated to a RESULTS depth of 1 in or an impenetrable layer plants without connecting rhirhizomeszomes were consid- transect excavations ered to have originated from seed plant plants arising from rhirhizomeszomes were more height length of rhizomesrhizomes and stem and rhi- abundant than those that grew from seedlings zome diameters were measured on all origi- P 025025.025 table 1 approximately 63 ofthe nally located plants and those plants to which excavated plants were connected by rhizomesrhizomes they were directly connected root distribu- to an established plant or rhizome system tion from each excavation was mapped within counts of annual rings established that the a grid and line sketches of each plant were rhizomatous connections were one to four drawn samples for age determination fer- years old usually a large established parent guson 1964 were taken from stem root rhi- plant was the source of rhirhizomeszomes connect- zome and connecting rhizome sections of ing either single plants or a series of sprouts 1990 SPROUTING AND SEEDLING establishment IN SAGEBRUSH 203

sprouts

ground e ad titio s bu s

subsurface connecting rhizochizo

0 scm 16cm

fig 1 graphic example ofan excavated sprout connected to a parent plant and other offspring original or oldest material is on far right

TABLE 1 A comparison of the number of rhizomatous ordingaffordingaftafiatt an advantage to taprootingtaprooting plants in to non rhizomatous plains silver sagebrush plants found at reaching each study site deeper more favorable conditions the three sites disturbed by fire or ice action study sitesiteasite1 were compared with the three undisturbed 1 2 3 4 5 6 total sites to learn whether the ratio of sprouts to Rhizomatous 2 13 13 15 7 13 14 757533.3 seedlings changed with disturbance no sig- rhizomatousnonrhizomatousNon 7 7 5 13 7 6 454511 nificantnificant differences P 05os05.05 in numbers of sites are numbered as follows I1 yellowstone river 2 lower blaekblackblace plants arising from rhirhizomeszomes due to distur- springs 3 lower flood 4 lignite creek 5 paddy faye 6 moon creek bance were found plants with rhizome connectioiiiconnections alive or dead to other plants significant P 025 differences between rhizomatous and rhizomatousnoninonnonrhizomatous generally an elaborate subsurface rhizome plant totals by student s t bestaretestaretesttegt are followed by different lettelletterss system was found that was older than aboveground stems there were significant P fig 1 however some plants were from a 05os05.05 age differences among plant stems tap- series of sprouts along a rhizome presently roots and parent rhizomesrhizomes table 2 over all terminated with a dead or decadent stump SIXsix study sites aboveground stems were one site lignite creek was different in that three to five years younger than taproots and it had a majority of connectednonnonconnected individuals associated rhizomesrhizomes parent rhirhizomeszomes with di- table 1 reduction ofavailable soil moisture rectly connected sprouts were significantly due to clay pan soils overlying extensive older than taproots taproots and rhirhizomeszomes gravel at lignite creek might explain the without direct connections to a parent plant difference plant water use would be less were not significantly different in age from favorable with this condition at the surface each other 204 C L WAMwamboltetalWAMBOLTBOLT ETETALAL volume 50

40 36

3 0

N U M 20 b e r 12 10

0 1 0500 50 5110051 100 101150101 150 151200151 200 201 and over centimeters

fig 2 number of distances encountered between parents and traceable sprouts of rhizomatous plains silver sagebrush over six study sites

TABLE 2 age relationships ofaboveofabove and belowbelowgroundground TABLE 3 growth relationships of above and below parts of plains silver sagebrush plants from the six study ground parts ofrhizomatous plains silver sagebrush plants sites from the six study areas

mean age standard number of mean range number of plant part years deviation samples plant part erncm cm samples stems 3413.4134 20 204 plant sprout height 32 115911 59 155 taproots 69691 31 28 lateral distance to 2 parent rhizome 88 37 68 parent connection 2 7878b 1427714 277 61 rhizome systemsystem3systems3 606 01 25 128 lateral spread of significant P 05 rdeanmeanrnean differences by duncan s multiple range test are rhizomerhizornezonne system3systems 1051056 1136911 369 90 followed by diorentdi&rentdiftei ent letterletterss significant P 05 mean differences by duncan s multiple range 210rizome fi test are uiizoine originatingkoin01 ignidtingoriginating hoinkoinholn0111 a parent plant 01or dead stump to whichtoshichtowhich sprout was followed by different letters dndirectlycctlycutly connected 2laterallateral distance from parentpaipal ent plant 01or i hizomerhizome to nearest sprout on rhizome rhizome other than in above sections in 2 system expressed as a mean ofcfallofallallailali plants with this growth habit total lateral extent ofallcfallof all rhizomesrhi zomes in an excavation expressed as a mean ofallcfallof allaliail plants with rhizome systems rhizome extension was greater than aboveground heights P 05os05.05 table 3 even in 100 cm in length followed by 0 50soem older plants had largest aerial the cm that the por- distance 20 tions rhizome length from the selected plant to the parent plant averaged 242.4 times that isolated plant excavations of plant height lateral spread of total the the extensive sprouting nature of plains rhizome system averaged 33 333.3 times that of silver sagebrush was apparent after excava- plant height tions had been completed in areas surround- figure 2 summarizes the number of dis- ing two large isolated plants most roots were tances encountered between parent plants part of a shallow complex underground net- and traceable sprouts the largest proportion work of interconnected rhirhizomeszomes that often 59 of these connections were from 50 to included several smaller nearly independent 1990 SPROUTING AND SEEDLING establishment IN SAGEBRUSH 205

3 2 2

1 1 1 1 2 19aj1j 2 3 1

NUMBERS IN r HEIGHT 4 CATEGORY 2araar 2 2 S 4sas cm 14 4 3sas 45 cm 13 S 2 3 25 3scm 17 2 2 IS zscm 26 1 0 IS cm 46 3 rhizome 0 aboveground shoots 40 canopy covercovar of mainmaln plantpiant 2 2 N 2 0 5.5 im

fig 3 diagram of an isolated plant excavation all aboveground shoots are shown individually and numbered according to height

systems fig 3 this was most apparent with management benefits of vegetative repro- older well established plants from which the duction include 1 an enhanced ability to uti- network originated fig 3 lize unevenly distributed resources and 2 an characteristic of the horizontal rhizome increased competitive ability to occupy adja- expansion were size classes decreasing in con- cent areas harper 1977 cook 1983 in addi- centric circles away from the parent plant tion sprouts are better able to resist invasions was considerable variation rhizome there in ofseedlingsofseedlings from other species while reducing complexity within these individual systems the probability of extinction is accom- excavations established that rhizomesrhizomes can this plished by spreading the risk sprout at least 3 from the parent plant among many in genetically identical individuals therefore a large number of progeny may cook 1983 an evolutionary arise asexually from one individual individ- strategy that employs asexual mechanisms is ual rhizomesrhizomes had from I1 to 52 sprouts consistent with the findings of no evidence indicated that all individual abrahamson 1980 who reported that increased environmental systems were of the same origin that is no severity generally common root connections could be traced shifted emphasis to vegetative reproduction however some might have been connected generally vegetative reproduction is most and later separated after mortality of connec- important where fire weather phenomena tive rhirhizomeszomes and other disturbances are common bostock and benton 1979 went 1979 abrahamson discussion 1980 legere and payette 1981 the sprout- vegetative reproduction is prevalent in ing nature of plains silver sagebrush is likely plains silver sagebrush and the causal agents an adaptation to its northern great plains are of interest and importance to rangeland habitat flooding with associated deposition 206 CclwamboltetalL WAM BOLT ET AL volume 50

along with ice scraping during winter events of plains silver sagebrush reproduction is is common plant production and subse- strongly related to available moisture as in- quently fuel loads for fires are relatively dicated by salisbury 1942 for wild garlic high in bottomlandsbottomlands inhabited by the taxon allium caricarinatumnatum L perhaps this influence consequently fires are common in plains of climate on reproduction might mask differ- silver sagebrush habitats it is logical that ences of sprout to seedling ratios expected plains silver sagebrush is a vigorous sprouter between disturbed and undisturbed sites in in response to the evolutionary influences our study these ratios did not vary signifi- of recurring disturbances this taxon is re- cantly table 1 the reproductive strategies ported to produce only 18 as many achenesacbenes of plains silver sagebrush partially explain the as big sagebrush artemisia tridentata nutt taxon s success and require consideration in harvey 1981 tisdale and hironaka 1981 managing its habitats for optimum balance which likely reflects a reliance on asexual re- between livestock forage production and suit- production able wildlife habitat abundant herbaceous vegetation in mesic flood plains produces substantial competition conclusions for seedlings vegetative sprouts may com- pensate through more rapid morphological we conclude that vegetative reproduc- development because sprouts have the ad- tion in plains silver sagebrush is the pri- vantage of a nutrient reserve from established mary means of plant establishment although plants the sprouting strategy increases sur- sprouting in this taxon has likely evolved with vival abrahamson 1980 habitat disturbances this study did not estab- although not rare in the communities stud- lish that a greater percentage of plants arising ied seedling establishment was found in only from sprouts should be expected on disturbed one third of the plants excavated table 1 than on undisturbed sites however annual this may be attributable to the inconsistency precipitation does appear to be related to the of specific environmental conditions required relative success in initiation and survival of for germination and seedling establishment seedlings and sprouts seedlings apparently environmental factors especially drought require more moisture for both germination might best explain differences in ratios of and survival than do sprouts sprouts and seedlings found in 1983 for example a three year drought at the study area acknowledgments occurred between 1979 and 1981 when the mean annual precipitation was 23023.0 cm and this paper was published with the approval preceded the wet year of 1982 with 41641.6 cm of of the montana agricultural experiment sta- precipitation the longtermlong term average precipi- tion as journal article J 2431 tation is 34834.8 cm this drought coincided with the ages of most plants examined in this study CITED the relatively moist years preceding 1978 literature with 44744.7 cm and following 1982 with 41641.6 abrahamson W G 1980 demography and vegetative cm this drought provided the periods of es- reproduction pages 89 106 in 0 T solbrig ed tablishment for seedlings at the study sites demography and evolution in plant populations blackwell publications as seedlings appear favored scientific oxford however just BEETLE A A 1977 recognition of artemisia subspe- during wet years sprouts were found to have clescieseles a necessity pages 35 42 in K L johnson the advantage in establishing during rela- ed proceedings ath6th annual wyoming shrub tively dry periods the cool wet growing sea- ecology workshop son of 1982 was followed by a warm dry 22322322.3 BOSTOCK S J ANDANDRR A BENTON 1979 the reproductive strategies of five perennial compositae journal of cm growing season in 1983 subsequently ecology 67 91 107 numerous seedlings and few sprouts were COOK R E 1983 clonal plant populations american produced during 1982 and few seedlings with scientist 71 244 253 an abundance of produced FERGUSONFFRGUSON C W 1964 annual rings in big sagebrush sprouts were in press 1983 seedlings from 1982 survived university of arizona tucson 95 appp few HARPER J L 1977 population biology of plants aca- beyond the dry second season therefore it demic press london new york san francisco appears that both the mode and the success 892 appp 1990 SPROUTING AND SEEDLING establishment IN SAGEBRUSH 207

HARVEY S J 1981 life history and reproductive strate- tionaldional biological program vol 1 cambridge uni- gies min artemisiaaftemArtemisiatsiaista unpublished master s thesis versity press cambridge london new york montana state university boemanbozeman melbourne LFGFRELEGERE A AND S PAYETTE 1981 ecology of black SALISBURY E J 1942 the reproductive capacity of plantsofplants spruce picea marianamartanamarlana clonal populations in the studies inm quantitative biology CG bell and sons hemiarctichemiarctic zone northern quebec population ltd london 224 appp dynamics and spatial development arctic and TISDALE E W AND M HIRONAKA 1981 the sagebrush alpine research 13 261 276 grass region a review of the ecological literature MCARTHUR E D AND A P PLUMMER 1978 biogeogra- university of idaho forest wildlife and range phy and management of native western shrubs A experiment station bulletin 209 31 appp case study section tridentataetrtdentataeTridentatae of artemisia WENT F W 1979 germination and seedling behavior of pages 229 243 in K T harper and J L reveal desert plants pages 477 479 in andlandaridlandaddland ecosys- eds proceedings intermountain biogeography tems structure functioning and management symposium great basin naturalist memoirs international biological program vol 1 cam- no 2 brigham young university press provo bridge university press cambridge london utah 268 appp new york melbourne MOTT J J 1979 flowering seed formation and dis- YOUNG J A AND R A EVANS 1972 population dynamics persal pages 627 645 in andlandaddlandaridland ecosystems of green rabbitrabbitbrushbrush proceedings western soci- structure functioning and management internainternal ety weed science 24 13 I1 great basin naturalist 503 1990 appp 209 242 bibliography OF NEVADA AND UTAH vegetation description

2 RP S bourgeron2 L D Engelking J S tuhy andandaandjJ D Brothersonbrotherson44

ABSTRACT listed in alphabetical order by author are 934 references to literature ofthe native vegetation ofnevada and utah this updates and expands the 1967 bibliography of christensen for utah A keyword citation index is provided

this bibliography includes 934 published hencensivehensivesive the only nonbonfieldnonfieldfield studies included and unpublished references to studies of are those explaining nevada utah s vegeta- nevada and utah s native vegetation 1851 tion patterns studies had to be conducted at 1989 vegetation is broadly defined as the least in part in nevada andor utah many aggregate of self reproducing plant species references pertinent to nevada andor utah found in an area or plant communities this but which were completely done in adjacent work updates and expands christensen s states are excluded readers are referred to 1967a 1967b bibliography of the vegetation the relevant bibliographies for these citations of utah papers cited in christensen 1967a the bibliography lists authors alphabeti- 1967b that did not refer directly to the study cally each reference has been assigned of the vegetation are not included here one or more of 28 keywords first when applica- the emphasis is on plant communities 1 I ble it includes a definition of their description species composition phys- a broad vegetation type alpine iognomy and ecology and the techniques desert forest grassland riparian shrubland and concepts used to recognize and analyze wetland woodland using the unesco the communities our definition ofvegetation muellermueiler dombois and ellen- ecology includes spatial and temporal rela- berg 1974 definition the use of riparian is tions of plant communities to the environ- restricted to communities along streams the ment broadly defined to include soil and cli- use of wetland to communities along all other mate and to each other at all levels from the water formations lakes bogs etc the use of blomes to small scale patterns occurring on a these broad terms does not substitute for a given site we omitted references dealing formal classification of the study vegetation solely with the flora of an area floras alone do they are solely provided for easy retrieval of not provide information on the assemblages of the appropriate source of information species and their relationships to the environ- the choice of the other keywords age size ment and to each other structure autecology baseline study bibli- the following subjects are covered ex- ographyography classification community analysis haubauhaustivelystively classification community analysis disturbance diversity patterns early explo- including structure and function distur- ration fire gradient analysis grazing habitat bance diversity pattern fire gradient analy- type inventory map model relict vegeta- sis grazing habitat studies inventory soil tion soil vegetation relationship succession vegetation analysis succession and zonation zonation reflects our own interests in vegeta- selected references to autecology population tion patterns and not necessarily the author s biology competition phenology manage- original emphasis the total number of key- ment and species range are included only words was kept small for easy use of the key when they specifically address some aspect of word citation index finally the keywords the nevada utah vegetation patterns cover- are used consistently for the same object of age of these tangential topics is not compre study throughout the bibliography

address reprint requests to the utah natural heritage program athe2thethe nature conservancy 134 union blvd suite 125 lakewood colorado 80228 atahutahutahvtah natural heritage program 1636 west north temple suite 316 salt lake city utah 84116 4botanybotany and range science department Brigbrighamharn young university provo utah 8460284601

209 210 P S BOURGERON ETALET AL volume 50

acknowledgments 11 ANDALEX RESOURCES 1989 vegetation information volume 1 chapter 3 section G 78319783 19 shirley badame worked on various aspects pages 58 62 in mining and reclamation plan for of the literature search jeff gruenert helped centennial project andalexandalea resources utah division of oil gas and mining number act007ACT 007 with the final search and contributed to the 019 salt lake city utah quality of the bibliography funding was pro- 12 ANDERSONANDEBSON B A AND A H HOLMGBENHOLMGREN 1976 vided by the utah natural heritage program mountain plants of northeasternofnortheastern utah utah agri- and the nature conservancy s rocky moun- cultural experiment station circular 319 logan utah 148 appp tain heritage task force this is a contribu- of the program 13 ANDERSONANDEKSON DAD A J A HENDERSON ANDANDRR L MAUK tion utah natural heritage 1977 indicator species for the habitat types of and RMHTF the authors would appreciate northern utah utah state university depart- being informed of any omissions ment offorestry and outdoor recreation publication 76 appp 14 ANDERSON D A AND E L MINER 1940 some bibliography OF UTAH AND soil plant relationships in the lumperjuniperjumper pinyon belt NEVADA vegetation proceedings of utah academy ofscience arts and letters 17 13 14 1 ALDER CG M 1970 age profiles of aspen forests in 15 ANDERSON D L 1974 ecological aspects ofcercoof cerco utah and northern arizona unpublished thesis carpus montanus raf communities in central university of utah salt lake city 31 appp utah unpublished thesis brigham young uni- 2 ALDON E F ANDANDTT J LORING TECHNICAL COORDI versityversity provoprove utah 84 appp NATORS 1977 ecology uses and management of 16 ANDERSON F K 1987 vegetation volume 1 part pinyon juniper woodlands USDA forest service 7 pages 717 1 7167 16 in mining and reclamation plan general technical report BMRM 39 rocky moun- for banning leadoutloadout facility soldier creek coal tain forest and range experiment station fort company utah division of oil gas and mining collins colorado 32 appp number act007034ACT 007034 salt lake city utah 3 ALEXANDER R R 1985 major habitat types com- 17 ANNABIANNABLE E C R 1985 vegetation and flora of the munity types and plant communities in the rocky funeral mountains death valley national monu- mountains USDA forest service general tech- ment california nevada university of nevada nical repolreportt RM 123 rocky mountain forest and cooperative national park resources studies range experiment station fort collins colo- unit national park serviceuniversityService University of nevada rado 105 appp contribution number 01607 las vegas nevada 4 ALEXANDER R R 1988 forest vegetation on na- 188 appp tional forests in the rocky mountain and inter- 18 ANNFANANNEAN T C 1980 A characterization of yampa mountain regions habitat types and community and green river ecosystems unpublished thesis types USDA forest service general technical utah state university logan 143 appp report RM 162 rocky mountain forest and 19 ARMSTRONG J D 1969 vegetation of the virgin range experiment station fort collins colo- mountains clark county nevada unpublished rado 47 appp thesis university of nevada las vegas 104 appp 5 ALLAN J S 1962 the plant communities oftheodtheofthe big 20 ASHBAUGH J H 1939 A preliminary ecological cottonwood canyon drainage unpublished the- taxonomic survey of southern washoe county sis university of utah salt lake city unpublished thesis university of nevada reno 6 ALLAN J S 1977 the plant communtiescommunitiescomm unties of arches 19 appp national park unpublished dissertation brig- 21 ATWOOD N D 1979 management programs for ham young university provo utah 98 appp plants on federal lands great basin naturalist 7 ALLMAN V P 1952 A preliminary study of the memoirs 3 81 86 vegetation in an exclosure in the chaparral oftheodtheof the 22 AUCLAIR ANA N A BUCHARD AND J pajackowskizajackowskiPAJACKOWSKI wasatch mountains utah unpublished thesis 1976 productivity relations in a carex dominated brigham young university provo utah ecosystem oecologiaoecologia2626 9 31 8 ALLMAN V P 1953 A preliminary study of the 23 AUSTIN D D 1987 plant community changes vegetation in an enclosure in the chaparral of the within a matuiematulemature pinyon juniper woodland great wasatch mountains utah proceedings of utah basin naturalist 47 96 99 academy of science arts and letters 30 637363 73 24 BABCOCK W H 1977 browse evaluation and sur- 9 ALLRED D M D E BECK AND C D JORGENSEN vey techniques for the uinta north slope moose 1963 biotic communities ofthe nevada test site herd unpublished thesis utah state university bnghambrighambangham young university science bulletin bio- logan logical series 23 1 52 25 BAILEY R G ANDANDCC T CUSHWA 1981 ecoregionsEcoregions 10 ANDALEX RESOURCES 1988 vegetation informa- of north america map USUSDIDI fish and wildlife tion volume 1 chapter 3 section G 7831978319783.19783 19 service washington DCD C pages 42 46 in mining and reclamation plan for 26 BAILEY R W C L FORSLING AND R J BECRAFT wildcat leadoutloadout facility andalexandalea resources 1934 floods and accelerated erosion in northern utah division of oil gas and mining number utah USDA miscellaneous publication 196 act007033ACT 007033 salt lake city utah washington DC 21 appp 1990 NEVADA UTAH vegetation bibliography 211

27 BAKERBAKEB F S 1925 aspen in the central rocky 42 BFATHBEATH 0 A 1943 toxic vegetation growing on mountain region USDA department bulletin the salt wash sandstone member of the morri-morn- 12912911 washington DC 47 appp son formation american journal of botany 30 28 BAKER F S C F KORSTIAN ANDNAND N J FETHEBOLFFETHEROLF 698 707 1921 snowshoe rabbits and conifers in the 43 BEATH 0 A C S GILBERT AND H F EPPSON 1940 wasatch mountains of utah ecology 2 304 310 the use of indicator plants in locating seleniferous 29 BAKER W L 1981 tuscaroraTuscaroi a mountains grassland areas in the western united states III111 further unpublished memorandum to dave livermore studies american journal of botany 27 564 573 prepared for the western regional office the 44 BEATH 0 A C S clibertGILBERTcilcligil BERT ANDANDHH F EPPSON 1941 nature conservancy san flanfranciscofraneiscoFianciscoelseo california the use of indicator plants in locating seleniferous 99ppappappp areas in western united states IV progress re- 30 BAMBERG S A A WALLACE E M ROMNEY AND port american journal of botany 28 887 900 R E HUNTER 1980 further attributes of the 45 BEATLEYBFATLEY J C 1965 ecology of the nevada test perennial vegetation in the rock valley area of the site IV effects ofthe sedan detonation on desert northern mojave desert great basin naturalist shrub vegetation in northeastern yucca flat memoirs 4374 37 40 1962 1965 united states atomic energy com- 31 BARKER J R 1983 vegetation information for the mission report UCLA 12157112 1571 university ofcali- deer creek mine volume 12 part 2 pages fornia school of medicine los angeles 55 appp 21012 101 21302 130 in mining and reclamation plan 46 BEATLEYBEATI EY J C 1966a ecological status ofintroduced foiforholbolhorbor deer creek mine utah power and light brome grasses bromus sppapp in desert vegetation company utah division of oil gas and mining ofsouthern nevada ecology 47 548 554 number act015018ACT 015018 salt lake city utah 47 BEATLEY J C 1966b winter annual vegetation 32 BARKER J R 1984 vegetation information foiformol the following a nuclear detonation in the northern des bee dove mines volume 1 part 2 pages mojave desert nevada test site radiation 21022 102 21202 120 in mining and reclamation plan foiroirolfor botany 6 69 82 des bee dove mine utah power and light 48 BEATLEY J C 1974 effects of rainfall and tempera- company utah division of oil gas and mining ture on the distribution and behavior of larrea number act015017ACT 015017 salt lake city utah tntritrldentata creosote bush in the mojave desert of 33 BARKER J R 1987 vegetation information for the nevada ecology 55 245 261 wilberg mine volume 1 paitpaltpart 2 pages 21012 101 2- 49 BEATLEY J C 1975 climates and vegetation pat- 140 in mining and reclamation plan for wilberg tern across the mojavegreatMojave Great basin desert transi- mine utah power and light company utah tion ofsouthern nevada american midland natu- division of oil gas and mining number ACT ralist 93 53 70 015019 salt lake city utah 50 BEATLEY J C 1976 vascular plants of the nevada 34 BARNES W C 1926 the story of the range USDA test site and central southern nevada ecological forest service reprinted in 1926 from part 6 of and geographic distributions technical informa- the hearings before a subcommittee of the com- tion center energy research and development mittee on public lands and surveys U S senate administration TID 2688268811 prepared for the divi- 69th congress ist session 60 appp sion of biomedical and environmental research 35 BARNEY M H ANDANDNN C frishknechtfrischknechtFRISH KNECHT 1974 veg- energy research and development administra- etation changes following fire in the pinyon tion 297 appp juniper types of west central utah journal of 51 bearleyBEATLEYBEAFLEY J C 1979 shrub and tree data for plant range management 279127 91 96 associations across the mojavegreatMojave Great basin desert 36 BARNHURST D ANDANDBB HOSEA 1986 utah big game transition of the nevada test site 1963 1975 range trend studies 1985 utah division of wild- unpublished report prepared for the U S de- life resources publication number 86586 5 salt lake partmentpartment of energy contract EY 76 S 02 2307 city utah 384 appp 52 appp 37 BARTOS D L ANDWAND W F MUEGGLER 1982 early 52 BEATLEY J C 1980 fluctuations and stability in succession following clearclearcuttingcutting ofaspen comcommu-mu climax shrub and woodland vegetation of the mo- nities in northern utah journal ofrange manage- jave great basin and transition deserts of south- ment 35 764 768 ern nevada israel journal of botany 28 149 168 38 BARIOSBARTOS D L F R WARD AND G S INNIS 1983 53 BEAVERBFAVER CRFFKCREEK COAICOAL COMPANY 1983a vegetation aspen succession in the intermountain west a resources volume 2 section 9 pages 919 1 9gigaiga inm deterministicdeter mimsticmimetic model USDA forest service gen- mining and reclamation plan for castle valley eral technical report INT 153 intermountain spur coal processing and leadoutloadout facility forest and range experiment station ogden beaver creek coal company utah division of utah 60 appp oil gas and mining number act007022ACT 007022 39 BATES C G 1924 forest types in the central rocky 54 beaver creek coal company 1983b vegetation mountains as affected by climate and soil USDA resources volume 2 section 9 pages 919 1 9299 29 inm bulletin 1233 washington DCD C 152 appp mining and reclamation plan for huntington 40 BATES J W 1982 desert bighorn sheep habitat canyon no 4 mine beaver creek coal com- utilization in Canyoncanyonlandslands national park utah pany utah division ofoil gas and mining num- unpublished thesis utah state university logan ber act015004ACT 015004 salt lake city utah 41 BEASLEY R S anoANDJAND J 0 klemmedson 1980 eco- 55 BEAVER CREEK COAL COMPANY 1986 vegetation logical relationships ofbristlecone pine amerleanamerican resources volume 2 section 9 pages 919 1 9609 60 in midland naturalist 104 242 252 mining and reclamation plan for gordon creek 3 212 P S BOURGERON ETETALAL volume 50

and 6 mines beaver creek coal company utah 71 BILLINGS W D 1950 vegetation and plant growth division of oil gas and mining number ACT as affected by chemically altered rocks in the west- 007017 salt lake city utah ern great basin ecology 31 62 74 56 BEAVERBEAVFR CRFEKCREEK COAL COMPANY 1988 vegetation 72 BILLINGS W D 1951 vegetational zonation in the resources volume 3 chapter 9 sections 919 1 939.39 3 great basin of western north america pages pages 919 1 9369 36 in mining and reclamation plan 101 122 in the ecological foundations of the re- for trail mountain no 9 mine beaver creek coal generation ofvegetationof vegetation in andaridarld zones UNESCO company utah division of oil gas and mining series B colloquescollo ques 9 number act015009ACT 015009 salt lake city utah 73 BILLINGS W D 1954 temperature inversions in 57 BEAVER CREEK COAL COMPANY 1989 vegetation the pinyon juniper zone of a nevada mountain resources volume 3 section 9 pages 919 1 9399 39 in range butler university botanical studies 12 plan mining and reclamation for gordon creek 2 112 118 company and 7 mines beaver creek coal utah 74 BILLINGS W D 1978 alpine phytogeography gas ACT division of oil and mining number across the great basin great basin naturalist 007016 salt lake city utah memoirs 2 105 117 58 BECK D E D M ALLRED J R MURDOCK C D 75 BILLINGS W D H K HUMPHREYS AND J B DAR- JORGENSON C L HAYWARD ANDANDWW W TANNER LING 1954 environmental studies in the cold 1964 nevada site desert ecology proceed- test deserts and deserts of the western great ofutahof utah academy ofscienceof science arts and semi ings letters basin of unpublished report 41 202 210 north america by 44 109 GM 59 BECKWITH LICUTLIEUT E G 1854 report of explo- duke university contract DA 1261 for U S army 112 rations for a route for the pacific railroad on the prepared the US appp line of the forty first parallel of north latitude in 76 BISHOP A B AND D B PORCELLA 1980 physical reports of explorations and surveys to ascertain and ecological aspects oftheofodthethe upper colorado river the most practicable and economical route for a basin pages 17 56 in W spofford A parker and railroad from the mississippi river to the pacific A kneese eds energy development in the ocean vol II11 house of representatives execu- southwest problems of water fish and wildlife tive document 91 33rd congress and2nd session in the upper colorado basin resources for the 132 appp future washington DCD C 60 BEESON C D 1974 the distribution and synecol- 77 bjerregaard R S N E WEST M M CALDWELL ogy of great basin pinyonpryonpmyon juniperjumper unpublished AND H F MAYLAND 1984 standing crops and thesis university of nevada reno 91 appp dynamics of phyphytomasstomass and minerals in two salt 61 BEETLEBELJTLE A A 1950 A study ofsagebrushofsagebrush the section desert shrub communities great basin naturalist tndentataetridentataeTridentatae ofartemisia university of wyoming 44327 337 agricultural experiment station bulletin 368 78 BLACKBURN W H 1967 plant succession on se- laramie wyoming 83 appp lected habitat types in nevada unpublished the- 62 BELLBLLL K L ANDANDRR E JOHNSON 1980 alpine flora of sis university of nevada reno 162162ppappp the bassukwassuk range mineral county nevada 79 BLACKBURN W H R E ECKERT JR AND P T Madromadronofioflo 27 25 35 TUELLER 1969 vegetation and soils of the crane 63 BELL K L AND L C BLISS 1979 autecology of springs watershed nevada agricultural experi- kobresiaKobresia bellardbeliardbellardibellardiibellardnbellardnii why winter snow accumulation ment station bulletin R 55 reno nevada 63 appp limits local distribution ecological monographs 80 BLACKBURN W H ANDPAND P T TUELLER 1970 pinyon 49 377402377 402 and juniper invasion in black sagebrush communi- 64 BENSON S B 1935 A biological reconnaissance of ties in east central nevada ecology 51 841 848 navajo mountain utah university of california 81 BLACKBURN W H P T TUELLER AND R E publications in zoology 40 439 456 ECKERT JR 1968a vegetation and soils oftheodtheof the mill 65 BERMANT D J 1982 trend and condition of major creek watershed nevada agricultural experi- range types oftheodtheofthe uinta national forest utah as ment station bulletin R 43 reno nevada 69 appp determined by quantitative analysis of protoprotoplotplot 82 BLACKBURN W H P T TUELLER AND R E transects unpublished thesis utah state univer- ECKERT JR 1968b vegetation and soils of the sity logan crowley creek watershed nevada agricultural 66 BESSEY G E 1960 aquatic plants of central utah experiment station bulletin R 42 reno nevada and their distribution unpublished thesis brig- 60goppappp ham young university provo utah 83 BLACKBURN W H P T TUELLER AND R E 67 betancourt J L 1984 late quaternary plant ECKERT JR 1968c vegetation and soils of the zonation and climate in southeastern utah great buckwaterduckwaterDuckwater watershed nevada agricultural basin naturalist 44 1 35 experiment station bulletin R 40 reno nevada 68 BIDDULPHBIDDULFH 0 1933 the anatomy of the transition 76 appp region ofofpinuspinus adulisedulis unpublished thesis brig- 84 BLACKBURN W H P T TUELLER AND R E ham young university provo utah ECKERT JR 1969a vegetation and soils oftheofodthethe cow 69 BILLINGS W D 1945 the plant associations of creek watershed nevada agricultural experi- the carson desert region western nevada but- ment station bulletin R 49 reno nevada 80 appp ler university botanical studies 7 89 123 85 BLACKBURN W H P T TUELLER AND R E 70 BILLINGS W D 1949 the shadscaleshadscale vegetation ECKERT JR 1969b vegetation and soils of the zone of nevada and eastern california in relation coils creek watershed nevada agricultural ex- to climate and soils american midland naturalist periperimentment station bulletin R 48 reno nevada 42 87 109 81 appp 1990 NEVADA UTAH vegetation bibliography 213

86 BLACKBURN W H P T TUELLER AND R E 100 BOWNS J E AND N E WEST 1976 blackbush ECKERT JR 1969 vegetation and soils of the coleognyneramosissimacoleognyne ramosissima torr on southwestern churchill canyon watershed nevada agricul- utah rangelands utah agricultural experiment tural experiment station bulletin R 45 reno station research report 27 logan utah 27 appp nevada 157pp157 appp 101 BRACKEN A F 1940 dry farming as developed 87 BLACKBURN W H P T TUELLERTUELLEK AND R E in the sagebrush zone proceedings of utah ECKERT JR 1969d vegetation and soils oftheodtheofthe pine academy of science arts and letters 17 25 32 and mathew canyon watersheds nevada agricul- 102 BRADLEYWGBRADLEY W G 1964 the vegetation of the desert tural experiment station bulletin R 46 reno game range with special reference to the desert nevada illliiI1 I1 I1 appp bighorn pages 43 67 in transactions of the 88 BLACKBURN W H P T TUELLERTUELLEB AND R E& desert bighorn council las vegas nevada ECKERT JR 1971 vegetation and soils of the 103 BRADLEY W G 1965 A study ofblackbrushofblackbrushblaekblackbrush com- rock springs watershed nevada agricultural munity on the desert game range pages 56 61 experiment station bulletin R 83 reno nevada in transactions of the desert bighorn council 116ppilg116 appp ath9th annual meeting 6 8 april 1965 redlands 89 BLACKHAWK COAL COMPANY 1981 vegetation re- california sources chapter 9 section 929.2 pages 919 1 9279 27 in 104 BRADLEY W G 1966 populations of two sonoran mining and reclamation plan for willow creek desert plants and deductions as to factors limiting mine blackhawk coal company utah division their northward extension southwestern natural- of oil gas and mining number act007002ACT 007002 ist 11 395 401 salt lake city utah 105 BRADLEYBBADLEY W CG 1967 A geographical analysis oftheofodthethe 90 BLAISDELL J P AND R C HOLMGREN 1984 flora of clark county nevada journal of the ari- managing intermountain rangelands salt desert zona academy ofscience 4 151 162 shrub ranges USDA forest service general 106 BRADLEYBBADLEY W G 1973 ecological study ofzion nat- technical report INT 163 intermountain forest ional park unpublished report prepared for the and range experiment station ogden utah USDI national park service Springdale utah 91 BLAISDELL J P R B MURRAY AND E D 183 appp 1982 mcarthur managing intermountain range- 107 BRADLEY W GGANDJAND J E DEACON 1965 the biotic lands sagebrush grass ranges USDA forest ser- communities of southern nevada university of vice general technical report INT 134 inter- nevada prepublication series 9 desert research mountain forest and range experiment station institute las vegas nevada 86 appp ogden 41 pp utah ap 108 bradley W G andaandjand J E deacon 1967 the biotic 92 BLAKSLEY J A 1987 avian habitat relationships communities of southern nevada nevada state in riparian zones of northern utah unpublished museum anthropological papers 13 part IV car- thesis university ofidaho moscow son city nevada 69 appp 93 BLANK D L 1984 forage quality comparison of 109 BRANSON F A 1966 geographic distribution and burned and unburned aspen communities un- factors affecting the distribution of salt desert published thesis utah state university logan shrubs in the united states pages 13 43 in pro- 94 BLEAK A T N C frischknecht A P PLUMMER ceceedings from the salt desert shrub symposium AND R E ECKERT JR 1965 problems in artificial 1 4 august 1966 USDI bureau eflandoflandofland manage- and natural revegetation ofthe andaridarld shadscaleshadscale veg- ment cedar city utah etation zone ofutahofutah and nevada journal ofrangeoforrangerange 110 BRANSON F A 1985 vegetation changes on west- management 185918 59 65 ern rangelands range monograph 2 society for 95 BOLEN E G 1962 plant ecology of spring fed salt range management denver colorado 76 appp marshes unpublished thesis utah state univer- 111 BRANSON F AARR F MILLER AND I1 S mcqueen sity logan 1967 geographic distribution and factors affect- 96 BOLEN E G 1964 plant ecology of spring fed salt ing the distribution of salt desert shrubs in the marshes in western utah ecological monographs united states journal of range management 29 3414334.143 166 287 296 97 BOSTICK V B W E NILES W A MCCLELLAN 112 breternitzbreternitzdD AAANDDAND D W MARTIN 1973 report E H OAKESANDJOAKES AND J R WILBANKS 1975 inventory of dolores river project reconnaissance 1972 of natural landmarks of the great basin vols I1 1973 unpublished report submitted to the USDI and II11 unpublished report prepared for the national park service midwest archaeological USDI national park service by university of center lincoln nebraska nevada las vegas 690 appp 113 BRIGGS G M 1978 A study of sedge dominated 98 BOUCEKMBOUCEK M M 1986 vegetation survey at the sum- areas in the uinta mountains unpublished thesis mit no 1 coal mine summit county utah vol- utah state university logan 89 appp ume 1 section 78319783.19 appendix 78319783.19 pages 114 BRIGGS G M AND J A MACMAHON 1982 struc- 1 15 in mining and reclamation plan for summit ture ofalpineofalpine plant communities near king s peak no I1 mine summit minerals incoroporated uinta mountains utah great basin naturalist 42 utah division of oil gas and mining number 50 59 act043001ACT 043001 salt lake city utah 115 BRIGGS G MANDJM AND J A MACMAHON 1983 alpine 99 BOWERS J E 1982 the plant ecology of inland and subalpine wetland plant communities of the dunes in western north america journal ofarid uinta mountains utah great basin naturalist 43 environments 5 199 220 523 530 214 P S BOURGERON ETALET AL volume 50

116 BRbritionbritronBRITTONBRITiTrONronTON CMC M ANDANDMM H RALPHS 1979 use of fire 132 brotherson J D ANDANDSS T OSAYANDE 1980 min- as a management tool in sagebrush ecosystesm eral concentrations inm true mountain mahogany pages 101 109 in the sagebrush ecosystem A and utah juniper and in associated soils journal of symposium april 1978 utah state university range management 33 182 185 logan 133 brotherson J D ANDANDKK P PRICE 1984 naturali- 117 brothersonBROThersonHEBSON J D 1974 A vegetation analysis of zation and habitat relationships of bitter night- areas involved in the proposed four seasons pro- shade solanumsolarium dulcamara inm central utah ject pages 60 104 in environmental report on great basin naturalist 44 317 323 proposed four seasons project unpublished 134 brotherson J D K P PRICE AND L 0 orourkeROURKE report prepared by brigham young university 1987 age in relationship to stem circumference center for health and environmental studies and diameter in cliffrose cowaniacoramaCocowamawania mexicansmexicanamexicana var for four seasons inc stansburstansburianastansbunanastansbunanaianalana in central utah great basin naturalist 47 118 brothersonBROThersonHEBSON J D 1978 vegetative and wildlife 334 338 assessment of the jordan aqueduct extension 135 brotherson J D L L RASMUSSEN AND R D bonneville unit central utah project unpub- BLACK 1986 comparative habitat and community lished report prepared for the rocky mountain relationships of atnplexatriplex conferticonferttfohaconfertifoliafolia and sarco- research co and USDI bureau of reclamation batus vermiculatus in central utah great basin naturalist 46 348 357 provo utah 45 appp 136 brotherson J D S R AND J R 119 brotbrothersonBROI hersonHEBSON J D 1981 aquatic and semiaquatic rushforth JOHANSEN 1983 effects of longtermlong term vegetation of utah lake and its bays great basin grazing on cryptogam crust cover navajo monu- naturalist memoirs 5 68 84 in national ment arizona journal of range management 120 D 1987 plant 36 brotherson J community zonation 579 581 in response to soil gradients in a saline meadow in 137 brotherson J D AND V WINKEL 1986 habitat near utah lake utah county utah gigreateat basin relationships of saltcedarsaltcedar tamarix ramosts naturalist 47 322 333 ramosiskamosis sima in central utah great basin naturalist 46 121 brotherson J D D L ANDERSONANDEBSON AND L A 535 541 SZYSKA 1984 cercocarpus habitat relations of Cercocarpus 138 BROWN D E C H LOWFLOWE AND C P PASE 1979 montanus true mountain mahogany in central A digitized classification system for the range biotic com- utah journal of management 3732137 321 324 munimunitiesties of north america with community se- 122 brotherson J D AND S J BARNESBARNFS 1984 habitat ries and association examples for the southwest relationships of clauxglaux marimartmaritimemaritimamaritinwmaritimafititimafimanw in central utah journal of the arizona nevada academy of sci- great basin naturalist 44 299 309 ence 14114 1 16 123 brotherson J D AND K J brothersonBRO IHERSON 1979 139 BROWN D E AND C H LOWE 1980 biotic com- ecological and community relationships of eri- munimunitiesties of the southwest USDA forest service ogonum corymbosumcorymbosum polygonaceae in the uinta general technical report RM 78 rocky moun- basin utah great basin naturalist 39 177 191 tain forest and range experiment station fort 124 brotherson J D AND W T brotherson 1981 collins colorado grazing impacts on the sagebrush communities of 140 BROWN L L 1962 establishment of forage plants central utah greatgi eat basin naturalist 41 335 340 on difficult salt desert range sites unpublished 125 brotherson J D J G CARMAN ANDANDLL A SZYSKA thesis university of nevada reno 1984 stem diameter age relationships of tamarix 141 BRUN J M 1962 A comparison of the line inter ramosissimaramosisstma in central utah journal of range ceptioncaptionception point frame and distance measurement management 37 362 364 methods for analyzing desert shrub vegetation 126 brotherson J D J N DAVIES AND L GREEN unpublished thesis utah state university lo- WOOD 1980 diameter age relationships of two gan species of mountain mahogany journal of range 142 BRUNERBBUNER A D 1979 predicting success of pre- management 33 367 370 scribed fires in pinyonpryonpmyon juniperjumper woodland in ne- 127 brotherson J D ANDWAND W E EVENSON 1983 veg- vada USDA forest service research paper INT etation communities surrounding utah lake and 219 intermountain forest and range experiment its bays utah lake vegetation studies unpub- station ogden utah 11 appp lished report prepared for the utah division of 143 BUCHANAN B A K T HARPER AND N C wildlife resources and USDI bureau of recla- frischknecht 1978 allelopathicAllelopathic effects of bur mation provo utah 401 appp buttercup tissue on the germination and growth of 128 brotherson J D AND D FIELD 1987 tamarixTamatixmarix various grasses and forbs in vitro and in soil great impacts of a successful weed rangelands 9 basin naturalist 38 90 96 110 112 144 BUCHANAN H 1960 the plant ecology of bryce 129 brotherson J D ANDANDWW J MASSLICH 1985 veg- canyon national park utah unpublished disser- etation patterns in relation to slope position in the tation university of utah salt lake city castle cliffs area of southern utah great basin 145 BUCHANAN H AND K T HARPER 1981 forests of naturalist 45 535 541 bryce canyon national park consequences ofpast 130 brotherson J D ANDANDCC MORDEN 1979 vegeta- management policies and recommendations for tion of utah lake shoreline and bays map in four the future unpublished final report part III111 pro- paipalpartsts press publishing limited provo utah ject CX 1200 8 boigbolg typescript 131 brothersonjbrothersonbrothertonj J D G NEBEKFR M SKOUGARD AND 146 BUDY J D andlAND J A YOUNG 1979 historical use of J FAIRCHILD 1978 plants of navajo national nevada s pinyon juniper woodlands journal of monument great basin naturalist 33 19 30 forest history 23 112 121 1990 NEVADA UTAH vegetation bibliography 215

147 BUNDERSONBUNDEBSON E D D J WEBER AND D L NELSON kane county utah unpublished thesis brigham 1986 diseases associated with juniperusJumperus osteoosoteo young university provo utah sperma and a model for predicting their occur- 163 CEDAR CREEKCBEEK associates incorporated 1987 rence with environmental site factors great basin draft vegetation information for the alton coal naturalist 46 427 440 mining project volume 6 chapter 3 appendix 148 BURKE M H 1934 plant distribution studies in the 363 6 B pages 1 41 in mining and reclamation plan wellsville range utah unpublished thesis utah foiformorpor alton mine utah international incorporated state agricultural college logan 33 appp utah division of oil gas and mining number 149 BURR G 0 1931 native vegetation of the pre- act025003ACT 025003 salt lake city utah historic lake bonneville basin journal of the 164 CHABOT B F AND W D BILLINGSBILLINCS 1972 origins american society Agronomyofagronomyof Agi onomy 23 407 413 and ecology ofthe sierran alpine flora and vegeta- 150 BUTLER J R ANDJAND J L ENGLAND 1979 vegetation tion ecological monographs 42 163 199 map of the southeastern uinta basin utah and 165 chamberlain R V 1911 the ethnobotany of the colorado USDI geological survey miscellaneous gosiutegosmte indians proceedings of the academy of investigative series 1141111411 washington DCD C science of philadelphia 63 249924 99 151 BUZZARD R F JR 1981 comparisonCompanson of two tradi- 166 CHAMBERS J C 1979 the effects of grazing on salt tional techniques for determining range trend and desert shrub species survival during a period of recomendationsrecommendations for improved procedures un- below average precipitation unpublished thesis published thesis utah state university logan utah state university logan 152 BYE R A 1972 ethnobotany oftheodtheofthe southern paiute 167 CHAPMAN V J 1960 salt marshes and salt deserts indians min the 1870 s in great basin cultural of the world plant science monograph inter- ecology a symposium reno nevada research science publishers inc new york 392 appp publications in the social sciences 8 168 CHOATE G A 1965 forests in utah USDAUS DA forest 153 CALDWELL M M ANDANDNN E WEST 1972 the plant service resource bulletin INT 4 intermountain ecology of utah s desert rangelands utah science forest and range experiment station ogden 331433 14 15 utah 61 appp 154 call C A and J R barker 1980 vegetation sur- 169 christensen E M 1949 the ecology and geo- vey of USU S fuel company property hiawatha graphic distribution of oakbrushoakbrush quercus gam utah volume 3 chapter 9 section 999.99gg 9 appendix beill in utah unpublished thesis university of IXIIX I1 pages 1 80 in mining and reclamation plan utah salt lake city 70 appp for hiawatha mines complex USU S fuel com- 170 christensen E M 1950 distributional observa- gas pany utah division ofrilof0ilof oil and mining num- tions of oakbrushoakbrush quercus gamberigambehi nutt in ber act00701act007011ACT 007010070111 salt lake city utah utah proceedings of utah academy of science 155 CALLISON J JR ANDANDJJ D brotherson 1985 habi- arts and letters 27 22 25 tat relationships of the blackbush community 171 christensen E M 1955 ecological notes on the coleogyne ramosissiramosissimaina of southwestern utah mountain brushblush in utah proceedings of utah great basin naturalist 45 321 326 academy of science arts and letters 32 156 CALLISON J JR J D brotherson AND J E 107111107 iiiililil111 BOWNS 1985 the effects affireoffireoffirehirebire on the blackbrushblackbrush 172 christensen E M 1957 photographic history of coleogyne ramosissimaramostssima community of south- the mountain brush on Y mountain central western utah journal of range management 38 utah proceedings of utah academy of science 535 538 arts and letters 34 154 155 abstract 157 CAWBFLLCAMPBELL V 0 1977 certain edaphic and biotic 173 christensen E M 1958 growth rates and vege- factors affecting vegetation in the shadscaleshadscale tation changes in the oak maple brush in lower community of the kaiparowits areaai ea unpublished provo canyon utah proceedings of utah thesis brigham young university provo utah academy of science arts and letters 35 59 appp 167 168 158 CANNON H L 1960 the development of botanical 174 christensen E M 1959a A climatic study ofsomeorsomeofsome methods of prospecting for uranium on the colo- northern desert shrub communities proceedings rado plateau USDI geological survey bulletin of utah academy of science arts and letters 36 1085 A washington DC 50 appp 174 abstract 159 CANON S K 1985 habitat selection foraging 175 christensen E M 1959b A comparative study of behavior and dietary nutrition of elk in burned the climates of mountain brush pinyon juniperjumper vs unburned aspen forests unpublished thesis and sagebrush communities in utah proceedings utah state university logan of utah academy of science arts and letters 36 160 CARMAN J G ANDJAND J D brotherson 1982 com- 174 175 abstract parisonsparisons of sites infested and not infested with 176 christensen E M 1961a the agropyron poa salt cedar tamarix penpentandrapentandriatandra and russian bunchbunchgrassgrass vegetation in central utah proceed- olive elaeagnus angustifoliaangusttfolia weed science 30 ings of utah academy of science arts and letters 360 364 38 113 abstract 161 CARTER K S& 1981 analysis of trend and methods 177 christensen E M 1961b the deciduous tree used to determine trend on southern utah nat- communities ofcentral utah bulletin of the eco- ional forest unpublished thesis utah state logical society ofamericaofamerica 42 62 63 abstract university logan 178 christensen E M 1962 the rate ofnaturalizationofnaturalization 162 CASTLE E &S 1955 the vegetation and its relation- tainarixoftmnanxof Tainarix min utah americal midland naturalist ship to the dune soils of the pink sand dunes of 685157685168 515751 57 216 P S BOURGERON ETETALAL volume 50

179 christensen E M 1963a1963d naturalization of rus- wildlife resources publication 67167 1 salt lake sianslan olive elaeagnus angustifoliaangustifolta L in utah city utah 171 appp american midland naturalist 70 133137133 137 196 coicolCOLESES F H AND J C PEDERSONPEDEBSON 1968 utah big 180 christensen E M 1963b the foothill bunchbunchgrassgrass game range inventory 1967 utah division of vegetation of centralofcentral utah ecology 44 156 158 wildlife resources publication 68268 2 salt lake 181 christensen E M 1964a changes in composition city utah 120 appp of a bromus tectoriumtectorumtectorum sporobolus cryptcryptandrusandrus 197 COLES F H AND J C PEDERSONPEDEBSON 1969 utah big pro- aristida longisetalongiseta community following fire game range inventory 1968 utah division ofot ceceedingceedingsedingedings of utah academy of science arts and wildlife resources publication 69269 2 salt lake letters 41 53 57 city utah 164 appp 182 christensenchbistensfn E M 1964b succession in a moun- 198 COLES F H ANDJAND J C PEDERSONPFDERSON 1970 big proceed- utah tain brush community in central utah game range inventory 1969 utah division of ings of utah academy ofscience arts and letters wildlife resources publication 70170 1 salt lake 41 10 13 erratum 412 ii11 city utah 127 appp 183 christensen E A 1967a bibliography of utah 199 COLLINS P D 1980 comparative life history and wildwindlandland brigham young botany and wildland conservation floral characteristics of desert and montane plant university science bulletin biological series communities inm utah unpublished thesis brig- 91 provo 135 appp 91 utah ham young university provo utah 163 appp 184 E M 1967b bibliography of utah christensen 200 COLLINS P D 1983 vegetation and reclamation of botany and wildlandwildwindlandland no II11 pro- conservation the wellington coal cleaning plant volume 3 ceceedings of utah academy of science arts and appendix H pages 1 44 in mining and reclama- 44 545 566 in letters tion plan for wellington prep plant kaiser coal 185 E M AND D 1972 christensen J brotherson corporation utah division of oil gas and min- preliminary botanical survey of the bonneville ing number act007012ACT 007012 salt lake city utah unit of the central utah project pages 5 134 in 201 COLLINS P D AND K T HARPER 1980 E M christensen et al eds preliminary survey vegetation map of the mt complex and san pitch of the biota of the bonneville of the central nebo unit mountains brigham young utah project unpublished report prepared for utah university press provoprove utah the USDI bureau of reclamation provo utah by brigham young university center for health 202 colllnsCOLLINScoicol LINS P D K T HARPER AND B K PENDLETONPCNDLETON and environmental studies provo utah 1983 comparative life history and floral charac- teristicste of desert and mountain floras in utah 186 christensen E M AND B F HARRISON 1961 in great basin naturalist 43 385 393 ecological study area at lily lake in the uinta mountains utah proceedings of utah academy 203 COLLINS W B 1977 diet composition and activi- ofscience arts and letters 38 36 49 ties of elk on different habitat segments in the lodgepole 187 christensen E M anumANDMAND M A hutchinson 1965 pine type uinta mountains unpub- historical observations of the ecology of rush lished thesis utah state university logan and tooelethoele valleys utah proceedings of utah 204 COLTHARP G B AND N E WEST 1966 effects of academy of science arts and letters 42 90 105 surface soil treatments on soil water and vegeta- pages 188 christensen E M AND H B JOHNSON 1964 pre- tion in utah s east desert area 88 97 in settlement vegetation and vegetational change in proceedings fromhiom salt desert shrub symposium three valleys in central utah brigham young uni- 1 4 august 1966 USDI bureau eflandoflandof land manage- versity science bulletin biological series 44 ment cedar city utah I1 16 205 COMMITTEE ON environmentalENVIRONMENTAI QUALITY 1970 189 christensen E M AND S L welghWELSHWFLSH 1963 pre- preserving nevada s environmental heritage un- settlement vegetation of the valleys of western published final report prepared for the governor s summit and wasatch counties utah proceed- natural resources council by the department ings of utah academy of science arts and letters of conservation and natural resources carson 4016340 163 174 city nevada 190 CHRISTIAN R 1962 plant geography of the beaver 206 COOK C W 1977 effects of season and intensity dam mountains unpublished thesis university of use on desert vegetation utah agricultural of utah salt lake city 153 appp experiment station bulletin 483 logan utah 191 clCLEMENTEMENT R C 1979 culture and species endan- 57 appp germent great basin naturalist memoirs 3 207 COOK C W AND L E HARRIS 1950 the nutritive II11 16 content ofthe grazing sheep s diet on the summer 192 CLOKEY I1 W 1951 flora of the charleston moun- and winter ranges of utah utah agricultural tains clark county nevada university of cali- experiment station bulletin 342 logan utah fornia press berkeley 66 appp 193 CLOVER E U anulANDLAND L JOTTER 1944 floristicfionFlonsticstie studies 208 COOK C W AND R HURST 1962 A quantitative in the canyon of the colorado and tributariestributaries measure ofofplantplantpiant association on ranges in good and american midland naturalist 32 591 642 poor condition journal ofrange management 15 194 COLE N J 1968 mule deer utilization of rehabili- 266 273 tated nevada rangelands unpublished thesis 209 COOK C W ANDANDCC E LEWIS 1963 competition university of nevada reno between big sagebrush and seeded grasses on 195 COLES F H AND J C PEDERSON 1967 utah big foothill ranges in utah journal of range manage- game range inventory 1966 utah divisondavison of ment 1624516 245243 250 1990 NEVADA UTAH vegetation bibliography 217

210 coCOOPOP MINING COMPANY 1988 vegetation vol- 227 COYNE P I1 AND C W COOK 1970 seasonal carbo- ume 2 chapter 9 sections 919.19gi 1 939 3 pages 919 1 hydrate reserve cycles in eight desert range spe- gag 9109 10 plus appendix 9 A pages gai 1 9 in min- cieseles journal of range management 13 438 444 and plan for canyon ing reclamation bear mine 228 critchfield W B AND G L allenbaugh 1969 coopco op mining company utah division of oil gas the distribution of pinaceae in and near northern and mining number act015025ACT 015025 salt lake nevada Madromadronomadroniofionionno 19 12 27 city utah 229 CROFT A R L WOODWARD AND D A ANDERSON 211 COOPco OP MINING COMPANY 1989 vegetation vol- 1943 measurement of accelerated erosion on ume 2 chapter 9 sections 919.19gi 1 939.39 3 pages 919 1 range watershed land journal of 4 9109 10 plus appendix 9 A pages gai 1 gag9 min- forestry in 112 116 ing and reclamation plan for trail canyon mine coopco op mining company utah divisondavison of oil 230 CROWTHER E G AND K T HARPER 1965 vegeta- gas and mining number act015021ACT 015021 salt lake tional and edaphic characteristics associated with city utah aspen strips in big cottonwood canyon pro- 212 COOPER S V P T MUELLER AND R B MURRAY ceedingsce of utah academy of science arts and 1978 habitat type classification ofpinyon junjuniperiperi letters 42 222 230 woodlands in the great basin unpublished re- 231 CURREY D R 1965 an ancient bristlecone pine port prepared for the agricultural experiment stand in eastern nevada ecology 46 564 566 station reno nevada 60 appp 232 DAINES L L 1917 on the flora of great salt lake 213 COTTAM W P 1926 an ecological study of the flora american naturalist 51 499 506 of utah lake utah unpublished dissertation 233 DASTRUP B C 1963 vegetational changes of the university of chicago 46 appp uinta basin since settlement unpublished thesis 214 COTTAM W P 1929a man as a biotic factor illus- brigham young university provo utah 118 appp trated by recent flonfloristicsticstie and physiographic 234 DAVIS J B 1971 estimating plant production from changes at the meadows washington mountain soil and flonfloristicstiestic correlates in hot creek valley county utah ecology 10 361 364 nevada unpublished thesis university of ne- 215 commCOTTAM W P 1929b some phytogeographical vada reno features of utah proceedings of utah academy 235 DAVIS J N 1976 ecological investigations in cer of science arts and 6 6 7 in letters cocarpus ledifoliusledifolius nutt of 216 COTTAM W P 1930 some unusual flonfloristic features communities utah sticstie unpublished thesis brigham young of the uintah mountains of university utah proceedings provo utah utah academy of science arts and letters 7 48 49 236 DEBYLE N V 1973 establishment report for mor- 217 COTTAM W P 1937 has utah lost claim to the ris creek research natural area within the davis lower sonoran zone science 85 563 564 county experimental watershed unpublished 218 COTTAM W P 1947 Is utah sahara bound bul- report prepared for USDA national forest ser- letin ofthe university ofutah 3711 vicevlee washington DCD C 7 appp 237 DEAN K C HAVENS 219 commCOTTAM W P 1954 prevernal leafing of aspen in R AND K T HARPER 1969 utah mountains journal oftheodtheof the arnold arboretum chemical and vegetative stabilization of a nevada 3523935 239 250 copper porphyry mill tailing USDI bureau of 220 COTTAM W P 1961 our renewable wild lands a mines report of investigations 7261 salt lake challenge university of utah press salt lake city utah 14 appp city 182 appp 238 DEAVER C F AND H S HASKELL 1955 pinyon 221 COTTAM W P 1976 the impact ofofmanman on the flora resources distribution ofpmyonof pinyon pinus ededuiseduliseduhsulis of the bonneville basin university of utah re- yield and resin potentialities navajo hopi reser- search paper 76176 1 salt lake city utah vationsvations arizona utah university of arizona press 222 COTTAM W P AND F R EVANS 1945 A compara- tucson 37 appp tive study oftheodtheofthe vegetation ofgrazedofgrazed and ungrazed 239 dellenbaugh F S 192619081926 1908 A canyon voyage canyons of the wasatch range utah ecology 26 the narrative of the second powell expedition 171 181 down the green colorado river from wyoming 223 COTTAM W P AND G STEWART 1938 range con- and the expeditions on land in the years 1871 and servation in wasatch county utah unpublished 1872 G P putnam s sons new york 277 appp cooperative western range survey report prepared 240 DESCHAMP J A P J URNESS AND D D AUSTIN for USDA forest service intermountain forest 1979 summer diets of mule deer from lodgepole and range experiment station ogden utah pine habitats journal of wildlifeofwildlife management 43 224 COTTAM W P ANDANDGG STEWART 1940 plant succes- 154 161 sion as a result of grazing and of meadow desicca- 241 DICE L R 1939 the sonoran biotic province tion by erosion since settlement in 1862 journal of ecology 20 118 129 forestry 38 613 626 242 DINA S J 1970 an evaluation of physiological re- 225 COTTAM W P J M TUCKER AND R DROBNICK sponse to water stress as a factor influencing the 1959 some clues to great basin postpostpluvialpluvial cli- distribution of six woody species in red butte mates provided by oak distribution ecology 40 canyon utah unpublished dissertation univer- 361 377 sity of utah salt lake city 117 appp 226 COVILLE F V 1892 descriptions of new plants 243 DINA S J L G KLIKOFF AND M B keddington from southern california nevada utah and ari- 1973 seasonal water potential patterns in the zona proceedings of the biological society of mountain brush zone utah american midland washington 7 65 80 naturalist 89 234 239 218 P S bourgebonetalbourgeronBouRceronGERON ET AL volume 50

244 dittberner PPLL AND M R OLSON 1983 the 258 EASTMOND R J 1968 vegetative changes in a plant information network PIN data base colo mountain brush community through eighteen radolado montana north dakota utah and years unpublished thesis brigham young uni- wyoming USDI fish and wildlife service FWS versity provo utah 64 appp OBS 8336 westelwesternn energy and land use team 259 EASTWOOD A 1896 report on a collection ofofplantsplants fort collins colorado 786 appp from san juan county in southeastern utah pro- 245 DIXON H 1935 ecological studies on the high ceceedings of the california academy of science plateaus of utah botanical gazette 97 272 320 series 2 6 271 328 246 DOUGLAS C L 1984 management significance of 260 EATON F D 1971 soil moisture depletion actual the flora and physiognomy of the cottonwood and potential evapotranspiration in an engelmann mountains death valley national monument spruce subalpine fir forest unpublished thesis california university of nevada cooperative na- utah state university logan utah tional park resources studies unit report 02207 261 ECKERTECKEBT R EK 1954 A study of competition be las vegas nevada twedentweeen whitesagewhitwhitecageesage and halogeton in nevada un- 247 DOUGLAS C L 1985 management significance of published thesis university of nevada reno vegetation and flora of the funeral mountains 262 ECKERTECKERF R E A T BLEAK J 11 ROBERTSON AND death valley national monument californianscaliforniane E A NAPHAN 1961 responses of agropyron vada university of nevada cooperative national statumcricrlcncristatumcnstatum A desertorumdesertorum and other range grasses park resources studies unit report 01608 las to different sites in eastern nevada ecology 42 vegas nevada 775 783 248 DOUGLAS C L 1986a management significance of 263 el ghonemyghonhmy A A 1980 socioecologicalSocioecological and soil plant succession after fire hunterhuntel mountain plant studies oftheodtheofthe natural vegetation in the north- death valley national monument university of ern mojave desert great basin interface great nevada cooperative national park resources basin naturalist memoirs 4 71 86 studies unit report 03402 las vegas nevada 264 EL GHONEMY A A A WALLACEWAI LACE AND E M ROM 249 DOUGLAS C L 1986b management significance of NEY 1980a frequencyfreFiequency distribution of numbers of soil analyses in relation to vegetation in the cott- perennial shrubs in the northern mojave desert onwood mountains death valley national mon- great basin naturalist memoirs 4 323632 36 ument california university of nevadaofnevada coopera- 265 EL GHONEMY A A A WALLACE AND E M ROM tive national park resources studies unit report NEY 1980b multivariate analysis oftheodtheofthe vegetation 03602 las vegas nevada in a two desert interface great basin naturalist 250 DROBNICK R 1958 the ecology ofaof a relic hybrid oak memoirs 4 40 57 in the great basin area of utah unpublished 266 EL GHONEMY A A A WALLACE E A ROMNEY thesis university ofutahofutah salt lake city 96 appp AND W VALENTINE 1980c A phytosociological 251 DROBNICK R 1960 utah big game range inventory study of a small desert area in rock valley ne- 1959 utah division ofwildlifeofwildlife resources depart- vada great basin naturalist memoirs 4 57 71 mental information bulletin 60160 1 salt lake city 267 ELLISONel LISON L 1941 notes on ecology of subalpine utah 59 appp range plants in central utah proceedings of utah 252 DROBNICK R 1961 utah big game range inventory academy of science arts and letters 18 12 1960 utah division ofwildlifeofwildlife resources depart- abstract mental information bulletin 61361 3 salt lake city 268 elELLISONLISON L 1948 subalpine vegetation of the utah 18 appp wasatch plateau unpublished dissertation uni- 253 DUFF D A 1979 riparian habitat recovery on big versity of minnesota minneapolis creek rich county utah page 91 in 0 B cope 269 ELLISON L 1949a the ecological basis for judging ed proceedings of the forum grazing and condition and trend on mountain range land jour- riparianstreamRiparian Stream ecosystems trout unlimited nal of forestry 47 786 795 inc denver colorado 94 appp 270 ELLISON L 1949b establishment ofvegetationof vegetation on 254 DUTTON C E 1880 report of the geology of the depleted subalpine range as influenced by micro high plateaus of utah atlas USUSDIDI geographical environment ecological monographs 199519 95 121 and geological survey USU S government print- 271 ELLISON L 1954 subalpine vegetation of the ing office washington DCD C 307 appp wasatch plateau utah ecological monographs 255 dzurfrdzureedzurec c R S 1980 carbon isotope ratios of soil 24 89 184 organic matter and their use in assessing com- 272 ELIISONELLISON L 1959 role ofplantofplantpiant succession in range munity composition changes in curlew valley improvement american association for the ad- utah unpublished thesis utah state university vancementvancement of science symposium volume on logan grasslands publication 53 307 321 256 EAGLEY V L 1980 rangeland resource inventory 273 ellELIELLISONlsonISON L 1960 influence ofgrazingofgrazing on plant suc- of the six county area of utah unpublished thesis cession ofrangelands botanical review 26 1 78 utah state university logan 274 ELLISON L AND C M ALDOUS 1952 influence of 257 EARTH environmental consultants incorpo- pocket gophers on vegetation of subalpine grass- rated 1981 soil survey and vegetation inventory land inm central utah ecology 33 177 186 of the proposed Sunnysunnysideside mine site and exten- 275 ELLISON L A R CROFT AND R W BAILEY 1951 sion of the pinnacle mine site volume 2 ap- indicators of condition and trend on high range pendix M pages 254 322 in mining and recla- watersheds of the intermountain region USDA mation plan for centennial project andalexandalea re- agricultural handbook 19 intermountain forest sources utah division of oil gas and mining and range experiment station ogden utah number act007019ACT 007019 salt lake city utah 66 appp 1990 NEVADA UTAH vegetation bibliography 219

276 ELLISON L ANDANDWW R HOUSTON 1958 production general technical report INT 157 intermoun- of herbaceous vegetation in openings and under tain forest and range experiment station og- canopiescanopies of western aspen ecology 39 337 345 den utah 277 EMPIRE engineering AND LAND SURVEYING 1986 293 EVERETT R L S H SHARROW annANDRAND R 0 MFFUWIGMEEUWIG vegetation resources volume 2 chapter 9 sec- 1983 pinyon juniper woodland understory distri- go tions 909.09 0 939.39 3 pages 919 1 9199 19 plus table 9149 14 bution patterns and species associations bulletin pages 9389 38 9419 41 in mining and reclamation plan of the torrey botanical club 110 454 463 for black jack no 1 new mine tech mining 294 EVERETT R L AND S H SHARROW 1985 under- corporation division of gas and utah oil min- story response to tree harvesting in singleleafsingle leaf ing number act019004ACT 019004 salt lake city utah pinyon and utah juniper great basin naturalist 278 ENGLAND J L 1979 measured and inferred mois- 4510545 losios105 112 ture gradient relationships a study of a montane 295 EVERETT R L S H SHARROW ANDDAND D THRAN 1986 steppe in central in utah unpublished thesis soil nutrient distribution under and adjacent to brigham young university provo 36 utah appp singleleafsingleleaf pinyon crowns soil science society of 279 ENGLE D M C D BONHAM BARTEL AND L E america journal 50 788 792 1983 ecological characteristics and control of 296 EVERETT R L AND K WARD 1984 early plant gambel oak journal of range management 36 on controlled 363 365 succession pinyon juniper burns northwest science 58 57 68 280 ERDMAN K S 1961 classification and distribution 297 EVERITT B L 1980 vegetation and of the native trees of utah unpublished thesis sediment mimi- gration in the henry mountains utah brigham young universityuniver sity provo utah 221 appp region pages 208 239 in M picard ed proceedings 281 ESPLIN A C J E GREAVES AND L A STODDART utah geological and mineral survey 1980 1937 A study of utah s winter range utah agri- henry mountains symposium geological cultural experiment station bulletin 277 logan association salt lake utah 49 appp city utah 298 FAUTIN R W 1941 biotic of the 282 EVANS C C 1986 the irelationshipelation ship ofcattle grazing communities northern desert shrub in to sage grouse use of meadow habitat on the shel- biome in western utah don national wildlife refuge unpublished unpublished dissertation university of illinois the- urbana sis university of nevada benoreno 299 FAUTIN R W 1946 283 EVANS F R 1936 A comparative study ofthe vege- biotic communities of the northern desert shrub tation of a grazed and an ungrazed canyon of the biome in western utah wasatch range unpublished thesis university of ecological monographs 1625116 251 310 utah salt lake city 300 FEARS R D 1966 the effects of intensity and season of use on recovery of 284 EVANS P A 1925 A plant ecology study in utah desert range plants unpublished dissertation university ofchicago unpublished thesis utah state university logan 285 EVANS P A 1926 an ecological study in utah botanical gazette 82 253 285 301 FENEMORE R M JR 1970 plant succession in a receding lake great 286 EVENSON W E J D brotherson AND R B bed in the western basin unpublished thesis WILCOX 1980 relationship between environ- university of nevada reno mental and vegetational parameters for under- 302 FERGUSONFFRGUSON C W 1962 annual ring studies of story and open area communities great basin desert shrubs unpublished progress report naturalist 40 167 174 financed by national science foundation grant 287 EVERETT R L 1968 use of the cottonwood in an 65568G 5568 5 appp investigation ofthe recent history of a flood plain 303 FERNANDEZ D A AND M M CALDWELL 1975 american journal ofscience 266 417 439 phenology and dynamics of root growth of three 288 everett R L 1983 understory seed rain on tree cool semisemidesertdesert shrubs under field conditions harvested and unharvested pinyon juniperjumper sites journal ofecology 63 703 714 journal of environmental management 17 304 foifolfolli01tllottLIOTT P F AND W P CLARY 1982 under 349 358 story overstory vegetation relationships an anno- 289 EVERETT R L 1985 creatgreat basin pinyon and ju- tated bibliography USDAUS DA forest service general niper communities and their response to manage- technical report INT 136 intermountain forest ment pages 536153 61 in proceedings ofthe cultural and range experiment station ogden utah physical and biological characteristics of range 39 appp livestock industry in the great basin sympo- 305 FIREMAN M AND H E HAYWARD 1952 indicator sium 38th annual meeting of the society for significance of some shrubs in the escalante range management 11 14 february 1985 salt desert utah botanical gazette 114 143 155 lake city utah 306 FISHER M 1978 A survey of plants and animals of 290 EVERETT R L 1986 understory seed rainram in har- hill and wendover bombing ranges western vested pinyon juniperjumper woodlands great basin utah unpublished report prepared for the de- naturalist 46 706 710 partmentpartment of wildlife science utah state univer- 291 EVERETT R L AND S KONIAK 1981 understory sity logan 70 appp vegetation in fully stocked pinyon juniper stands 307 FLOWERS S 1934 vegetation of the great salt lake great basin naturalist 41 467 475 region botanical gazette 95 353 418 292 EVERETT R L ANDANDSS II11 SHARROW 1983 response 308 FLOWERS S 1942 plant life of the great salt lake ofunderstory species to tree harvesting and fire in region news bulletin ofthe mineralogical society pinyon juniper woodlands USDA forest service of utah 33 36 56 220 P S bourgeronetalbourgeronBouRceronGERON ET AL volume 50

309 FLOWERSFLOWI RS S 1959 vegetation ofglenofgrenofglengien canyon uni- 326 GIUNTA B C 1979 utah big game range inventory versity of utah anthropological papers 40 21 61 1977 utah division of wildlife resources publi- 310 FLOWERSfl owersOWCRS S 1960 vegetation of flaming gorge cation number 79379 3 salt lake city utah 257 appp reservoir basin pages 1 98 in studies oftheofodthethe flora 327 GIUNTA B C 1980 utah big game range inventory and fauna offlamingofflaming gorge reservoir basin utah 1978 utah division of wildlife resources publi- and wyoming university ofutahofutah anthropological cation number 80980 9 salt lake city utah 333 appp papers 48 salt lake city utah 328 GIUNTA B C 1982 utah big game range inventory 311 FOLKS F N 1969 evaluation of five methods for 1980 utah division of wildlife resources publi- sampling desert and wetland vegetation unpub- cation number 82582 5 salt lake city utah 262 appp lished thesis utah state university logan 329 GIUNTA B D BARNHURST B HOSEA T HOWICK 312 forslingFORSI ING C L AND E V STORM 1929 the utiliza- AND J TEEGARDEN 1986 utah big game range tion of browseofbrowse forage as summer range for cattle inm trend studies 1984 utah division of wildlife re- southwestern utah USDAUS DA circular 62 washing- sources publication number 86686 6 salt lake city ton DC 29 appp utah 443 appp 313 forslingFoRsFORSI ingINClingLINC C L 1931 A study of the influence of 330 GIUNTACIUNTA B C AND R MUSCLOW 1983a utah big herbaceous plant cover on surface runoffrun off and soil game range trend studies 1981 utah division of erosion in relation to grazing on the wasatch wildlife resources publication number 83183 1 salt plateau in utah USDAUS DA agricultural technical lake city utah 189 appp bulletin 220 71 appp 331 GIUNTA B C AND R MUSCLOW 1983b utah big 314 FOSBERG F R 1938 the lower sonoran in utah game range trend studies 1982 utah division of science 87 39 40 wildlife resources publication number 83283 2 salt 315 posteFOSTEFOSILRR R H 1968 distribution of the major plant lake city utah 421 appp communities in utah unpublished dissertation 332 GIUNTA B C AND R MUSCLOW 1984 utah big brigham young university provo utah 124 appp game range trend studies 1983 utah division of 316 FOSTIFOSTER R R M 1980 threatened and endangered wildlife resources publication number 84384 3 salt plant inventory wells resource area nevada lake city utah 399 appp unpublished final report prepared for USDI 333 GIUNTA B C R STEVENS K R JORGENSENJOBGENSEN AND bureau of land management elko nevada un- A P PLUMMER 1978 antelope bitterbrushbitterbrusbbitterbrush an der contract YA 512 ctrct9 125 by intermountaininter mountain important wildlandwildwindlandland shrub utah division of research wildlife research publication 781278 12 salt lake 317 FREEMANFRLLMAN J andaandjAND J L MAHONEYMAHONFY 1977 geothermal city utah 47 appp areas inm nevada the distribution of vascular 334 GLEASON H A AND A CRONQUIST 1964 the natu- plants near the thermal springs surveyed ral geography of plants columbia university mentzelia 3 7 14 press new york 420 appp 318 frischknecht N C 1975 native faunal relation-i 335 GOEBEL C J 1960 effect of range condition and ships within the pinyon juniper ecosystem pages plant vigor upon the production and nutritive 55 65 inm pinyon juniper ecosystem A sympo- value of forage unpublished dissertation utah siumslum utah state university logan state university logan 319 frischknecht N C ANDandlANULL E HARRIS 1968 graz- 336 GOETZ H 1968 vegetation and soil responses ing intensities and systems on crested wheatwheatgrassgrass to nitrogen feireibel tihzationfertilization on different range sites in central utah response ofofvegetationvegetation and cattle unpublished dissertation utah state university USU S department of agriculture technical bul- logan letin 1388 washington DCD C 47 appp 337 GONZALES A H 1964 patterns of livestockoflivestock behav- 320 cardinerGARDINERGARDINCK H G 1984 dynamics of and land ior and forage utilization as influenced by environ-environ perennial plant populations with an examination of mental factors on a summer mountain range potential causal agents unpublished dissertation unpublished dissertation utah state university utah state universityumveiumpei sity logan logan 321 GATES D H 1956 ecology ofplantof plantpiant distribution on 338 GOODRICH S 19819811 A flonfloristicstic study of central ne- the salt deserts of utah unpublished disserta- vada unpublished thesis brigham young uni- tion utah state university logan versityversity provo utah 400 appp 322 catesGATFsGATES D H L A STODDART AND C W COOK 339 GOODWIN D L 1963 vegetation and soils relation 1956 soil as a factorbactor influencing plant distributiondistibisti ibution ships oftheofodthethe shadscaleshadscale zone in utah pages 242425942525 in on salt deserts of utah ecological monographs proceedings of the american society of range 2615526 155 175 management 12 15 february 1963 rapid city 323 GENWALGLNWAL COAL COMPANY 1988 vegetation refer- south dakota ence area volume 3 chapter 9 item 929 2 pages 340 GRAHAM E H 1937 botanical studies in the uinta 1 8 in mining and reclamation plan for crandall basin of utah and colorado annals of the canyon mine gendalgenwal coal company utah divi- carnegie museum vol 26 carnegie institute sion of oil gas and mining number act015ACT 015 pittsburgh pennsylvania 432 appp 032 salt lake city utah 341 GRAYBOSCH R A AND H BUCHANAN 1983 vegeta- 324 GERMANOgl RMANO D J AND D N LAWHEAD 1986 species tive types and endemic plants oftheofodthethe bryce canyon diversitydiver sitssity and habitat complexity does vegetation breaks great basin naturalist 43 701 712 organize vertebrate communities min the great 342 GREENWOOD L R AND J D brothersonBROT hersonHEBSON 1978 basin great basin naturalist 46 711 720 ecological relationships between pinyonpinsonyon juniper 325 ginglnGIMGIMORDGIFFORDORD G F AND F E BUSBY FDS 1975 the and true mountain mahogany stands in the uintah pinyon juniper ecosystem a symposium utah basin utah journal of range management 31 state university logan 194 appp 164 167 1990 NEVADA UTAH vegetation bibliography 221

343 GRIFFITHS D 1902 forage conditions in the north- 359 HARNISS R 0 AND K T HARPER 1982 tree dy- ern border of the great basin USDAUS DA bureau of namics in seraiseral and stable aspen stands in central plant industry bulletin 15 utah USDA forest service research paper INT 344 cruellGRUELLGRUFLL G E S BUNTINGBUNFING ANDANDLL neuenschwan 297 intermountain forest and range experiment DERDEB 1985 influence offire on curlleafcurlcurlleanleaf mountain station ogden utah 7 appp mahogany in the intermountamintermountain west pages 360 HARPER K T 1959 vegetational changes in a shad 58 72 inTO J E lotanlotanandjand J K brownbi own compilers scale winterfat plant association during twenty fire s effects on wildlife habitat symposium pro- three years of controlled grazing unpublished ceceedings USDAUS DA forest service general techni- thesis brigham young university provo utah cal report INT 186 intermountain forest and 72pp72 appp range experiment station ogden utah 361 HARPER K T 1967 the vegetational environment 345 HAAS R H H L MORTON AND P J TORELL 1962 of the bear river number 2 archaeological site influence of soil salinity and 24 D treatments on university of utah anthropological paper 87 salt establishment of desert wheatgrasswheatgrass and control of lake city utah halogeton and other annual weeds journal of 362 HARPER K T 1976a permanent plot native legetavegeta range management 15 205 210 tion studies pages 20 43 in A C hill et Aal eds 346 HALL H H 1954 the impact ofmanofman on the vegeta- vegetation air pollution investigations in the tion and soil of the upper valley allotment vicinity of four corners and san juan power garfield county utah unpublished thesis uni- plants new mexico university of utah research versity of utah salt lake city institute environmental studies laboratory salt 347 HALL H H 1971 ecology ofaofa subalpine meadow of lake city the aquarius plateau garfield and wayne coun- 363 HARPER K T 1976b work plan ecological impact ties utah unpublished dissertation brigham study of weatherofweather modification in the uinta moun young university provo utah tainsbains utah unpublished report prepared for division of resources 348 HALVERSON BR M 1974 ERTS 1 imagery for vege- utah ofwaterwater salt lake city and the USDI tation mapping and phenology detection in ne- bureau of reclamation office of vada unpublished thesis university of nevada atmospheric resources management denver reno colorado 46 appp 364 HARPER K T 1977 ecology 349 HANKS L M 1982 pollinator community of a the utah project eco- the logical sand dune ecosystem unpublished thesis uni- impact of weather modification studies in the mountains versity of nevada reno uinta utah unpublished interim report prepared for utah division of water re- 350 HANSEN D J 1977 interrelations ofofvalleyvalleyvailey vegeta- sources salt lake city and the USDI of tion stream soils and solar irradiation bureau regimen reclamation denver colorado along the rock creek in the uinta mountains of 365 HARPER K T 1978 ecology study bi- utah unpublished dissertationdisser tahon university of the uinta ennial report unpublished for michigan ann arbor report prepared utah divisondavison of water resources salt lake city 351 HANSEN D &S P dayanandanDAY ANANDAN P B KAUFMAN AND and USDI bureau ofreclamationofreclamation denver colo- J D BROTHEBSON 1976 ecological adaptations of brothersonherson rado 24 appp salt maishmarshmalsh glassgrass distichlis spicatespicata Gramgramineaemeae 366 HARPER K T 1979 some reproductive and life and environmental factors affecting growth and its history characteristics of rare plants and implica- distribution journal 635 american of botany 63 tions for management great 650 basin naturalist memoirs 3 129 138 352 HANSEN H P 1947 postglacial vegetation of the 367 HARPER K T 1985 predicting successional rates in great basin journal of northern american botany utah aspen forests pages 96 100 in proceedings 34 164 171 34164 oftheofodthethe 1985 society ofamericanofamerican foresters national HANSON C A 1962 perennial 353 atriplex of utah and convention 28283131 july 1985 bethesda mary- the northern deserts unpublished thesis brig- land 445 appp ham young university provo utah 368 HARPER K T AND C S CUMERCLIMER 1985 factors af- 354 HANSON H C 1950 ecology of the grassland II11 fecting productivity and compositional stability of botanical review 16 283360283 360 artemisia steppes in idaho and utah USA 355 HANSON W R 1939 the ecology of agropyron pages 592 594 in proceedings of xvthxyth inter- inerweinerwthermetnerme on protected and heavily grazed range land national grassland conference august kyoto in cache valley utah unpublished thesis utah japan state university logan 33 appp 369 HARPER K T D C FREEMAN W K OSTLER AND 356 HANSON W BR ANDLAND L A STODDART 1940 effects of L G KLIKOFF 1978 the flora of great basin grazing upon bluebunchblue bunch wheat grass journal of mountain ranges diversity sources and disper- the american society of ofagronomyagronomy 32 279 289 sal ecology great basin naturalist memoirs 2 357 HARNER R F AND K T HARPER 1973 mineral 81 103 composition of grassland species of the eastern 370 HARPER K T AND R A JAYNES 1986 some edaphic great basin in relation to stand productivity and compositional characteristics of artemisiaofartemisia tritrl canadian journal ofbotany 51 2037 2046 dentata and associated plant communities in 358 HARNERHARNFR R F AND K T HARPER 1976 the role of southeastern utah pages 265 272 in E D area heterogeneity and favorabilityfavorability inm plant spe- mcarthur and B L welch compilers proceed- cies diversity ofpinyonof pryonpmyon juniperjumper ecosystems ecol- ings symposium on the biology of artemisiaofartemisia and ogy 57 1254 1263 chrysothamnus 9 13 july 1984 provo utah 222 P S BOURGERON ETETALAL volume 50

USDA forest service general technical report university center for health and environmental INT 200 intermountain research station og studies provo utah den utah 382 HARPER K T R A WOODWARD AND K B 371 HARPER K T J R MURDOCK AND S L WELSH mcknight 1980 interrelationships among pre- 1974 botanical evaluation of proposed power cipitation vegetation and streamstreamflowflow in the uinta plant sites in garfield and wayne counties utah mountains utah encycliaencyclicencycha57Encyeneycliaellaeila 57 56 86 unpublished report prepared for environmental 383 HARRIS M L 1926 an ecological study of tim systems department of westinghouse corpora- palogospanogos creek from aspen grove to wildwood tion salt lake city utah unpublished thesis brigham young university 372 HARPER K T J R MURDOCK AND S L WELSH provo utah 82 appp 1975 the vegetation of salt wash area wayne 384 HARRISON BFB F 1980 botanical survey threatened county utah unpublished report prepared for endangered and other rare plants of the schell environmental systems department of westing- resource area ely district nevada unpub- house corporation salt lake city utah 30 appp lished report prepared for USDI bureau of land 373 HARPER K T W K OSTLER AND DCD C ANDERSON management by intermountaininterintel mountain research cor 1978 the utah ecology project ecological impact horationporationporation provo utah 101 appp of weather modification studies in the uinta 385 HASSAN M A 1983 effects of wildfire on seed mountains utah unpublished special report rainram and soil seed reserve dynamics of a good prepared for utah division of water resources condition sagebrush grass rangeland in central salt lake city and USDI bureau ofreclamation utah unpublished thesis utah state university denver colorado logan 374 HARPER K T W K OSTLER AND K B McKMCKNIGHTnightNICHT 386 HASSAN M A AND N E WEST 1984 soil seed 1979 the uinta ecology study interim report reserve dynamics in burned and unburned sage unpublished report to utah division of water brush grass vegetation ofcentral utah bulletin of resources salt lake city and USDI bureau of ecological society america 65 180 reclamation denver colorado 76 appp 387 HAYWARD C L 1943 biotic communities of the 375 HARPER K T W K OSTLEROSTLEB K B McKMCKNIGHTnightNICHT AND coniferous forests of the southern wasatch and D L HUNTER 1981a potential ecological impacts uinta mountains utah unpublished thesis uni- of snow pack augmentation in the uinta moun- versity of illinois urbana tains utah unpublished report prepared for 388 HAYWARD C L 1945 biotic communities of the USDI bureau of reclamation denver colo- southern wasatch and uinta mountains utah rado 291 appp great basin naturalist 6 1 124 376 HARPER K T W K OSTLER K B McKMCKNIGHTnightNICHT AND 389 HAYWARD C L 1948 biotic communities of the D L HUNTER 1981b effects oflate lying snow on wasatch chaparral utah ecological monographs an alpine herblandherslandherbland in the uinta mountains pages 1847318 473 506 179 203 in K T harper W K ostler K B 390 HAYWARD C L 1952 alpine biotic communities of mcknight and D L hunter eds potential eco- the uinta mountains utah ecological mono- logical impact of snow pack augmentation in the graphs 22 93 120 uinta mountains of utah unpublished report 391 HAYWARD C L D E BECK ANDWAND W W TANNER prepared for USDIUS DI bureau of reclamation den- 1958 zoology ofthe upper colorado river basin ver colorado part I1 the biotic communities brigham young 377 HARPER K T AND J L REVEAL EDS 1978 inter- university science bulletin biological series 13 mountain biogeography a symposium great 1 74 basin naturalist memoirs 212 1 268 392 HLINZE D H R E ECKERT AND P T TUELLER 378 HARPER K T &S C SANDERSON AND E D 1962 the vegetation and soils of the steptoe mcarthur 1987 vegetation communities of watershed unpublished report prepared for zion national park pages 191 196 in university USDI bureau of land management 40 appp of wyoming national park service research cen- 393 HENDERSON J A 1978 ECOSYM potential vege- ter eleventh annual report laramie wyoming tation classification appendix report 7 in J A 379 HARPER K T F J WAGSTAFF AND L M KUNZLER henderson L S davis and E M ryberg eds 1985 biology and management ofthe gambel oak ECOSYM an ecosystem classification and data vegetative type a literature review USDA forest storage system foiforhoiboi natural i esourcesresources management service general technical report INT 179 inter- unpublished report prepared for utah state uni- mountain forest and range experiment station versityversity logan 12 appp ogden utah 394 HENDERSONHENDFRSON J A R L MAUK D L ANDERSON 380 HARPER K TTSS walteWAITEWAITEANDCAND C LAMB 1973 vegeta- T A DAVIS AND T J KECK 1977 preliminary tion ofthe proposed power generating plant sites forest habitat types of the uinta mountains utah pages 1 33 in evaluation of three alternative unpublished report prepared for department of power plant sites in utah and wyoming for the forestry and outdoor recreation utah state uni- utah power and light company salt lake city versityversity logan 94 appp utah 395 HENDERSONHENDFRSON J A S A SIMON AND S B HART 381 HARPER K TTBB W WOOD AND J D brotherson VIGSEN 1977 plant community types and habitat 1974 A vegetational analysis of areas involved in types of the pricepi iceieelee district mantilamanti la sal national the proposed four seasons project pages 60 104 foiestforest unpublished report prepared foiformoimor depart- in unpublished environmental report on proposed ment of forestry and outdoor recreation utah four seasons project prepared for brigham young state university logan 1990 NEVADA UTAH vegetation bibliography 223

396 HENDERSON J A AND N E WEST 1977 410 HOLMGRENHOLMGBEN A H 1948 handbook of the vascular ECOSYM vegetation classification appendix plants of the northern wasatch lithotype process report 6 in J A henderson and L S davis eds company san francisco 202 appp ECOSYM a classification and data storage system 411 HOLMGREN A H 1962 the vascular plants of the for natural resources management unpublished green river flomfrom the flaming gorge to split report prepared for utah state university logan mountain gorge unpublished report prepared 8989ppappp for the national park service by utah state uni- 397 henningson DURHAM & richardson INC 1981a versityversity logan 40 appp preliminary FEIS grazing MXM X environmental 412 holHOIHOLMGRENMGREN A H 1979 strategies for preservationpiepreservation technical report 40 unpublished report pre- of rare plants gleatgreat basin naturalist memoirs 3 pared for USAF norton air force base califor- 95 100 nia 168 appp 413 HOLMGREN A H ANDJAND L REVEAL 1966 checklist 398 henningson DURHAM & richardson INC 1981b of the vascular plants of the intermountain re- preliminary FEIS native vegetation MXM X envi- gion USDAUS DA forest service research paper INT ronmentalronmental technical report 14 unpublished re- 32 intermountain forest and range experiment port prepared for USAF norton air force base station ogden utah 160 appp california 197pp197 appp 414 HOLMGREN R C 1975 the desert experimental 399 hfnningsonhenningson DURHAM & richardson INC 1981 range description history and program pages preliminary FEIS aquatic habitats and biota 18 22 in andaridarld shrubland proceedings oftheodtheofthe third MXM X environmental technical report 16 unpub- workshop of the united statesaustraliaStates Australia rangel- lished report prepared for USAF norton air ands panel tucson arizona 1973 force base california 70 appp 415 HOLMCRENHOLMGREN R C AND S F BREWSTER JR 1972 400 HERMAN F 1953 A growth record of the utah distribution of organic matter reserve in a desert juniper journal of forestry 5151193193 195 shrub community USDAUS DA forest service re- 401 HESSING M B C D JOHNSON AND R P BALDA search paper INT 130 intermountain forest and 1982 early secondary succession of a pinyon range experiment station ogden utah juniper woodland in a northern arizona popowerlinewerline 416 HOLMGBENHOLMGREN R C AND S S HUTCHINGS 1972 salt corridor juniperus monomonospermamonospermysperma pinus edadulisedulisulis desert shrub response to grazing use pages southwestern naturalist 2727127.11 9 153 164 in wildlandWildwindlandland shrubs their biology and 402 HIEBERTHIEBERTRR D ANDJAND J L HAMRICK 1984 an ecolog- utilization USDA forest service general techni- ical study of bristlecone pine pinus longaevalongaeva in cal report INT GTR 1 intermountain forest and utah and eastern nevada great basin naturalist range experiment station ogden utah 44487 494 417 HOSEAHOSFA B ANDANDDD BARNHURST 1988 utah big game 403 HIGGINS L C 1967 A flora of the beaver dam range trend studies 1986 utah divisondavison of wild- mountains unpublished thesis brigham young life resources publication number 88588 5 salt lake university provo utah 305 appp city utah 305 appp 404 HILL A C K T HARPER AND W H EDWARDS 418 HOUSTON W R 1951 A preliminary study of some 1972 ecological evaluation ofproposedofproposed routes for factors affecting forage production in the aspen the huntington sigurd power transmission line type ofcentralofcentral utah unpublished thesis univer- unpublished report prepared for utah power and sity of utah salt lake city light company salt lake city utah 419 HOUSTON W R 1954 A condition guide for aspen 405 HILTONJHILTON J W 1941 effects ofcertainofcertain micro ecologiecology ranges of utah nevada southern idaho and cal factors on the germinability and early develop- western wyoming USDA forest service res- ment of errotaeurota lanata northwest science 15 earch paper 32 intelinterintermountainmountain forest and range 86 92 experiment station ogden utah 25 appp 406 HOFFMAN W J 1877 the distribution ofvegetationofvegetation 420 HUBBARD K G 1980 relating fire occurrence to in portions of nevadaofnevada and arizona american nat- weather conditions on the great basin rangelands uralist 111133611.336336 343 journal ofrangeorrangeof range management 33 360 362 407 HOLLAND J 1982 A floristic and vegetation analysis 421 HUFF C L 1962 utah big game range inventory of the newberry mountains clark county ne- 1961 utah division ofwildlifeofwildlife resources depart- vada unpublished thesis university of nevada mental information bulletin number 62562 5 salt las vegas reproduced in the lake mead report lake city utah 45 appp series by university of nevada cooperative na- 422 HUFF C L 1963 utah big game range inventory tional park resources studies unit las vegas 1962 utah division ofwildlifeofwildlife resources depart- nevada mental information bulletin number 63263 2 salt 408 HOLLAND J S W E NILES AND P J LEARY 1979 lake city utah 161 appp vascular plants of the lake mead national recre- 421423 HUFF C L AND R BLOTTER 1964 utah big game ation area lake mead technical report 3 uni- range inventory 1963 utah division of wildlife versity of nevada cooperative national park re- resources departmental information bulletin sources studies unit las vegas nevada 223 appp number 64264 2 salt lake city utah liollo110hoppnoppappp 409 HOLLAND J S W E NILES AND D R SCHRAM 424 HUFF C L AND J E BOWNS JR 1965 utah big 1980 A guide to the threatened and endangered game range inventory 1964 utah division of vascular plants of the lake mead national recre- wildlife resources publication number 65165 1 salt ation area lake mead technical report 4 uni- lake city utah 111 appp versity of nevada cooperative national park re- 425 HUFF C L AND F H COLES 1966 utah big game sources studies unit las vegas nevada 75 appp range inventory 1965 utah division of wildlife 224 P S bourgeronBoURbourgeronetalceronGERON ET AL volume 50

resources publication number 66366 3 salt lake 443 JENSEN A E 1989 soil characteristics of moun- city utah 96 appp tainous northeastern nevada sagebrush commu- 426 HULL A C JR AND M K HULL 1974 presettle- nity types great basin naturalist 49 469 481 ofcache ment vegetation of cache valley utah and idaho 444 JENSEN M E L S PECK AND M V WILSON 1988 journal of range management 27 27 29 A sagebrush community type classification for 427 HUNT C B P AVERITT AND R L MILLER 1953 mountainous northeastern nevada rangelands geology and geography of the henry mountains creatgreat basin naturalist 48 422 433 region utah USDI geological survey profes- 445 JENSEN S E ANDahdjJ S TUHY 1981 soils investiga- sional paper 228 washington DCD C ahde tion of riparian communities of east smiths fork 428 HUTCHENS S K ANDANDKK T HARPER 1965 A method and henrys fork drainages north slope for determining indicators in aspen com- uinta grazing in mountains utah unpublished presented munitiesties proceedings of colorado wyoming report muni to USDA service ogden utah 35 appp academy of science arts and letters 42 319 forest C 429 IBRAHIM K 1962 plant communities of the shad 446 JOHNSON A 1970 common native trees ofutahof utah scale zone in grand county as related to soil and USDA forest service special report 22 utah geology proceedings of utah academy of sci- agricultural experiment station logan utah ence arts and letters 37 39 191 193 109 appp 430 IBRAHIM K M 1963 ecological factors influencing 447 JOHNSON H B 1964 changes in the vegetation of plant distribution in the shadscaleshadscale zone of south- two restricted areas of the wasatch plateau as eastern utah unpublished dissertation utah related to reduced grazing and complete protec- state university logan tion unpublished thesis brigham young univer- 431 IBRAHIM KMK M N E WEST AND D L GOODWIN sity provo utah 123 appp 1972 phytosociological characteristics of peren- 448 JOHNSON J L ANDANDRR D PFISTER 1982 A survey of nial atriplex dominated vegetation of southeast- potential ecological natural landmarks of the mid- ern utah vegetatiovegetation 24 13 22 dle rocky mountains unpublished report pre- 432 IRVINE J R 1970 riparian environmental legetavegeta pared for USDI national park service by USDA tion interrelationships along the escalante river forest service intermountain forest and range glen canyon national recreation area utah experiment station ogden utah 197 appp unpublished thesis utah state university lo- 449 JOHNSON K L 1986 sagebrush types as ecological gan 82 appp indicators to integrated pest management IPM in 433 IRVINEIRVINL J R AND N E WEST 1976 riparian enviedvi the sagebrush ecosystem of western north amer- ronmentalnonmental vegetation interrelationships along the icaleaiea pages 1 10 in J A onsager ed integrated lower escalante glen canyon national ica in river pest management in rangeland state oftheodtheofthe art inm area utah unpublished report pre- in recreation sagebrush ecosystem 27 28 march 1984 pared for the national park service utah 82 appp university of nevada reno 434 IRVINE J R AND N E WEST 1979 riparian tree distribution and along the 450 JOHNSON K L ED 1989 rangeland resources of species succession coop- lower escalante river utah southwestern natu- utah utah rangeland committee report ralist 24 331 346 erative extension service utah state university 103 appp 435 ISAACSON H E 1967 ecological within logan provinces patterns the pinyon juniperjumper type of the great basin and 451 JOHNSON N K 1978 of asian geography colorado plateau unpublished thesis utah state and specialism in the intermountain region university logan great basin naturalist memoirs 2 137 160 436 IVES R L 1942 atypical subalpine environments 452 JOHNSON R V 1929 A taxonomic ecological survey ecology 23 89 96 of the truckee meadows unpublished thesis 437 JACKSON L E 1985 fionflonsticfloristicFlonstic analysis of the distri- university of nevada reno bution of ephemeral plants in tretreelinetreehnetreebineeline areas of the 453 KAISER COAL corporation 1985a vegetation vol- western U S arctic and alpine research 17 ume 3 chapter 8 sections 808.08 0 838.38 3 pages VIII- 251 260 I VIII 21 plus appendices VIIIIVIII I1 VIII 3 VIII 5 438 JACKSON L E AND L C BLISS 1984 phenology in mining and reclamation plan for horse canyon and water relations ofthreeof three plant life forms in a dry mine kaiser coal corporation utah divisondavison of tree lined meadow ecology 65 1302 1314 oil gas and mining number act007013ACT 007013 salt 439 JAREK L B 1986 A photographic history ofvegetaof vegeta- lake city utah tion changes on the humbolt national forest 454 KAISER COAL corporation 1985b vegetation re- unpublished thesis university ofnevadaofnevada benoreno sources book 7 chapter 9 sections 919.19gi 1 939.39 3 440 JATKAR S A S R RUSHFORTH ANDANDJJ D BROTHER pages 1 8 plus tables IX 1 IX 38 plus appendix SON 1979 diatom floristics and succession in a IX 1 in mining and reclamation plan for sunny peat bog near lily lake summit county utah side mine kaiser coal corporation utah divi- great basin naturalist 39 15 43 sionslon of oil gas and mining number act007ACT 007 441 JAYNES R A ANDANDKK T HARPER 1978 patterns of 007 salt lake city utah natural revegetation in and southeastern utah 455 KAY P A ANDANDCC G OVIATT 1978 pinus longaevalongaeva in journal of range management 31 407 411 the stansbury mountains utah great basin nat- 442 JENSEN G H 1940 the relation of some physical uralist 38 49 50 and chemical factors oftheofodthethe soil to the productivity 456 KEARNEY T H L J BRIGGS H L SHANTZ J W and distribution ofccrtainofcertain waterfowl food plants at MCLANE ANDANDRR L PIEMEISEL 1914 indicator sig- bear river migratory waterfowl refuge unpubunput nificancenificance of vegetation in tooelethoele valley utah lisheddished thesis utah state university logan journal ofagriculturalofagricultural research 1 365 417 1990 NEVADA UTAH vegetation bibliography 225

457 KENDEIGH S C 1954 history and evolution of 472 KLINE L G 1973 bromegrassBromegrass productivity in rela- variousvailous concepts of plant and animal communities tion to precipitation on shrubs canopy cover and inm north america ecology 35 152 172 soil nitrogen content unpublished thesis utah 458 KENNEDY P B 1910 plant distribution on the state university logan truckee carson project fallenfallonfailon nevada muh- 473 KONIAK S 1985 succession in pinyon juniper lenbergialenbergia 6 woodlands following wildfire in the great basin 459 KERR C W ANDANDJJ A HENDERSONHENDEBSON 1979a upland great basin naturalist 45 556 566 vegetation classification and map for a test area 474 KONIAK S 1986 successional processes in pinyon manti la sal national forest appendix report 13 juniper woodlands in the great basin unpub- in J A henderson and L S davis eds lished thesis university of nevada reno ECOSYM a classification and information system 475 KORSTIAN C F 1921 effect oflateofoblatelate spring frost upon for wildlife resource management unpublished forest vegetation in the wasatch mountains of for report prepared utah state university logan utah ecology 2 47 52 128 appp 476 KUNZLERKUNZLEB L M AND K T HARPER 1980 recovery 460 KERR C W ANDANDJJ A HENDERSONHENDFRSON 1979b upland of gambel oak after fire in central utah great potential vegetation classification and map for a basin naturalist 40 127 130 test area manti sal national forest appendix la 477 KUNZLER L M K T HARPER AND D B KUNZLER report 15 min J A henderson L S davis and 1981 compositional similarity within oak E M ryberg eds ECOSYM a classification and the brush great information system for wildlife resource manage- type in central and northern utah ment unpublished report prepared for utah basin naturalist 41 147 153 state university logan 68 appp 478 KURZIUS M 1981 vegetation and flora of the 461 KING S AND A MUIR 1971 utah big game range grapevine mountains death valley national inventory 1970 utah division of wildlife res- monument californianevadaCalifornia Nevada unpublished the- ources publication number 71371 3 salt lake city sis university of nevada las vegas utah 208 appp 479 LAMBERT C B 1940 the distribution ofplantof plantpiant types 462 KING S AND B C OLSON 1972 utah big game of the escalante desert of utah with relation to range inventory 1971 utah division of wildlife soil condition unpublished thesis brigham resources publication number 721072 10 salt lake young university provo utah city utah 451 appp 480 LAMARCHE V C AND H A MOONEYMOONFY 1972 recent 463 KIPPING K L 1969 ground photographic tech- climatic change and development of the bristle- niques for range condition and trend analysis un- cone pine P longaevalongaeva bailey krummholz zone published thesis university of nevada reno mount washington nevada arctic and alpine 464 KLEBENOW D A R BEALL A BRUNER R MASON research 4 61 72 B ROUNDY W STAGER anukANDKAND K WARD 1977 sum- 481 LANNER R M 1971 conifers oftheodtheof the bear lake area mary progress report fy1977 controlled fire as a and mountains south oftheodtheofthe great salt lake great management tool in the pinyon juniper woodland basin naturalist 31 85 89 university of nevadaofnevada max fielschfleischmannFleischmarm college of 482 LANNER R M 1975 pinyon pines and junijunipersjuniperuspers of agriculture miscellaneous publication 29 un3aunaa the southwestern woodlands pages 1 7 in G F 9 c76 reno nevada 155 appp gifford and F E busby eds pinyon juniper 465 KLFINERKLEINER E F 1966 A study of the vegetational ecosystem A symposium utah state university communities of red butte canyon salt lake logan county utah unpublished thesis university of 483 LANNER R M ANDANDEE R HUTCHISON 1972 relict utah salt lake city 54 appp stands of pinyon hybrids in northern utah great 466 KLEINER E F 1968 comparative study of grass- basin naturalist 32 171 175 lands ofcanyonlands national park unpublished 484 LANNER R M ANDANDSS B vanderwallVANDER WALL 1980 dis- dissertation university of utah salt lake city persal of limber pine seed by clark s nutcracker 58 appp journal of forestry 78 637 639 467 KLEINER E F 1983 successional trends in an un- 485 LANNER R M AND R WARNICK 1971 conifers of grazed and grassland over a decade journal of the san francisco mountain san rafael swell range management 3611436 114 118 and roan plateau great basin naturalist 31 468 KLEINER E F AND K T HARPER 1966 an investi- 177 180 gation of association patterns of prevalent grass- 486 LARRISON E J AND D R JOHNSON 1973 density land species in red butte canyon salt lake changes and habitat affinities of rodents of shad county utah proceedings of utah academy of scale and sagebrush associations great basin nat- science arts and letters 43 29 36 uralist 33 255 264 469 KLEINER E F AND K T HARPER 1972 environ- 487 LAVIN M 1981 the flonsticsfloristicsfloristiesflori sties of the headwaters of ment and community organization in grasslands of the walker river california and nevada unpubl- canyonlandsCanyon lands national park ecology 53 299 309 ished thesis university of nevada reno 470 KLEINER E F AND K T HARPER 1977a occur- 488 LAWTON P M 1979 an investigation of the envi- rence ofoffouroffousfourmour major perennial grasses in relation to ronmentalronmental relationships of selected forest habitat edaphic factors in a pristine community journal of types in northern utah unpublished thesis utah range management 30 286 289 state university logan 102 appp 471 KLEINER E F AND K T HARPERHARPFR 1977b soil prop- 489 LAYCOCK W A 1969 exclosure and natural areas erties in relation to cryptogamic groundcovergroundcover in on rangelands in utah USDA forest service re- canyonlandsCanyonlands national park journal of range search paper INT 62 intermountain forest and management 30 202 205 range experiment station ogden utah 44 appp 226 P S BOURGERON ETETALAL volume 50

490 LAYCOCK W A AND P W CONRAD 1981 res- tionaldional park pages 195 199 in R M linn ed ponses ofvegetationofvegetation and cattle to various systems proceedings first conference of scientific re- of grazing on seeded and native mountain range- search in the national parks vol I1 9 13 novem- lands in eastern utah journal of range manage- ber 1976 new orleans lousiana USDI na- ment 345234.5234 52 58 tional park service transactions and proceeding 491 LEARYLEARYKK DANDPD AND P M PETERSONPETEBSON 1984 soil analy- series number 5 ses in relation to vegetation in the cottonwood 506 LORAIN G E 1970 70 mm large scale aerial photog- mountains death valley national monument raphy for range resources evaluation unpub- university of nevada cooperative national park lished thesis university ofnevada reno 93 appp vegas resources studies unit report 03601 las 507 LOTAN J E 1975 the role ofconeofeonecone serotinaserotinyserotiny in lodge- nevada pole pine forests pages 471 495 in proceedings 492 LEARYPJLEARY P J 1986 plant succession after fire hunter symposium on management of lodgepole pine mountain death valley national monument washington state university pullman university of nevada cooperative national park 508 louderbough E T ANDLAND L D POTTER 1982 man- resources studies unit technical report 03403 cos shale some physical and chemical properties las vegas nevada which affect vegetative communities on shale out- 493 LEARY P J AND W E NILES 1976 investigation of crops pages 547 550 in proceedings symposium vegetational communities in joshua tree national on surface mining hydrology sedimentologySediment ology and monument university of nevada cooperative reclamation university of kentucky lexington park resources studies national unit technical 509 LOVEJOY T E 1979 epoch of biotic vegas the impover- report 2 las nevada ishment great basin naturalist memoirs 3 5 10 494 lesperance A L J A YOUNG R E ECKERT JR 510 LUDWIG J A 1965 relationships of grasslands to AND R A EVANS 1978 great wildryewildrye basin edaphic gradients of the foothill zone salt lake rangeman s journal 5 125 127 county utah unpublished thesis university of 495 LEWIS M E 1970 alpine rangelands of the uinta utah salt lake city mountains ashley and wasatch national forests 511 LUDWIG J A 1969 environmental interpretation region 4 ofthe US service unpublished forest of foothill grassland of northern report prepared for USDA service region communities forest utah unpublished dissertation university of IV ogden utah 75 appp utah salt lake city 100 appp 496 LEWIS M E 1971 flora and major plant communi- 512 MACDOUGAL D T 1908 ties of the ruby east humboldt mountains with sagebrush deserts of nevada pages 28 special emphasis on lamoille canyon unpub- and utah 30 in botanical features lished report prepared for USDA forest service of north american deserts carnegie institute of washington publication 99 region IV ogden utah 62 appp 497 LEWIS M E 1973 wheeler peak area species list 513 MACMAHON J A ANDTAND T F WIEBOLDT 1978 ap- inventinventoriesinventoriedoried and cataloged as of januaryofjanuary 1973 un- plying biogeographic principles to resource man- published report prepared for USDA forest ser- agement a case study evaluating holdridge s life zone great vice region IV ogden utah 17 appp model basin naturalist memoirs 2 498 LEWIS M E 1975a flora of the santa rosa moun- 245 257 tains unpublished report prepared for USDA 514 MADANY M H 1980 Peterpetersborosboro prairie unpub- forest service region IV ogden utah 19 appp lished report on field project prepared for depart- 499 LEWIS M E 1975b plant communities of the jar ment of range science utah state university bidge mountain complex humboldt national logan utah 19 appp forest unpublished report prepared for USDA 515 MADANY M H 1981 land use fire regime interac- forest service region IV ogden utah 22 appp tions with vegetation structure ofseveralofseveral montane park 500 llnsdalemLINSDALEMAAJJ T HOWELLhowellandjAND J M LINSDALE forest areas of zion national unpublished 1952 plants of the toiyabe mountains area ne- thesis utah state university logan 103 appp vada wasmann journal of biology 10 129 200 516 MADANY M H no date ponderosa pine forests of 501 livingstonlivingstonblivingstonbaB EANDFE AND F SHREVE 1921 the distri- the colorado plateau unpublished report pre- bution of vegetation in the united states as re- pared for dr N E west department of range lated to climatic conditions carnegie institution science utah state university logan publication 284 washington DC 590590ppappp 517 MADANY M H ANDANDNN E WEST 1980a fire history 502 LOEW 0 1876 report on the geographic distribu- of two montane forest areas of zion national tion of vegetation in the mohave desert pages park pages 50 56 in proceedings oftheodtheofthe fire his- 442 444 in G M wheeler chief of engineers tory workshop USDA forest service general annual report US geographical and geological technical report RM 81 rocky mountain forest survey west oftheodtheofthe 100th mendanmeridan appendix 7 and range experiment station fort collins col- 503 LOOPE L L 1969 subalpine and alpine vegeta- orado tion of northeasternofriortheastern nevada unpublished thesis 518 MADANY M H AND N E WEST 1980b fire duke university durham north carolina regime and land use interactions in ponderosa 504 LOOMLOOPE W L 1977 relationships of vegetation to pine forests of zion national park utah bulletin environment in canyonlandsCanyon lands national park utah oftheodtheof the ecological society of americaofamerica 61 122 unpublished dissertation utah state university 519 MADANY M H AND N E WEST 1982 montane logan vegetation of several relict mesas in zion national 505 LOOMLOOPE W L AND N E WEST 1979 vegetation park utah abstracts of the annual meetings of in relation to environments of canyonlandsCanyon lands na the society for range management 35 35 1990 NEVADA UTAH vegetation bibliography 227

520 MADANY A H AND N E WEST 1983 livestock 535 MASON L R ANDANDVV MORTENSEN 1977 vegetation grazing fire regime interactions within montane and soils of zion national park unpublished re- forests of zion national park utah ecology 64 port prepared for USDAUS DA soil conservation ser- 661 667 vicevlee salt lake city utah 521 MADANY A H ANDNAND N E WEST 1984 vegetation 536 MASON L R AND N E WEST 1970 timber top of two relict mesas in zion national park journal mesa a relict area in zion national park proceed- of range Mmangement27angement 27 456 461 ings of utah academy of science arts and lett- 522 MAJOR J 1948 elements ofaofa range condition classi- ers 47 284 285 ficationfication of the aspen type inm utah proceedings of 537 MAUK R L ANDANDJJ A HENDERSONHENDEBSON 1984 conifer- the utah academy ofscienceofscienceselence arts and letters 25 ous forest habitat types of northern utah USDA 173 174 forest service general technical report INT 523 MALANSON G P 1980 habitat and plant distribu- 170 intermountain forest and range experiment of narrones tions in hanging gardens the barronesNarrones zion station ogden utah 89 appp national park great naturalist 40 utah basin 538 MCDONALD J E 1946 measuring the forage 178 182 crop of range lands by vegetation analysis un- MALANSON G 1982 assembly of hanging 524 C P the published thesis utah agricultural college effects and state gardens of age area location ameri- logan can naturalist 119 145 150 539 mcgraw J F 1980 landsatlandsaw computer aided 525 MARIAH associates 1981 vegetation baseline data analysis techniques for range analysis alton coal lease study area volume 6 vegetation mapping unpublished thesis of reno chapter 3 appendix 363 6 A in mining and reclama- university nevada 540 MCKELL C M 1950 A study tion plan for alton mine utah international in- ofplantof plantpiant succession inm corpcorporatedorated utah division offilofoilofoil gas and mining the oak brush quercus gamberigambehigamb ehi zone after fire number act025003ACT 025003 salt lake city utah unpublished thesis university of utah salt lake 526 MARIAH associates 1989 vegetation data report city 79 appp volume 4 chapter 9 section 9 1 pages 1 47 in 541 mclaughlin S P 1986 fionfloristicFlon sticstie analysis of the mining and reclamation plan for castle gate coal southwestern united states great basin natural- mine castle gate coal company utah division ist 46 46 65 offilofoilof oil gas and mining number act007004ACT 007004 542 MCMILLAN C 1948 A taxonomic and ecological 527 MARKHAM B S 1939 A preliminary study of the study of the flora of the deep creek mountains of vegetation cover in spanish fork canyon utah central western utah unpublished thesis uni- unpublished thesis brigham young university versity of utah salt lake city 99 appp provoprove utah 79 appp 543 MCNULTY I1 B 1947 the ecology of bitterbitterbrushbrush 528 MARR J W AND D BUCKNER 1973 ecological in utah unpublished thesis university of utah analyses of potential shale oil products pipeline salt lake city corridors in colorado and utah unpublished re- 544 MEEUWIG R 0 AND S COOPER 1981 site quality port to colony development operation atlantic and growth of pinyon juniper stands in nevada richfleldrichfield company denver colorado by journal of forestry science 27 593 601 thorne ecological institute and the university of 545 MEEUWIG R 0 R B MURRAY P T TUELLER AND colorado boulder S V COOPER 1977 stratifying the pinyon juniper 529 MARR J W D L BUCKNER AND C MUTEL 1974 woodlands according to biotic potentials unpub- reaction of vegetation and soils to impact from lished report prepared for nevada agricultural construction and operation ofpipelinesofpipelines in western experiment station reno colorado and eastern utah unpublished report 546 MEINERS W R 1965 some geologic and edaphic prepared for colony development operation characteristics useful to management gramingprogramingprogrammingpro atlantic richfield company denver colorado within pinyon juniper type woodlands unpub- 138 appp lished thesis utah state university logan 530 MARSTON R B 1963 some comparative hydrologic 547 MEINKE R J 1975 A preliminary ecological and characteristics of aspen and mountain brush com- historical survey of north and south cainevilleCaineville munimunitiesties on steep mountain watersheds in north- mesas wayne county utah unpublished report ern utah unpublished dissertation utah state prepared for USDI bureau oflandeflandof land management university logan richfield utah 127 appp 531 MARTIN W E 1963 close in effects of an under- 548 MENZIES C W 1935 effects ofovergrazingof overgrazing on the ground nuclear detonation on vegetation I1 im- mortality of desert browse on the utah west mediate effects of eratcratcrateringcratencratenngeringng throwoutthrow out and blast desert unpublished thesis brigham young uni- project SEDAN USU S atomic energy commis- versityversity provo utah sionslon report PNE 228f 549 MERRIAMMERBIAM C H 1893 notes on the distribution of 532 MASON L R 1963 using historical records to de- trees and shrubs in the deserts and desert ranges termine climax vegetation journal of soil and of southern california southern nevada north- water conservation 18 190 194 western arizona and southwestern utah north 533 MASON L R 1971 yield and composition of utah s american fauna 7 285 394 range sites USDAUS DA soil conservation service 550 MEYER J C AND R D COMER 1985 reclamation portland oregon investigations on disturbed lands at glen canyon 534 MASON L R H A ANDREWS J A CARLEY AND national recreation area unpublished report E D HAACKE 1967 vegetation and soils of no prepared for USDI national park service by man s mesa relict area journal ofrange manage- thorne ecological institute boulder colorado ment 20 45 59 10 appp 228 P S BOURGERON ETALET AL volume 50

551 MEYER S E 1978 some factors governing plant western woodlands and range ecosystems sym- distributions in the mojave intermountain transi- posium 18 19 march 1976 USDA forest ser- tion zone great basin naturalist memoirs 2 vice utah agricultural experiment station lo- 197 207 gan utah 552 MILLER R F F A BRANSON I1 S MCQUEEN ANDCAND C 567 MUEGGLERMUEGGLEB W F 1985 vegetation associations T SNYDERSNYDEK 1982 water relations in soils as related pages 45 55 in debyle and winokur eds aspen to plant communities in ruby valley nevada ecology and management in the western united journal ofrange management 35 462 468 states USDA forest service general technical 553 MILLER W M 1888 the investigation oftheodtheofthe inter- report RM 119 rocky mountain forest and dependence ofplantofplantpiant life and climatic conditions of range experiment station fort collins colo- nevada nevada agricultural experiment station rado bulletin 2 reno nevada 568 MUEGGLER W F 1988 aspen community types of 554 MILTON NMN M ANDANDTT L PURDY 1983 plant and soil the intermountain region USDA forest service relationships in two hydrothermally altered areas general technical report INT 250 intermoun of the great basin great basin naturalist 43 taintam research station ogden utah 135 appp 457 469 569 MUEGGLER W F AND D C BARTOS 1977 grind- 555 MITCHELLMITCHCLL J E 1965 relationship between soil stone flat and big flat exclosures a 41 year record mineralogy and plant distribution within two com- ofchangesofchanges in clearcut aspen communities USDA munimunitiesties of the shadscaleshadscale zone in utah unpub- forest service research papelpaper INT 195 rocky lished thesis utah state university logan mountain forest and range experiment station 556 MITCHELL J E AND N E WEST 1965 effects ogden utah of physical soil propertiespropelproper ties on the distribution of 570 MUEGGLER W F AND R B CAMPBELL JR 1986 two shadscaleshad scale zone species in northern utah bul- aspen community types of utah USDA forest letin of the ecological society of america 46 46 service research paper INT 362 intermountain abstract forest and range experiment station ogden 557 MITCHELL J E N E WEST AND R W MILLER utah 1966 soil physical properties in relation to plant 571 MURDOCK J R 1951 alpine plant succession near community patterns in the shadscaleshad scale zone of mount emmons uinta mountains utah unpub- northwestern utah ecology 47 627 630 lished thesis brigham young university provo 558 MOONEY H H 1973 plant communities and vege- utah tation of the white mountains pages 7 17 in R 572 MURDOCKMUBDOCK J R &S L WELSH ANDANDBB W WOOD EDS M lloyd and R S mitchell A flora of the white 1975 navajo kaiparowitskaipaiowits environmental baseline mountains california and nevada university of studies 1971 1974 unpublished report prepared california press berkley for brigham young university center for health 559 MOONEYMOONLY H H G ST ANDRE AND R D WRIGHT and environmental studies provo utah 864 appp 1962 alpine and subalpine vegetation of the 573 NABI A A 1978 variation in successional status of white mountains of california american mid- pinyon juniper woodlands in the great basin un- land naturalist 68 257 273 published thesis utah state university logan 560 MOONEY M J 1985 A preliminary classification of 574 nachlinger J L 1985 the ecology of subalpine high elevation sagebrush grass plant vegetation in meadows in the lake tahoe region california and northern and central nevada unpublished thesis nevada unpublished thesis university of ne- university of nevada reno 118118ppappp vada reno 561 MOORE R R 1964 A study ofstandonstandofstand structure in the 575 NATIVE PLANTS incorporated 1983 J B king uneven aged stands in the engelmann spruce mine soil and vegetation data volume 1 section subalpine fir types on the utah state university 78319783 19 in mining and reclamation plan for J B forest unpublished dissertation utah state uni- king mine western states minerals corporation versityversity logan utah division of oil gas and mining number 562 MORDEN C J D brotherson AND B N SMITH act015002ACT 015002 salt lake city utah 1986 ecological differences of c3ca and c4ca plant 576 NEELY E E ANDANDMM E BARKWORTH 1984 vegeta- species from central utah in habitats and mineral tion on soils derived from dolomite and quartzite composition great basin naturalist 46 140 147 in the bear river range utah a comparative 563 MORETTIMORCTTI A C 1979 vegetation and soil factors in study bulletin of the torrey botanical club 3 relation to slope position a study ofplantof plantpiant commu- 179 192 nities on foothill knolls in the uinta basin of utah 577 NEESE E 1980 vegetation of the henry moun- unpublished thesis brigham young university tains pages 219 236 in proceedings UGA 1980 provo utah 31 appp henry mountains symposium 564 MORETTI M C ANDANDJJ D brotherson 1982 vege- 578 NEILSON R P 1986 on the interface between cur- tation and soil factors in i elationrelation to slope position rent ecological studies and the paleobotany of of foothill knolls in the uintaumtaomta basin of utah great pinyonpryonpmyon juniperjumper woodlands unpublished paper basin naturalist 42 81 90 presented to pinyon juniperjumper conference 13 16 565 MOZINGO H 1987 shrubs of the great basin a january 1986 USDA forest service reno ne- natural history university of nevada press las vada vegas 342 appp 579 NEILSON R P AND L H WULLSTEIN 1985 com- 566 MUEGGLER W F 1976 ecological role of fire in parativeparative drought physiology and biogeography of western woodland and range ecosystems pages quercus gamgamberigambehigambelttbelttbeitt and quercus turbturbinellamellameliameila ameri- 1 9 in proceedings use prescribedofofpiescnbed burning in can midland naturalist 114 259 272711 1990 NEVADA UTAH vegetation bibliography 229

580 NEILSON R P AND L H WULLSTEIN 1986 micro 595 NORTH AMERICAN EQUITIES NVN V 1984 vegetation habitat affinities of gambel oak seedlings great information volume 1 section 78319783 19 plus ex- basin naturalist 46 294 298 hibit 6 pages 1 22 in mining and reclamation 581 NELSON J V 1973 the environment oftheodtheof the vegeta- plan for blazon mine no 1 north american eq- tion of the upper sonoran and transition zones in uities NVN V utah division of oil gas and mining the uinta basin unpublished thesis university number act007021ACT 007021 salt lake city utah of utah salt lake city 106 appp 596 NORTHCUTT B E 1978 the plant ecology of butler 582 NELSON D L AND C F TIERNAN 1983 winter wash southeastern utah unpublished thesis injury of sagebrush and other wildlandwildwindlandland shrubs in university of coloradoofcolorado boulder 118 appp the western united states USDA forest service 597 NORTON B E 1978 the impact ofsheepof sheep grazing on research paper INT 314 intermountain forest longtermlong term successional trends in salt desert shrub and range experiment station ogden utah vegetation of southwest utah pages 610 613 inm 17 appp proceedings of the first international rangeland 583 NELSON L P AND M E JENSEN 1987 sagebrush congress society for range management den- grass community types of the humboldt national ver colorado forest unpublished report prepared for USDA 598 NYGREN L R 1963 summer habits and habitats of forest service humboldt national forest elko chukar partridge in northern utah unpublished nevada 80 appp thesis utah state university logan 53 appp 584 NELSON M G 1976 the montane coniferous 599 0 FARRELLOFARRELL M J 1973 seasonal activity patterns forests of the northern deep creek range utah population dynamics and spatial relationships of unpublished thesis university of utah salt lake rodents in a sagebrush community unpublished city 73 appp dissertation university of nevada reno 600 FARRELLOFARRELL T P ANDLAND L A EMERY 1976 ecology of 585 NELSON N F 1954 factors in the development 0 and restoration of waterfowlofwaterfowl habitat at ogden bay the nevada test site a narrative summary and refuge weber county utah utah state fish annotated bibliography university of nevada applied ecology and game department publication 6 salt lake and physiology center desert city utah 87 appp research institute report NVO 167 boulder city nevada 586 NELSON N F 1955 ecology of great salt lake 601 OLSEN P F 1974 wildlife resources oftheodtheofthe oil marshes proceedings of utah academy of sci- utah shale area utah of wildlife ence arts and letters 32 37 40 division research publication 74274 2 salt lake city utah 147 appp 587 NELSON R 1976 plants ofzion national park un- 602 OLSON B C 1975 utah big game range published report prepared for the inventory zion natural 1974 utah division of wildlife resources pub- history association Spspringdalespnngdale utah ringdale licationlication number 751475 14 salt lake city utah 588 NETTLETON W D B R BRASHER E L SPENCER 230 appp L N LANGAN ANDANDFF F PETERSON 1986 differen- 603 OLSON B C 1976 utah big game range inventory tiation of closely related xerolls that support dif- 1975 utah division of wildlife resources pub- ferent sagebrush plant in communities in nevada licationlication number 761076 10 salt lake city utah soil science society of america journal 50 128 appp 1277 1280 604 OLSON B C 1977 utah big game range inventory 589 NEWBERRY J S 1861 sage plain in J C ive ed 1976 utah division of wildlife resources publi- of report topographical engineers upon explo- cation number 77377 3 salt lake city utah 210 appp the colorado of rations of river the west house 605 OLSON B C ANDANDJJ R LOGAN 1973 utah big game part executive document 90 3 36th congress range inventory 1972 utah division of wildlife serial volume 1058 with ist session set maps resources publication number 73173 1 salt lake washington DCD C city utah 288 appp NICKENS P R 1977 590 wood rat knoll a multicom- 606 OLSON B C ANDANDJJ R LOGAN 1974 utah big game ponent site in butler wash southeastern utah range inventory 1973 utah division of wildlife unpublished report prepared for USDI bureau resources publication number 741674 16 salt lake of land management by the department of city utah 369 appp anthropology denver university denver 607 OOSTINGCOSTING H J 1956 the study of plant communi- colorado ties ad2d edition W H freenanbreenan and company 591 NIXON E S 1961 an ecological study of an exclo- san francisco 440 appp sure on mountain brush vegetation oftheodtheof the wasatch 608 OOSTINGCOSTING H J AND W D BILLINGS 1943 the red mountains utah unpublished thesis brigham fir forest of the sierra nevada abietum magmfimagnifimagnific provo young university 245 appp cae ecological monographs 13 259 274 592 NIXON E S 1967 A comparative study of the 609 ORR H K 1957 effects of plowing and seeding on mountain brush vegetation in utah great basin some forage production and hydrologic character- naturalist 27 59 66 istics ofaof a subalpine range in central utah USDA 593 NIXON E S 1977 A mountain Cercocercocarpuscarpus popu- forest service research paper 47 intermountain lation revisited great basin naturalist 37 97 99 forest and range experiment station ogden 594 NIXON E S AND E M christensen 1959 an eco- utah 23 appp logical study afanofanofan exclosure in the mountain brush 610 OSTLER W K AND K T HARPER 1978 floral ecol- vegetation of the wasatch mountains utah pro- ogy in relation to plant species diversity in the ceceedings of utah academy of science arts and wasatch mountains of utah and idaho ecology letters 36 182 183 abstract 5984859 848 861 230 PR S BOURGERONBOURGEBON ETALET AL volume 50

611 OSTLER W KKKK T HARPERhakperandkAND K B MCKNIGHT 626 PETERSONPETFRSON D L 1954 inventoryreinventoryremventorypreinventoryRe of surface soil 1981 the effects oflate lying snow on a subalpine and plant characteristics morris watershed un- danthonia deschampsia carex meadow pages published thesis university of utah salt lake 136 178 in K T harper W K ostler K B city 48 appp mcknight and D L hunter eds potential eco- 627 PETERSONPFTERSON P A 1984 flora and physiognomy of logical impact of snow pack augmentation in the the cottonwood mountains death valley na- uinta mountains of utah unpublished report tional monument california university of ne- prepared for USDI bureau ofreclamationoffleclamation den vada cooperative national park resources stud- ver colorado ies unit report 02206 las vegas nevada 612 OSTLER W K K T HARPER K B mcknightMCKNICHT AND 628 PETERSONPETEBSON S J ANDANDWW R KEAMMERER 1980 vege- D C ANDERSON 1982 the effects of increasing tation resources for the emery utah deep mine snowsnowpackpack on a subalpine meadow in the uinta project volume 6 chapter 9 sections 919.19gi 1 949.49 4 mountains utah journal of arctic and alpine pages 919 1 9249 24 plus appendices 919 1 929 2 in min- research 14 203 214 ing and reclamation plan for emery deep mine 613 PADGETTPADGETTWW GANDMG AND A E MANNING 1988 prelimi- consolidation coal company utah division of nary riparian community type classification for oil gas and mining number act015015ACT 015015 salt nevada draft unpublished report prepared for lake city utah USDA forest service region IV intermountain 629 PETERSONPETEBSON S R 1969 waterfowl ecology and uti- region ecology and classification program og- lizationlisailsalizailzation of uinta mountain water areas unpub- den utah lished thesis utah state university logan 614 PADGETT W G A P youngblood AND A H 58 appp WINWARD 1988 riparian community type classi- 630 PFISTER R D 1972 vegetation and soils in the ficationfi of utah USDA forest service inter- subalpine forests of utah unpublished disser- mountain region publication r4ra ECOL 88 01 tation washington state university pullman ogden utah 304 appp 98 appp 615 PAMMEL L H 1903 some ecological notes on the 631 PHILLIPS B G A M PHILLIPS III111 AND M A vegetation oftheofodthethe uinta mountains proceedings of SCHMIDT BERNZOTTBEBNZOTT 1987 annotated checklist of iowa academy of sciences 10 57 70 vascular plants of grand canyon national park 616 PAMMEL L H 1913 the grasses of the uinta grand canyon natural history association mono- mountains and adjacent regions proceedings of graph 7 flagstaff arizona iowa academy of science 20 133 149 632 PHILLIPS T A 1979 northnor th cedars pinyonpryonpmyon juniperjumper 617 PANIK H R 1976 the vertebrate structure of a studies USDA forest service range improve- pinyon juniper woodland community in north- ment notes november 1979 intermountain re- western nevada unpublished thesis university gion ogden utah 12 appp of nevadaofnevada reno 633 PICKFORD G D 1932 the influence of continuedofcontinued 618 PARRY C C 1875 botanical observations in south- heavy grazing and of promiscuous burning on ern utah in 1874 american naturalist 9 spring fall ranges in utah ecology 13 159 171 139 146 634 PISTER E P 1979 endangered species costs and 619 PASSEYHpassey H BANDVB AND V K HUGIE 1962 sagebrush on benefits great basin naturalist memoirs 3 relict ranges in the snake river plains and north- 151 158 ern great basin journal of range management 635 PLAIR T B 1934 vegetative cover of the utah 1527315.273 278 parks zion bryce nature notes 63 33 35 620 PASSEY H B AND V K huciehuclehugleHUGIE 1963 variation in 636 PLATEAU MINING COMPANY 1987 vegetation infor- bluebunchbluebunch wheatgrasswheatgrass in relation to environments mation volume 1 section 783 19 pages 783- and geographic location ecology 444415844.158158 161 68 783 gid plus tables 17 46 in mining and 621 PASSEY H B V K HUGIE E W WILLIAMS AND reclamation plan for star point mine plateau D E BALL 1982 relationships between soil mining company utah division of oil gas and plant community and climate on rangelands oftheofodthethe mining number act007006ACT 007006 salt lake city intermountain west USDA soil conservation utah service technical bulletin 1669 salt lake city 637 platouPLAIOU K A 1985 plant successional patterns on utah 123 appp seraiseral sagebrushgrasssagebrush grass ranges in northern nevada 622 PAVLIK B M 1985 sand dune flora of the great unpublished thesis university of nevada reno basin and mojave deserts of california nevada 638 PLUMMER A P 1958 restoration ofjuniper pinyon and oregon madronoM 3219732.19732 197 213 ranges in utah pages 207 211 in proceedings 623 PEDERSON J C AND K T HARPER 1979 chemical society of american foresters annual meeting composition of some important plants of south- 1958 salt lake city utah society of american eastern utah summer ranges related to mule deer foresters bethesda maryland reproduction great basin naturalist 3912239 122 128 639 PLUMMER A P D R christensen ANDANDSS B MON 624 pendletonbPENDLETONB K 1981 aspects oftheofodthethe floral biology SEN 1968 restoring big game range in utah utah of a subalpine meadow of the uinta mountains division of fish and game publication 68368 3 salt utah unpublished thesis brigham young uni- lake city utah 183 appp versityversity provo utah 54 appp 640 PLUMMER A P D R christensen R STEVENS 625 PERSKE G K 1960 the altitudinal zonation of AND K R JORGENSEN 1970 highlights results plants and animals on the eastern slope of the and accomplishments of game range restoration carson range unpublished thesis university of studies utah state division of fish and game nevada reno 127 appp publication 70370 3 salt lake city utah 94 appp 1990 NEVADA UTAH vegetation bibliography 231

641 POTTER L D ANDANDNN B PATTISONPATFISON 1976 shoreline ties to release from grazing pressure great basin ecology of lake powell institute of geophysics naturalist 46 148 156 and planetary physics lake powell restoration 658 RATLIFF J M 1972 remote sensing evaluations of project bulletin 26 235 appp phreatophytic vegetation in nevada unpublished 642 POTTER L D R C REYNOLDS JR ANDANDEE T LOUD thesis university of nevada reno ERBOUGH 1985a mancos shale and plant commu- 659 RAWLEY E V 1976 small islands of the great salt nity relationships analysis of shale soil and vege- lake utah department of natural resources tation transects journal of andaridarld environments 9 publication 6196 19 salt lake city utah 109 appp 147 165 660 REAM R D 1960 an ordination of the oak commu- 643 POTTER L D R C REYNOLDS JR ANDANDEE T LOUD nities of the wasatch mountains unpublished ERBOUGH 1985b mancos shale and plant commu- thesis university of utah salt lake city 52 appp nity relationships field observations journal of 661 REAM R D 1964 the vegetation of the wasatch andaridarld environments 9 137 145 mountains utah and idaho unpublished disser- 644 POWELL J W 1879 report on the lands of the and tation university of wisconsinofwisconsin madison 190 appp region of the united states with a more detailed 662 REESE G A 1981 diversity and production of account of the lands of utah ad2d ed UUSS govern- herbaceous vegetation in northern utah subalpine ment printing office washington D C 195 appp chronochronosequencesequence unpublished thesis utah state 645 PRATT 0 1975 orson pratt s journal 1849 1850 university logan E J watson publishing company salt lake city 663 REINER R L 1981 effects of grazing horses on a utah northern utah big game winter range unpub- 646 PRATT P P 1888 life and travels of parley P pratt lished thesis utah state university logan russel brothers chicago illinois 664 REMY J AND J BRENCHLEY 1861 A journey to 647 PREECE S J JR 1950 floristicfionFlonstiestic and ecological fea- great salt lake city with a sketch of history tures oftheodtheof the raft river mountains of northwesternofnorthwestern religion and customs of the cormonsmormons and an utah unpublished thesis university of utah introduction to the religious movement in the salt lake city united states vol I1 508 appp vol II11 605 appp 648 PRESLEY A R 1978 synoptic patterns and precipi- W jeffs publishing london tation in the pinyon juniper zone of the great 665 REVEAL J L 1944 single leaf pinyon and utah basin unpublished thesis university of nevada juniper woodlands of western nevada journal of reno 149 appp forestry 42 276 278 649 PRICE K P ANDANDJJ D brotherson 1987 habitat 666 REVEAL J L 1979a biogeography of the inter- and community relationships ofchnroseofcliffrose cowaniacoramaCocowamawania mountain region a speculative appraisal mexicansmexicanamexicana var stansburianastansburiana in central utah mentzelia 4 1 92 great basin naturalist 47 132 151 667 REVEAL J L 1979b biogeography of the inter- 650 PRIGGE B A 1975 flora of the clark mountain mountain region journal oftheodtheof the northern nevada range san Bernardbernardinomo county california un- native plant society 4 1 49 published thesis california state university los 668 REYNOLDS R V R 1911 grazing and floods a study angeles 105 appp of conditions in the manti national forest utah 651 PROCTOR D V 1971 A phytosociological study of USDA forest service bulletin 91 USU S govern- the north slopes of the uinta mountains unpub- ment printing office washington DC 16 appp lished report prepared for the USDA forest ser- 669 RICE B AND M WESTOBY 1978 vegetative re- vice wasatch national forest salt lake city sponses of some great basin shrub communities utah 506 appp protected against jack rabbits or domestic stock 652 PUDNEY R A 1972 food preferences of mule deer journal of range management 31 28 34 on the ruby butte study area unpublished the- 670 RICHAN F P 1957 the geography ofofpavantdavantpavant valley sis university of nevada reno utah unpublished thesis university of utah 653 RALSIONRALSTON G L 1969 plant zonation on dune sands salt lake city of washoe lake washoe county nevada un- 671 RICHMOND G M 1962 quaternary stratigraphy of published thesis university of nevada reno the la sal mountains utah USDI geological 654 RANGELAND RESOURCES international INC 1979 survey professional paper 324 USU S government final report for field search for rare plants in the printing office washington DCD C 135 appp reno nevada ES area unpublished report pre- 672 RICKARD W H 1959 crossgross vegetation pattern pared for bureau of land management reno within the nevada test site journal of the colo nevada 78 appp rado wyoming academy of science 4 32 655 RASMUSSEN D I1 1933 animal ecology of the 673 RICKARD W H 1963 vegetational analyses inm a kaibab national forest unpublished thesis uni- creosote bush community and the radioradioecologicecologic versity of illinoisofillinois urbana implications pages 39 44 in V schultz and 656 RASMUSSEN L L AND J D brotherson 1984 A W kelmenthelmentKelment eds radioecologyRadioecology reinhold habitat relationships ofsandsage arfemisiafilifoartemisia ftlifo publishing corp new york and AIBSA I1 B S lia in southern utah pages 58 66 in proceed- washington DCD C ings symposium on the biology of Artemisia and 674 RICKARD W H ANDJAND J C BEATLEY 1965 canopy chrysothamnus USDA forest service INT 200 coverage of the desert shrub vegetation mosaic of intermountain range and experiment station the nevada test site ecology 46 524 529 ogden utah 675 RICKARD W H AND J R MURDOCK 1963 soil 657 RASMUSSEN L L ANDANDJJ D brotherson 1986 re- moisture and temperature survey ofaof a desert vege- sponse ofwinterfat Ceratceratoidesoides lanata comminicommuni tation mosaic ecology 44 821 824 232 P S bourgeronetalBOURGERON ET AL volume 50

676 RICKARD W H AND L M SHIELDS 1963 an early site U S atomic energy commission report stage in the plant recolonization ofaof a nuclear target 1291612 916 washington DCD C area radiation botany 3 41 44 692 ROMNEY E M AND A WALLACE 1980 ecotonal 677 RIDD K 1978 vegetation of the uinta mountains distribution ofsaltof saltsait tolerant shrubs in the northern above 7000 feet unpublished map prepared for mojave desert great basin naturalist memoirs department of geographygeogiaphyGeogiaphy university of utah 4 132 138 salt lake city 693 ROMNEY E M A WALLACE AND R B HUNTER 678 RIPARIAN AREA management FIELD TASK FORCE 1980 the pulse hypothesis in the establishment of 1986 riverine riparian inventory and monitoring Atleartemisiamisiamisla seedlings at kahutepahute mesa nevada technical i eferenceefereferencedeferencerencerenee unpublished report prepared great basin naturalist memoirs 4 22 26 for USDI bureau of land management carson 694 ROMNEY E M A WALLACE H KAAZ AND V Q city nevada HALE 1980 the role ofshrubsofshrubs on redistribution of 679 ROBERTS R C 1950 chemical effects of salt toler mineral nutrients in soil inm the mojave desert ant shrubs on soils fourth international congress great basin naturalist memoirs 4 122 132 of soil science 1 404 406 695 ROUNDY B A AND J A YOUNG 1985 salt deserts 680 ROBERTSON J H 1947 response ofrangeoforrangerange grasses to of the great basin pages 39 49 in proceedings different intensities ofcompetitionofcompetition with sagebrush selected papers presented at the 38th annual artemisia tntritrldentatadentato nutt ecology 28 1 16 meeting of the society for range management 681 ROBERTSON J H 1965 range types and grazing 11 15 february 1985 salt lake city utah history ofnevadaofnevada pages 1 2 in proceedings oftheofodthethe 696 rushforthRUSHFOBTH S R L L ST CLAIR J D brotherson 18th annual meeting american society of range AND G NEBEKER 1982 ecology of lichen com- management 9 12 feblfebruaryFebi uary 1965 las vegas munimunitiesties in zion national park bryologist 85 nevada 185 192 682 ROBERTSON J H 1971 changes on a sagebrush 697 RYDBERG P A 1911 report ofaofa trip to southeastern grass range in nevada ungrazed for 30 years jour- utah in the summer of 1911 journal of the new nal of range management 24 397 400 york botanical gaidengardengalden 12143 237 253 683 ROBERTSON J H AND P B KENNEDY 1954 half 698 RYDBERG P A 1913 phytogeographical notes on century changes on northern nevada ranges jour- the rocky mountain region I1 alpine region bul- nal ofrangeoforrangerange management 71177 117 121 letin of the torrey botanical club 40 677 686 684 ROBERTSON R L 1976 ruffed grouse habitat pref- 699 RYDBERG P A 1914a phytogeographical notes on erences and effects of livestock grazing on habitat the rocky mountain region III111 formations in the utilization unpublished thesis utah state uni- alpine zone bulletin of the torrey botanical soci- versityversity logan 62 appp ety 41 459 474 685 ROBINETTEROBINETTC W L AND C ALDOUS 1964 zion na- 700 RYDBERG P A 1914b phytogeographical notes on tional park vegetation studies unpublished re- the rocky mountain region II11 origin oftheodtheofthe alpine port prepared for USDI fish and wildlife service flora bulletin of the torrey botanical society 41 Springdale utah 89 103 686 ROBOCKER W C 1957 some characteristics of soils 701 RYDBERG P A 1915a phytogeographical notes on and associated vegetation infested with haloge- the rocky mountain region V grasslands of the ton unpublished paper presented at the loth subalpine and montane zones bulletin oftheodtheofthe tor- annual meeting american society of range man- rey botanical club 42 629 643 agement great falls montana 29 january 702 RYDBERG P A 1915b phytogeographical notes on I1 february 1957 6pp6appappp the rocky mountain region IV forests of the 687 ROBOCKER W C 1958 some characteristics ofsoilsof soils subalpine and montane zones bulletin oftheodtheofthe tor- and associated vegetation infested with haloge- rey botanical club 42 11 25 ton journal ofrangeoforrangerange management 11 212 220 703 RYDBERG P A 1916a phytogeographical notes on 688 ROBOCKER W C 1961 the influence of perennial the rocky mountain region VI distribution oftheodtheofthe vegetation and precipitation on occurrence of subalpine plants bulletin of the torrey botanical halogeton glomeratusglomeratus and bromus tectoriumtectorumtectorum in club 43 343 364 andaridarld and semisemiaridarid regions journal of range man- 704 RYDBERG P A 1916b vegetative life zones of the agement 14 28 34 rocky mountain region memoirs oftheodtheofthe new york 689 ROGERS B S AND A TIEHM 1979 vascular plants botanical garden 6 477 499 oftleoftbeofthe sheldon national wildlife refuge with spe- 705 RYDBERG P A 1917 phytogeographical notes on cial reference to possible threatened and endan- the rocky mountain region VII formations in the gered species unpublished report prepared subalpine zone bulletin of the torrey botanical for USDI fish and wildlife service portland club 44 431 455 oregon 706 RYDBERG P A 1919 phytogeographical notes on 690 ROGERS G F 1982 then and now a photographic the rocky mountain region VIII distribution of history of vegetation change in the central great the montane plants bulletin oftheodtheof the torrey botani- basin desert university of utah press salt lake cal club 46 295 327 city 152 appp 707 RYDBERG P A 1920 phytogeographical notes on 691 ROMNEY E M V Q HALEHALF A WALLACE 0 R the rocky mountain region IX wooded forma- LUNTLUNF J D CHILDRESS H KAAZ G V ALEXANDER tions oftheodtheofthe montane zone oftheodtheof the southern rockies J E KINNEAR ANDTAND T L ACKERMAN 1973 some bulletin oftheofodthethe torrey botanical club 4744147 441 454 characteristics of soil and perennial vegetation in 708 RYDBERG P A 1921 phytogeographical notes on northern mojave desert areas of the nevada test the rocky mountain region X grasslands and 1990 NEVADA UTAH vegetation bibliography 233

other open formations of the montane zone of the 725 SHIELDS L M ANDANDPP V WELLS 1963 recovery of southern rockies bulletin oftheodtheofthe torrey botanical vegetation atomic target areas at the nevada test club 48 315 326 site pages 307 310 in radioecologyRadio ecology proceedings 709 SALISBURY F B 1952 plant soil relations on mate- oftheodtheof the first national symposium on radloradioecologyradioccologyRadioecology rial derived from hydrothermally altered rock in rinhold publishing corporation new york the marysvaleMarysvale region of utah unpublished the- 726 SHIELDS L M PR V WELLS AND W H RICKARD sis university of utah salt lake city 1963 vegetational recovery on atomic target areas 710 SALISBURY F B 1964 soil formation and vegetation in nevada ecology 44 697 705 on hydrothermally altered rock material in utah 727 SHREVESHBEVE F 1942 the desert vegetation of north ecology 45 1 9 america botanical review 8 195 246 711 SAMPSON A W 1925 foothill montane alpine the 728 SHUPEJSHUPE J BANDJB AND J D brotherson 1985 differen- flora and pages 1 its environment 24 31 in I tide tial effects of cattle and sheep grazing on high strom ed of and nevada U S na- flora utah US mountain meadows in the strawberry valley of tional herbarium contributions 25 washington central utah great basin naturalist 45 142 149 DCD C 665 appp 729 SHUPE J B J D AND S R RUSH- 712 SARGENT C S 1879 the forests of central nevada brotherson FORTH 1986 patterns of vegetationofvegetation surrounding american journal ofscienceofscience and arts 17 417 426 springs in goshen bay utah county 713 SAVAGE D E 1968 the relationship ofsageofossagesage grouse utah hydrobiologia 139 97 107 to upland meadows in nevada unpublished the USA 13997 730 SHUTE D AND N E WEST 1978 sis university of nevada reno shutedandn the application of ECOSYM classifications 714 SCHIER G A 1975 deterioration ofaspenofaspen clones in vegetation to rangelands the middle rocky mountains USDA forest ser- near price utah appendix reports 14 and 16 in vice research paper INT 710 intermountain J A henderson and L S davis eds forest and range experiment station ogden ECOSYM an ecosystem classification and data utah 14 appp storage system for natural resources management 715 SCHIMPF D J J A HENDERSON AND J A MAC unpublished report prepared for utah state uni- MAHON 1980 some aspects of succession in the versityversity logan 53 appp spruce fir forests of northern utah great basin 731 SIGLERJSIGLER J WANDJW AND J S TUHY EDS 1982 ecological naturalist 40 1 26 surveys altex oil leases southern utah octo- 716 SCHRAMM D R 1982 Floriflonsticsfloristiesfloristicssties and vegetation of ber 1981 september 1982 unpublished report the black mountains death valley national mon- prepared for altex oil by W F sigler & associ- ument unpublished report prepared for univer- ates inc logan utah sity of nevada las vegas 732 SIGLER J W AND J S TUHY 1983 resource sur- 717 SCHULTZ B 1987 ecology of curlleafcurlcurlleanleaf mountain veys tar sands triangle wayne and garfield mahogany Cercocercocarpuscarpus ledifoliusledifolius in western counties utah unpublished report prepared and central nevada population structure and dy- for santa fe energy company houston texas namics unpublished thesis university of ne- 506 appp vada reno 733 SILKER A R 1972 maintenance and survival of 718 SCHULTZ L M E E NEELY ANDANDJJ S TUHY 1987 vegetation on the sunrise campground cache flora of the orange cliffs of utah great basin national forest unpublished thesis utah state naturalist 47 287 298 university logan 41 appp 719 SCIENCE applications incorporated 1980 vege- 734 SIMMS P L 1967 the effect of stand density on tation survey at the knight mine sevier county characteristics ofwheatgrassesofwbeatgrasses unpublished dis- utah volume 1 section 78319783.19783 19 appendix 783 H sertationsertation utah state university logan in mining and reclamation plan for knight mine 735 SIMPSONSIMPSONJJ H 1859 report ofexplorationofexploration across the utah international incorporated utah division of great basin of the territory of utah for a direct oil gas and mining number act041005ACT 041005 salt wagon route from camp floyd to genoa in carson lake city utah valley USU S government printing office wash- 720 SHANTZ H L 1916 plant succession in tooelethoele val- ington DC ley utah pages 233 236 in F E clements 736 SINGH T 1967 quantitative analysis of perennial carnegie institute publication 242 washington atriplex dominated vegetation of southeastern DCD C utah unpublished dissertation utah state uni- 721 SHANTZ H L 1925 plant communities in utah and versityversity logan 178 appp nevada pages 15 23 in I1 tidestromTidwidestromestrom flora of 737 SINGH T AND N E WEST 1965 ordination of utah and nevada contributions of the USU S na- shadscaleshadscale zone vegetation of southeastern utah tional herbarium 25 washington DCD C 665 appp bulletin of the ecological society of america 46 722 SHANTZ H L AND R L PIEMEISEL 1940 types of 198 vegetation in escalante valley utah as indicators 738 SINGH T AND N E WEST 1971 comparison of of soil conditions USDA technical bulletin 713 some multivariate analyses of perennial atriplex washington DC 46 appp vegetation in southeastern utah vegetatiovegetation 23 723 SHEETER G R 1968 secondary succession and 289 313 range improvements after wildfire in northeastern 739 SKOUGARD M G 1976 vegetational response to nevada unpublished thesis university of ne- three environmental gradients in a salt playa near vada reno goshen utah county unpublished thesis brig- 724 SHIELDS L M AND P V WELLS 1962 effects of ham young university provo utah 75 appp nuclear testing on desert vegetation science 135 740 SKOUGARD M G AND J D brotherson 1979 38 40 vegetational response to three environmental 234 P S BOURGERONbourgeronetalET AL volume 50

gradients in the salt playa near goshen utah 755 STEIN B A AND S F WARRICK EDS 1979 granite county utah great basin naturalist 394439 44 58 mountains resource survey university ofcaliforofcalifor- 741 SKOUSEN J J N DAVIS AND J D brotherson nia environmental field program publication 1 1986 comparison of vegetationofvegetation patterns resulting santa cruz california 361 appp from bulldozing and two way chaining on a utah 756 STEVENS G L 1963 appraisal of the timber re- pinyon juniper big game range great basin natu- source potential oftheofodthethe dixie national forest un- ralist 46 508512508 512 published thesis utah state university logan 742 SMILEY F J 1915 alpine and subalpine vegetation 75pp75 appp of the lake tahoe region botanical gazette 59 757 STEVENS R 1983 description and discussion of 265 286 field tour sites pages 47 49 in proceedings ofthe 743 SMITH C B& 1986 grazing effects and site factors in first utah shrub ecology workshop utah state relation to grazing use of salt desert shrub desert university college of natural resources logan experimental range unpublished thesis utah 50soppappp state university logan 758 STEWART G 1935 plant cover and forage conditions 744 SMITH G L 1973 A flora of the tahoe basin and on spring fall and winter ranges largely on public neighboring areas wasman journal of biology 31 domain lands of the intermountain region utah 1 231 juniper 6 9 13 745 SOLDIER CREEK COAL COMPANY 1986 vegetation 759 STEWART G 1941 historic records bearing on agri- volume 2 part 3 section 373.7 pages 31613 161 31923 192 cultural and grazing ecology in utah journal of in mining and reclamation plan for soldier forestry 39 362 375 canyon mine soldier creek coal company 760 STEWARTSTEWABTGG 1948 utah s biological heritage and the utah division of oil gas and mining number need for its conservation proceedings of utah act007018ACT 007018 salt lake city utah academy of science arts and letters 25525.525 5 22 746 SOUTHARD ARA R 1958 some characteristics of fine 761 STEWART G W P COTTAM AND S S HUTCHINGS soil profiles under quaking aspen in cache na- 1940 influence of unrestricted grazing on north- tional forest unpublished thesis utah state ern salt desert plant associations in western utah university logan journal ofagriculturalofagricultural research 60 289 317 747 SPENCEJSPENCE J R& 1990 A preliminary review of various 762 STEWART G AND W KFLLERKELLER 1936 A correlation vegetation classification schemes for use on the method for ecology as exemplified by studies of colorado plateau with suggestions for future con- native desert vegetation ecology 1750017 500 514 sideration unpublished report prepared for capi- 763 STEWART G AND J WIDTSOE 1943 contribution of tol reef national park torrey utah 29 appp forest land resources to the settlement and devel- 748 ST JOHN C P 1970 establishment report for red opment oftheodtheofthe mormon occupied west journal of butte canyon research natural area within forestry 4163341.63341 633 640 wasatch national forest salt lake county utah 764 STODDARTstoddartlaLLAA 1941 the jalousepalouse grassland associ- unpublished report prepared for USDA forest ation in northern utah ecology 22 158 163 service washington DC 66ppappappp 765 STODDART L A 1945 range lands of utah county 749 ST CLAIR L L S R RUSHFORTH AND J D broth- utah and their utilization utah agricultural ex- erson 1986 the influence of mieromicromicrohabitatmicrobabitathabitat on perimentperiment station bulletin 317 utah state agricul- diversity distribution and abundance of corti tural college logan 32 appp colous lichens in zion national park utah and 766 STODDART LAL A A H holmgrenandcHOLM GREN AND C W COOK navajo national monument arizona Mycomycotaxontaxon 1949 important poisonous plants of utah utah 26253 262 agricultural experiment station special report 2 750 STAGER D W 1977 mule deer response to succes- utah state agricultural college logan 21 appp sional changes in the pinyon juniper vegetation 767 STODDART L A P B LISTER G STEWART T D type after wildfire unpublished thesis univer- PHINNEYPBINNEY AND L W DARSON 1938 range condi- sity of nevada reno tions in the uinta basin utah utah agricultural 751 STANSBURY H 1851 exploration and survey of the experiment station bulletin 283 utah state agri- valley oftheodtheof the great salt lake ofutahof utah senate doc- cultural college logan 34 appp ument volume 2 number 3 US government 768 STOKES W L 1977 subdivisions oftheodtheof the major phys- printing office washington DC ioiographic provinces in utah utah geology 4 752 STANTON W D 1933 the ecology and florlflorifloristicsfloristiessties of 1 17 the henry mountains proceedings of the utah 769 STUTZ H C 1951 an ecological study of a sphag- academy of science arts and letters 10 25 num lake in the subalpine forest of the uinta 753 STEELE P L 1958 the comparative mineral re- mountains of utah unpublished thesis brigham quirequirementsments of pioneer and climax range grasses young university provoprove utah 53 appp unpublished thesis bnghambrighambangham young university 770 STUTZ H C 1979 the meaning of rare and provo utah endangered in the evolution ofwesternofwestern shrubs 754 STEELE R R D PFISTER R A RYKER AND J A great basin naturalist memoirs 3 119 128 KITTAMS 1974 preliminary habitat types of the 771 summerfield H B JR 1969 pedological factors challis salmon and sawtooth national forests related to the occurrence ofbigosbigof big and low sagebrush unpublished cooperative habitat type classifica- in northern washoe county nevada unpub- tion report prepared for USDA forest service lished thesis university of nevada reno region IV and intermountain forest and range 772 SUMMIT COAL COMPANY 1987 vegetation re- experiment station ogden utah 7272ppappp sources volume 1 chapter 9 sections 919.1gi 939.3 1990 NEVADA UTAH vegetation bibliography 235

pages 1 6 plus appendix 919 1 in mining and recla- service manti lasal national forest moab mation plan for boyer mine summit coal com- utah pany utah division offilofoilof oiloll011 gas and mining num- 789 THORNE R F 1982 the desert and other transmon- ber act043008ACT 043008 salt lake city utah tane plant communities of southern california 773 TANNER V M 1940a A chapter on the natural his- aliso 10 219 257 tory of the great basin 1800 to 1855 great basin 790 THORNE R F B A PRIGGE AND J henrickson naturalist 1 33 61 1981 flora of the higher ranges and the kelso 774 TANNER V M 1940b A biotic study of the kai dunes oftheodtheof the eastern mojave desert in california barowitzparowitzparowitz region of utah great basin naturalist 1 aliso 10 71 186 97 126 791 TISDALE E W AND M HIRONAKA 1981 the sage 775 TANNER V M ANDANDCC L HAYWARD 1934 A biologi- brush grass region a review oftheodtheofthe ecological liter- cal study oftheodtheofthe la sal mountains utah report 1 ature university of idaho forest wildlife and utah academy of sciences arts and letters 11 range experiment station bulletin 33 moscow 209 235 idaho 776 TAUSCH R J 1973 plant succession and mule deer 792 TOWER J D 1970 vegetation stagnation in three utilization on pinyonpryonpmyon juniper chamingschainings in nevada phase big game exclosures and its effects in deter- unpublished thesis university ofnevada reno mining forage utilization unpublished thesis 777 TAUSCH R J 1980 allometric analysis of plant university of nevada reno growth in woodland communities unpublished 793 TOWNSEND T W 1966 plant characteristics re- dissertation utah state university logan lating to the desirability of rehabilitating the 778 TAUSCH R J ANDANDPP T TUELLER 1977 plant succes arctostaphylos patula ceanothus velutinus sionslon following chaining of pinyon juniper wood- ceanothus proprostratusstratus association on the east slope lands in eastern nevada journal of range man- oftheodtheofthe sierra nevada unpublished thesis univer- agement 304430 44 49 sity ofnevada reno 779 TAUSCH R J AND P T TUELLER 1989 evaluation 794 TRUEBLOOD R W 1954 the effect of grass reseed of pinyon sapwood to phytomassphytomass relationships ingmg in sagebrush lands on sage grouse populations over different site conditions journal of range unpublished thesis utah state agricultural col- management 42 209 212 lege logan 77 appp 780 TAUSCH R J N E WEST AND AAA A NABI 1981 tree 795 TUELLER P T 1973 secondary succession discli- age and dominance patterns in great basin max and range condition standards in desert pinyon juniper woodlands journal ofrangeoforrangerange man- shrub vegetation pages 57 65 in D N cydershyders agement 34 259 264 ed andaridarld shrublandsShrublands proceedings of the third 781 TAUSCH R J AND N E WESTWFST 1985 patterns of workshop of the usaustraliaUSU S australiaaustralla rangelands pinyon and juniper establishment on a southwest- panel 26 march 5 april 1973 tucson arizona ern utah site abstracts oftheodtheof the annual meetings of 796 TUELLER P T 1975 the natural vegetation of ne- the society for range management 78 vada low deserts high mountains provide con- 782 TAUSCH R J AND N E WEST 1986 morphological trasts from shadshadscalescale shrub to ancient bristle variationprecipitationvariation precipitation relationships of great cones mentzeliaMentzeha 1 3 6 23 28 basin single needled pinyon pages 86 91 in 797 TUELLER P T 1985 sagebrush dominated vegeta- pinyon juniper conference proceedings USDAUS DA tion of the great basin pages 24 30 in proceed- forest service general technical report INT ings 38th annual meeting of the society for range 215 intermountain forest and range experiment management 11 15 february 1985 salt lake station ogden utah 91 appp city utah 783 TAUSCH R J AND N E weestWEST 1988 differential 798 TUELLERTUELLEB P T C D BEESONBEFSON R J TAUSCH N E establishment ofofpinyonpinyon andjuniper following fifefirefnan e WEST ANDANDKK H REA 1979 pinyon juniper wood- american midland naturalist 119174ilg119 174 184 lands of the great basin distribution flora and 784 TECHNICAL COMMITTEE 1971 vegetation of the vegetal cover USDA forest service research pa- lake tahoe region A guide for planning unpub- per INT 229 intermountain forest and range lished report prepared for the tahoe regional experiment station ogden utah 22 appp planning agency and USDA forest service plan- 799 TUELLER P T ANDANDWW H BLACKBURN 1974 condi- ning team south lake tahoe california 43 appp tion and trend of the big sagebrushneedle 785 TEPEDINO V J 1979 the importance of bees and andthreadand thread habitat type in nevada journal of other insect pollinatorspollinators in maintaining floral spe- range management 27 36 40 cies composition great basinbaemba&m naturalist memoirs 800 TUELLER P T ANDJAND J E CLARK 1975 autautecologyAntecology 31393 139 150 of pinyon juniper species of the great basin and 786 TEW R K J T LOTT ANDANDTT M BLISS 1988 ecosys- colorado plateau pages 27 39 in G F gifford tem stratification on the fishlikefishlakeFishlake national forest and F E busby eds the pinyon juniperjumper unpublished report prepared for USDA forest ecosystem A symposium utah state university service fishlikefishlakeFishlake national forest richfield utah logan 787 THATCHER A P AND V L HART 1974 spy mesa 801 TUELLER P T AND R E ECKERT JR 1987 big yields better understanding of pinyon juniper in sagebrush artemisia dentatatntritridentatetndentatatridentata vastyanavaseyanavaseyana and range ecosystems jouljournalnalnai of range management longleaf snowberry symphoricarpos oreoreophilusophilus 2735427 354 357 plant associations in northeastern nevada great 788 THOMPSON R 1979 life zones plant associations basin naturalist 47 117 131 and habitat types fenonferronpenonperron price planning units 802 TUELLERtuellTUELI EK P T D H HEINZE AND R E ECKERT unpublished report prepared for USDA forest 1966 A tentative list of existing nevada plant 236 P S BOURGERONbourgeronetalET AL volume 50

communities a third approximationappi oximation unpub- report 15 pages 102 108 and 199 203 in vol I1 lished report prepared for university of nevada energy mineral rehabilitation inventory and anal- department of range wildlife and forestry ysis in the mountain coal fields ofGarneldGarfield county reno 14 appp utah unpublished report prepared for USDI bu- 803 TUELLERTULLLER P T ANDJAND J D TOWFRTOWER 1979 vegetation reau of land management by utah state univer- stagnation in three phase big game exclosures sity logan 203 appp journal of range management 32 250 263 816 VALE T R 1973 the sagebrush landscape of the 804 tuhytuny J S ANDANDJJ A MACMAHON 1988 vegetation intermountain west unpublished dissertation and relict communities of glen canyon national university ofcalifornia berkeley 508 appp recreation area unpublished final report pre- 817 VALE T RK 1975 presettlement vegetation in the pared by state for USDI national sagebrushsagebi ush grass area of the intermountain west utah university range park service rocky mountain region lakewood journal of management 28 32 36 colorado 299 appp 818 VALENTINE J F 1961 important utah range grasses state university 805 USDA 1871 great salt lake pages 559 569 utah extension circular the 281 logan utah in 1870 report oftheodtheofthe commission on agricultureaghAgnculture in 819 VALLEY CAMP OF UTAH incorporated 1984 vege- unpublished report prepared for USDA wash- volume 6 78319783.19783 19 pages 15d D C tation section ington DC 15n32 in mining and reclamation plan for belinabelmabeimabellna 806 USDA SOIL conservation SERVICE 1981 land re- mines complex valley camp of utah incorpo- and land areas source regions majormaior resource rated utah divisondavison ofoilofoiloll011 gas and mining num- states offil of the united USDA soil conservation ber act007001ACT 007001 salt lake city utah service agriculture handbook 296 washington 820 VALLEY engineering incorporated 1988 vege- DCD C 156 appp tation and terrestrial wildlife report volume 3 SERVICE 807 USDA SOIL conservationconslrvation 1988a major chapter 9 item 919 1 pages 1 38 in mining and land resource area 28a great salt lake area reclamation plan for crandall canyon mine nevada site descriptions unpublished internal gendalgenwal coal company utah division of oil gas technical guide section HE reno nevada and mining number act015032ACT 015032 salt lake 808 USDA SOIL conservation SERVICE 1988b major city utah land resource area 28b central nevada basin and 821 VAN BALENBALFN J A 1973 plant covelcover types of the range nevada site descriptions unpublished upper bear river drainagediadramage utah unpublished internal technical guide section IIB reno thesis university of utah salt lake city nevada 822 VAN PELT N S 1978 woodland parks inm southeast- 809 USDI BUREAU OF LAND management 1972 his- ern utah unpublished thesis university ofutahofutah tory of range fires in the uinta basin utah un- salt lake city published report 823 VAN PEITPELT N S 1988 baseline description of a na- 810 USDI NATIONAL PARK SERVICE 1974 wilderness tive sagebrushgrasssagebrush grass reference area at wildcat recommendation lake mead national recreation leadoutloadout volume 1 appendix I1 pages 1 5 in area arizonaanzona nevada unpublished report pre- mining and reclamation plan for wildcat leadoutloadout pared for USDI national park service denver facility andalexandalea resources utah division offilofoilofoil serviceseisel vice center denver colorado 80 appp gas and mining number act007033ACT 007033 salt lake 811 USU S fueiFULIFUEL COMPANY 1980 vegetation volume 3 city utah chapter 9 sections 9199.1gi 1 939.39 3 pages IX 1 IX 76 in 824 VAN PELT N S RB STEVENS AND N E WEST 1989 mining and reclamation plan for hiawatha mines comparative survival and growth of immature complex USU S fuel company utah division of utah juniper after chaining in central utah ab- oil gas and mining number act007011ACT 007011 salt stracts of the annual meeting of the society for lake city utah range management 287 812 utaiiutainUT AH DIVISION OF wllwilWILDLIFEDI IFE RESOURCES AND ECOI 825 VEST E D 1962a biotic communities in the great OGY consultants INC 1977 annotated bibliog- salt lake desert university of utah institute of raphy of natural resource informationmfoimafoi mation southern environmental biological research ecology and utah USDI fish and wildlife service biological Epiepizoologyzoology series 73 salt lake city utah services program FWSOBS 7734 fort collins 122 appp colorado 265 appp 826 VEST E D 1962b the plant communities and asso- 813 utaiiutainUTAH environmentenvironmental AI AND agricultural CONSUL ciated fauna of dugway valley inm western utah TANTS 1973 environmental setting impact miti- unpublished thesis university ofutahof utah salt lake gation and recommendations for a proposed oil city products pipeline between lisbon valley utah 827 WAGNER W W 1973 cesium 137 accumulation and parachute creek Coloiacoloradodo unpublished re and distribution inm plants and soils of an alpine port prepared for colony development operat- community of utah unpublished dissertation ions denver colorado university of utah salt lake city 267 appp 814 UTAIIUTAH international incorporated 1987 legetavegeta 828 WAKEFIELD H 1933 A study oftheodtheofthe plant ecology of tion volume 14 section 6 pages 616 1 6536 53 in salt lake and utah valleys before the mormon mining and reclamation plan for alton mine immigration unpublished thesis brigham young utah international incorporated utah division of university provo utah 54 appp oil gas and mining number act025003ACT 025003 salt 829 WAKIFIEIDWAKEFIELDWAKI fieldFIEID H 1936 A study of the native vegeta- lake city utah tion of salt lake and utah valleys as determined 815 UTAH STATESTATL university 1978 rehabilitation po- by historical evidence proceedings of utah tential foiroiforror the henry mountain coal field emblaEMRIAEMBIA academy of science aitsaltsarts and letters 13 11 18 1990 NEVADA UTAH vegetation bibliography 237

830 walkkrWALKERWALKEH G R AND J D brothersonBROThersonHEBSON 1982 habi- 847 WELSH S L 1979 endangered and threatened tat relationships ofbasinofbasin wildrye in the high moun- plants of utah a case study great basin naturalist tain valleys ofcentralofcentral utah journal ofrangeoforrangerange man- memoirs 3 64 80 agement 35 628 633 848 WELSH S L N D ATWOOD L C HIGCINSHIGGINS AND S 831 WALLACE A AND E M ROMNEY 1980 the role of GOODRICH 1987 A utah flora great basin natu- pioneer species in revegetation of disturbed ralist memoirsmemon s 9 894 appp great desert areas basin naturalist memoirs 4 849 WELSH S L AND E M christensen 1957 an 31 33 ecological study of the vegetation ofthe dinosaur 832 WALLACE A E M ROMNEY AND R B HUNTER national monument proceedings of the utah 1980 the challenge of a desert revegetation of academy of science arts and letters 34 55 56 disturbed desert lands great basin naturalist 850 WELSH S L ANDANDRR J KASS 1985 vegetation moni- memoirs 42164 216 225 toring of reclaimed slopes and corresponding ref- 833 WALLIS A 1982 mixed C an oveloverviewview oftheodtheofthe grass- erence areas at the skyline mine 1985 appendix lands of america pages 195 208 A C north in volume a2aa pages 1 29 mining and reclama- nicholson A mclean and T E baker eds in tion plan for skyline mine company proceedings oftheodtheofthe grassland ecology and classifi- utah fuel utah division of oil gas and mining number cation symposium 2 4 june 1982 kamloopsKamloops ACT 007005 salt british columbia province of british columbia act007005 lake city utah 851 WELSH S ministry of forests victoria L AND J R MURDOCKMUBDOCK 1980 report of plant analysis 834 WARD K V 1977 two year vegetation response vegetation community threatened and endangered plant soils and reclama- and successional trends iolloifoilorfor spring burns in the species plans energy company pinyon juniper woodland unpublished thesis tion for coastal states university ofnevada reno mckinnon properties skyline project carbon emery 835 WARNER J H ANDANDKK T HARPER 1972 understory counties utah appendix volume a2aa 1 plan characteristics related to site quality for aspen in pages 135 in mining and reclamation for utah brigham young university science bul- skyline mine utah fuel company utah divi- gas letin biological series 16 1 20 sionslon of oil and mining number act007ACT 007 836 WATSON S 1871 botany in C king ed report 005 salt lake city utah of the geological exploration of the fortieth paral- 852 WELSH S L annANDANDJJ R MURDOCK 1982 report of lel volume 5 USU S army corps of engineers revegetation success and vegetation monitoring of professional paper 18 525 appp reclaimed and lefereneeferencereference areas at skyline mine 837 WEBB G M ANDanujANDJJ D brotherson 1988 eleva- during the 1982 field season appendix volume tional changes in woody vegetation along three a2aa pages 1 27 plus appendices I1 II11 illIII111 in min- streams in washington county utah great basin ing and reclamation plan for skyline mine utah naturalist 48 512 529 fuel company utah division of oil gas and 838 WEBB R H J W STEIGER ANDANDRR M TURNER 1987 mining number act007005ACT 007005 salt lake city dynamics of mojave desert shrub assemblages in utah the panamint mountains california ecology 68 853 WEISHWELSH S L J R MURDOCK L JUARROS AND 478 490 E NEESE 1981 report of studies of vegetation 839 WEHRLIE LLJJ 1973 vegetation analysis of a sierra and soils for coastal states energy company con- nevada montane coniferous forest unpublished vulsion canyon mine ofsouthernofsouthern utah fuel com- thesis university of nevada reno pany SUFCO sevier county utah volume 5 840 WEIR G H 1976 palynology flora and vegetation pages 1 11 and 39 78 in mining and reclamation of hovenweepHovenweep national monument implications plan for convulsion canyon mine southern utah for aboriginal plant use on cajon mesa colorado fuel company utah division of oil gas and and utah botany 3749 B abstract mining number act041002ACT 041002 salt lake city 841 WELLS P V 1960 physiognomic intergradation of utah vegetation on the pine valley mountains in south- 854 WILSHWELSH S L ANDCAND C A TOFT 1981 biotic commu- western utah ecology 41 553 556 nities of hanging gardens in southeastern utah 842 WELLS P V 1961 succession in desert vegetation national geographic society research reports 13 on streets of a nevada ghost town science 134 663 681 670 671 855 WESTWFST N E 1966a ecology and management ofsaltofsaltsait 843 WELLS P V 1983 paleobiogeographyPalcopaleobiogeography of montane desert shrub ranges a bibliography wyoming islands in the great basin since the last glacioplu range management 228 147 167 revised and vial ecological monographs 53 341 382 reprinted 1968 utah agricultural experiment 844 WELLS P V ANDannlANDLL M SHIELDS 1964 distribution station mimeograph series 505 30 appp of larrea didaridivaridivaricatedtvancatadivaricatadtvancatacata in relation to a temperature 856 WEST N E 1966b research on the ecology of the inversion at yucca flat southern nevada south- utah salt desert areas pages 160 169 in proceed- western naturalist 9 51 55 ings of the salt desert shrub symposium USDI 845 WELSH S L 1957 an ecological survey oftheodtheofthe vege- bureau of land management cedar city utah tation ofoftheodthethe dinosaur national monument utah 857 WEST N E 1971 pinyon juniper ranges unpub- unpublished thesis brigham young university lished report prepared for utah state university provo 86 appp department of natural resources logan 28 appp 846 WELSH S L 1970 vegetation of the colorado 858 WESTWFST N E 1974 the shrublandsshrub lands of utah utah plateau its phytogeographic distinction from the science 35 4 6 great basin unpublished manuscript prepared 859 WEST N E 1977 an ecologistecologistss thoughts toward for brigham young university provo utah 29 appp determining the recreational carrying capacity of 238 PR S BOURGERONbourgeronetalET AL volume 50

white water sections of the upper colorado river can terrestrial vegetation cambridge university drainage pages 125 145 in L E royer W H press new york becker and R schreyer eds managing colo- 872 WEST N E 1989 vegetation types of utah pages rado river whitewater the carrying capacity strat- 18 51 in K johnson ed rangelands of utah egy institute for the study ofoutdoorofoutdoor recreation utah state university press logan and tourism utah state university logan 873 WEST N E AND M M BAASHER 1968 determina- 860 WEST N E 1978a basic syn ecological relationships tion of adequate plot size by use of mean distance of sagebrush dominated lands in the great basin between salt desert shrubs southwestern natu- and colorado plateau pages 33 41 in G F gif- ralist 136113 61 74 ford and E busby eds F the sagebrush eco- 874 WEST N E AND J E BOWNS 1988 shrubland system a symposium utah state university col- grassland shrubsteppeshrubsteppe and woodland vegetation lege ofnaturalofnatural resources logan 251 appp of the northern mohave desert southwestern 861 WEST N E 1978b the changing woodlands of the utah and northwestern arizona pages 73 81 in great basin utah state university college of vankat ed resourcespeo J L vegetation of the southwest- natural resources edge natural ern united states A handbook for the 1988 excur- pie 1 13 16 sion oftheodtheofthe international association for vegetation 862 WEST N 1979 survival of E patterns major peren- science special publication 1 north american nials in salt deseideseldesert t shrub communities of south- section international association for vegetation western utah journal of range management 32 science 442 445 875 WEST N E D CAIN AND G GIFFORD 1973 biol- 863 WEST N 1982 char- E approaches to syn ecological ogy ecology and renewable resource manage- of wildlandswildlands the acterization in intermountain ment of the pigmy conifer woodlands of western pages 633 641 C ed west inttntin T brann in place north america a bibliography utah agricultural principles A resource inventories and practices experiment station research report 12 logan national workshop university of maine orono utah 36 appp society ofamericanamerleanofamerican foresters 9 14 august 1981 876 WEST N E AND M M CALDWELL 1983 snow mcclean virginia as a factor in salt desert shrub vegetation patterns in 864 WEST N E 1983a great colorado plateau basin curlew valley utah american midland natural- sagebrush semidesertdesert pages 331 349 N E semi in ist 109376109log 376 379 west ed temperate deserts and semi deserts 877 WEST N E G B COLTHARP WORKMANWOBKMAN ecosystems oftheofodthethe world vol 5 elsevier publish- J P R W WEINVWEIN V B HANCOCKHANCOCKKK SIMONSON AND KEITH ing company amsterdam slmonsonandjJ 1973 vegetational micro climate edaphic hy- 865 WEST N E 1983b western intermountainintel mountain sage- microclimate drodrologiclogic and economic effects of contour furrows brush steppe pages 351 374 in N E west ed temperate deserts and deserts ecosystems and gully plugs on frail lands near cisco utah semi utah agricultural experiment of the world vol 5 elsevier publishing com- station technical pany amsterdam bulletin manuscript logan utah 878 WEST GASTO 866 WEST N E 1983c intermountain salt desert shrub N E AND J 1978 phenology of the aerial of shadscaleshadscale and fat lands pages 375397375 397 in N E west ed temper- portions winterwinterfat in curlew valley journal of range manage- ate deserts and semi deserts ecosystems of utah the world vol 5 elsevier publishing company ment 31 43 45 amsterdam 879 WEST N E AND K I1 IBRAHIM 1968 soil legetavegeta 867 WEST N E 1983d colorado plateau moravianmohavian tion relationships in the shadshadscalescale zone of south- blackbrushblackbrush semisemidesertdesert pages 399 412 in N E eastern utah ecology 49 445 456 west ed temperate deserts and semi deserts 880 WESTNWEST N E ANDANDWW L LOOPE 1980 occurrence of ecosystems oftheodtheofthe world vol 5 elsevier publish- wildfireswildfires in zion national park in relation to ing company amsterdam ecosystem and record taking variables pages 868 WEST N E 1983e southeastern utah galleta 21 28 in proceedings of second conference on three awn shrub steppe pages 413 421 in N E scientific research in the national parks vol X west ed temperate deserts and semi deserts fire ecology USDA national park service transa- ecosystems oftheodtheofthe world vol 5 elsevier publish- ctions and proceedings series 6 washington DC ing company amsterdam 881 WEST N E AND M 11 MADANY 1981 fire history 869 WEST N E 1984 successional patterns and pro- oftheofodthethe horse pasture plateau zion national park ductivity potentials ofpmyonof pinyon juniper woodlands university of utah national park service cooper- pages 1301 1332 in natural resources council ative studies unit final report 225 appp national academy of sciences developing strate- 882 WEST N E R T MOORE K A VALENTINE L W gies for rangeland management westviewWestview press LAW P R OGDENOCDEN F C PINKNEY P T TUELLER boulder colorado AND A A BEETLE 1972 galleta taxonomy ecol- 870 WEST N E 1987 human impacts on vegetation of ogy and management of hilariahilatiahipatia jamesiijamesie on west- the semisemiaridandaridarld temperate intermountain region of ern rangelands utah agricultural experiment western north america page 398 in abstract of station bulletin 487 logan utah the XIV international botanical congress berlin 883 WEST N E F D PROVENZA P S JOHNSON AND west germany M K OWENS 1984 vegetation change after 13 871 WEST N E& 1988 intermountain deserts shrub years of livestock grazing exclusion on sagebrush steppes and woodlands pages 209 230 in M G semidesert in west central utah journal ofrange barbour and W D billings eds north ameriamerlamen management 37 262 264 199011990 NEVADA UTAH vegetation bibliography 239

884 WEST N E K H REA AND R J TAUSCH 1975 896 WILCOX R B 1977 canopy influence as a factor in basic ecologicalsynsynecologicalsynechological relationships in juniper determining understory community composition pinyon woodlands pages 41 53 in G F gifford unpublished thesis brigham young university and F E busby eds proceedings oftheodtheofthe pinyon provo utah 42 appp juniper ecosystem symposium utah agricul- 897 WILCOX R B J D brotherson AND W E tural experiment station logan EVENSON 1981 canopy influence on understory 885 WEST N E R J TAUSCH AND A A NABI 1979 community composition northwest science 55 patterns and rates ofofpmyonpinyon juniper invasion and 194 201 degree of suppression of understory vegetation in 898 WILLIAMS G 1969 analysis ofhydrologic edaphic the great basin USDAUS DA forest service inter- and vegetative factors affecting infiltration and mountain region range improvement notes og- erosion on certain treated and untreated pinyon den utah 14 appp juniper sites unpublished dissertation utah 886 WEST N E R J TAUSCH K PI REA AND A R state university logan SOUTHARD 1978a soils associated with pinyon 899 WILLIAMS J E D BOWMAN J BROOKS A juniper woodlands of the great basin pages ECHELLE R EDWARDS D A hendrickson AND 68 88 in C T youngberg ed forest soils and J J LANDYE 1985 endangered aquatic ecosys- land use proceedings oftheodtheofthe fifth north american tems in north american deserts with a list of forest and soils conference colorado state uni- vanishing fishes of the region journal of arizona versityversity fort collins colorado nevada academy of science 20 1 62 887 WEST N E R J TAUSCH K H REA AND P T 900 WILSON R 0 1985 aerial and near surface spectral TUELLER 1978b taxonomic determination dis- characteristics ofeightofeight nevada rangeland commu- tributiontribution and ecological indicator values of sage- nities unpublished thesis university of nevada brush within the pinyon juniper woodlands of the reno great basin journal of range management 31 901 WINDELLWlNDELL J T B E wilWlLWILLARDLARD D J COOPER S Q 87 92 FOSTER C F knudhansenKNUD HANSEN L P RINK AND G N 888 WEST N E R J TAUSCH K H REA AND P T KILADIS 1986 an ecological characterization of TUELLER 1979 phytogeographical variation with- rocky mountain montane and subalpine wet- in juniper pinyon woodlands of the great basin lands USDI fish and wildlife service biological in K T harper and J L reveal coords inter- report 8611 298 appp mountain biogeography a symposium great 902 WINN D S 1976 terrestrial vertebrate fauna and basin naturalist memoirs 2 119136ilg119 136 selected coniferous forest habitat types on the 889 WEST N E AND P T TUELLER 1972 special north slope of the uinta mountains unpublished approaches to studies of competition and succes- report prepared for USDA forest service sion in shrub communities pages 165 171 in wasatch national forest ogden utah 145 appp C M mckell J P blaisdell and J R goodin 903 WOOD B W 1966 an ecological life history of eds wildlandWildwindlandland shrubs their biology and utiliza- budsagebuddage in western utah unpublished thesis tion USDA forest service general technical re- brigham young university provo utah 85 appp port INT 1 intermountain forest and range ex- 904 WOODWOODBB WANDJW AND J D brotherson 1984 ecologi- perimentperiment station ogden utah cal adaptation and grazing response of budsagebuddage 890 WESTWESTNN EEANDNAND N S VAN PELT 1987 successional artemisia spinescentspinescensspinescens in southwestern utah patterns in pinyon juniper woodlands pages pages 75 92 in proceedings symposium on the 43 52 in R L everett compiler proceedings biology ofartemisiaof artemisia and chrysothamnus USDA pinyon juniper conference USDA forest ser- forest service general technical report INT vice general technical report INT 215 inter- 200 intermountain research station ogden mountain research station ogden utah utah 891 WHEELER N C AND P curlesGURIESCURIES 1982 biogeogra- 905 WOOD B W S L WELSH AND J R MURDOCK phy of lodgepole pine canadian journal ofbotany 1975 the vegetation of the kaiparowits region 601805 1814 an overview pages 249249302302 in J R murdock 892 WHIPPLEWHIFFLE A W 1854 report of exploration for a S L welsh and B W wood eds navajo railway route near the 35th parallel of latitude kaiparowits environmental baseline studies from the mississippi river to the pacific ocean 1971 1974 unpublished report prepared for 37th congress and2nd session house document brigham young university center for health and 129 volume 4 washington DC 154154ppappp environmental studies provo utah 864 appp 893 WHITE J A 1971 climate plant soil and water 906 WOODBURY A M 1933 biotic relationship ofofzionzion relationships to a wildlife plan unpublished the- canyon utah with special reference to succes- sis university ofnevada reno sion ecological monographs 3 148 245 894 WHITE W N 1932 A method ofestimatingofestimating ground- 907 WOODBURY A M 1938 the lower sonoran in water supplies based on discharge by plants and southwestern utah science 87 484 485 evaporation from soil results of investigations in 908 WOODBURY A M 1947 distribution of pigmy escalante valley utah USDI geological survey conifers in utah and northeastern arizona ecol- water supply paper 659a denver colorado ogy 281139811328 113 126 105 appp 909 WOODBURY A M 1956 ecological checklists 895 wieslander A E 1932 vegetation type maps great salt lake desert series unpublished re- of california and western nevada university of port prepared for university of utah salt lake california press berkeley city 240 P S BOURGERONbourgeronetalET AL volume 50

910 WOODBURY A M S D DURRANT AND S FLOWERS developing strategies for rangeland management 1959 survey of vegetation in the glen canyon national academy of science westviewWestview press reservoir basin university ofutahanthropologiof utah anthropologi- boulder colorado cal papers 36 salt lake city utah 53 appp 926 YOUNG J A R A EVANS B A ROUNDY AND J A 911 WOODBURY A M S D DURRANT S FLOWERS bbownBROWNBKOWN 1986 dynamic landformslandforms and plant com- 1960 A survey of the vegetation in the flaming munimunitiesties in a pluvial lake basin great basin natu- gorge reservoir basin university of utah ant- ralist 46 1 21 hropologicalthrop ological papers 45 salt lake city utah 927 YOUNG J A R A EVANS P T TUELLER 1976 great 121 appp basin plant communities pristine and grazed 912 WOODBURY W V 1966 the history and present pages 186 215 in R elston and P headrick status of the biota ofanahoofanabo island pyramid lake eds holocene environmental change in the nevada unpublished thesis university of ne- great basin nevada archeologicalArchearchaeologicalological survey re- vada reno search paper 6 reno nevada 913 WORKMAN G W N E WEST B HAWS AND 928 YOUNG S 1958 Exclosures in big game manage- C HURST 1983 aquatic and riparian ecological ment in utah journal of range management 11 relationships oftheodtheofthe north fork oftheodtheofthe virgin river 186 190 and its deep creek crystal creek tributary 929 youngblood A P AND R L MAUK 1985 conifer- zion national park utah unpublished report ous forest habitat types of central and southern prepared for department of fisheries and utah USDA forest service general technical wildlife utah state university logan 272 appp report INT 187 intermountain research station 914 WRIGHTWRIGHTHH A 1964 an evaluation ofseveralofseveral factors ogden utah 89 appp to determine why sitanionSitanion hystixhystrix is more resistant 930 ZAMORAZAMOBA B 1968 artemisia arbuscula artemisia to than burning stipa comatacamata unpublished dis- longilongilobaloba andarteimwand atzeartemisiaarzeATtemisiamisla nova plant associations min sertationsertation utah state university logan central and northern nevada unpublished thesis 915 WRIGHT H A 1980 the role and use of fire in the university of nevada reno desert grass shrub USDA ser- semidesertsemi type forest 931 ZAMORA B AND P T TUELIERTUELLER 1973 general artemisia vice technical report INT 85 inter- arbuscula A longilobalongiloba and A nova habitat types and range mountain forest experiment station in northern nevada great basin naturalist 33 ogden utah 23 appp 225242225 242 916 WRIGHT H A L F NEUEN SCHWANDER AND C M neuenschwander 932 ZAN M 1968 evaluation of the effects of reduced BRITTON 1981 role and use of fire in sage the transpiration upon soil moisture retentions in an brush grass and pinyon juniper plant communi- aspen stand general throughout the growing season in ties USDA forest service technical re- northern utah unpublished thesis utah state 58 and range port INT intermountain forest university logan 49 appp experiment station ogden utah 933 ZARN M 1977 ecological characteristics ofpinyon 917 WYCKOFF J W 1973 effects of soil texture on the woodlands on the colorado plateau species diversity in an arielarlei grassland of the east- juniper and A literature survey USDI bureau of man- great basin great basin naturalist 33 land ern agement technical note 310 colorado 163 168 denver 183 appp 918 YAKE S AND J D brotherson 1979 differentia- 934 ZOHNER K D 1967 A guide to the biogeographic tion ofserviceberry habitats in the wasatch moun- in literature ofutah unpublished thesis university tains of utah journal of range management 32 of utah salt 83 appp 379 386 lake city 919 YOUNG J A 1981 demography and fire history ofaof a western juniper stand journal of range manage- KEYWORD CITATION INDEX ment 3450134.50134 501 505 920 YOUNG J A AND J D BUDY 1979 historical use of age size structure 1 125 126 134 561 nevada s pinyon juniper woodlands journal of alpine 3 62 63 74 114 115 164 376 390 480 495 forestry history 23 112 121 503503559571698699700559 571 698 699 700 711 742 827 921 YOUNG J A AND R A EVANS 1970 invasion of autecology 21 22 41 46 48 61 63 71 100 104 121 medusmedusaheadahead into the great basin weed science 122 123 133 135 137 169 176 219 231 250 18188989 97 279 333 351 353 355 366 379 400 402 405 922 YOUNG J A AND R A EVANS 1973 downy brome 438 455 494 507 543 580 616 620 649 656 intruder in the plant succession ofbigosbigofbig sagebrush 687 688 717 734 766 797 800 818 830 833 communities in the great basin journal of range 844 859 882 891 903 904 918 921 management 26 410 415 baseline study 7 8 10 11 16 18 19 20 30 31 32 33 923 YOUNG J A AND R A EVANS 1974 population 36 46 53 54 55 56 57 65 75 76 89 98 112 dynamics of green rabbitbrushrabbitbrush inm disturbed big 117 118 119 143 151 154 156 160 163 173 sagebrush communities journal of range man- 174 175 185 186 193 195 196 197 198 200 agement 272712727.127127 132 204 205 206 208 210211210 211 221222221 222 227 229 924 YOUNG J A AND R A EVANS 1978 population 236 242243248243249243242 243 244 246 247 248 249 251 252 dynamics after wildfireswildfires in sagebrush grasslands 257 258 262 266 267 277 281302281 302 303 305 journal of range management 31 283 289 320 323 326 327 328329328 329 330 331 332 335 925 YOUNG J A R A EVANS AND R E ECKERT 1985 337 357 362 363 364 365 371 372 373 374 successional patterns and productivity potentials 375 380 381382381383381 382 384 397 398 399 404 408 of the sagebrush and salt desert ecosystems in 417 421 422 423 424 425 433 448 453 454 1990 NEVADA UTAH vegetation bibliography 241

461 462 475 486 490 504 525 526 527 529 fire 35 93 116 142 156 159 181 248 292 296 344 533 538 562 572 575 595 602 603 604 605 385 386 420 464 473 476 492 515 517 518 606 611 623 628 636 648 651 685 711 719 520 540 566 633 723 750 783 809 834 880 731 732 733 745 758 769 772 809 811 813 881914915916919924881914915916 919gig 924 814 819 820 823 850 851 852 853 856 873 forest 1 3 4 13 27 28 37 39 41 93 125 145 159 160igo gog 1341327283739 4193125145159160 877885894906912932877 885 894 906 912 932 168 177 178 179 203 219 228 230 240 260 bibliography 183 184 304 600 791 812 855 875 276 280 286 304 359 367 383 387 394 402 biogeography 9 49 64 66 67 68 72 74 76 104 105 418 419 428 446 459 460 475 481 485 488 106 107 109 111 123 127 129 131 144 145 496 507 515 516 517 518 520 522 537 561 148 150 155 164 167 168 169 170 180 186 567 568 569 570 584 608 630 655 668 702 189 190 192 202 213 215 225 228 232 238 707 712 714 715 746 754 756 763 835 839 242 244 245 246 249 263 264 268 271 280 902929932902 929 932 284 285 297 306 307 308 309 310 315 317 gradient analysis 120 265 278 286 510 530 545 551 321 334 338 340 341 348 351 352 361 365 563564637660 736 737 738 740 741 839 371 372 377 403 406 409 411 426 427 430 563564637660736737738740741839 427430 grassland 29 34 91 110 113 176 180 214 224 262 431 434 437 444 451 456 458 478 479 482 274 278 282 283 319 340 345 347 354 355 485 487 491 495 496 497 498 499 500 501 356 357 368 438 466 467 468 469 470 510 502 503 505 512 514 518 523 527 542 547 511 514 574 598 611 612 549 551 556 558 559 565 577 579 583 584 616 624 637 682 701 596 598 608 615 620 627 631 635 647 650 708 713 728 734 753 764 817 818 833 651 658 661 666 667 670 672 685 689 690 874916917924 695 698 699 700 701 702 703 704 705 706 grazing 24 34 40 90 91 102 124 136 159 166 194 707 708716708 716 718 721 727 729 739 742 744 203 206 207 208 222 223 224 240 253 269 749 752 755 760 771 775 789 790 796 798 270 273 275 282 283 300 312 313 318 319 802 805 806 807 808 810 816 825 840 843 335 337 343 355 356 360 416 418 428 447 845 846 864 866 867 872 878 887 891 899 450 463 489 490 520 548 569 591 594 597 907908910925934907got 908 910gioglo 925 934 623 633 652 657 663 668 669 681 682 683 classification 13 51 6969138212138 212 280 293393293 393 396396444444 684 728 743 750 759 761 765 767 776 792 457 459 460 522 546 560 613 614 730 747 883904927883 904 927 754786788754 786 788 habitat type 341378921221371552123943953 4 13 78 92 122 137 155 212 394 395 community analysis 3 4 7 8 15 18 20 23 25 27 29 488537684754788799902929931488 537 684 754 788 799 902 929 931 30 35 40 42 48 49 52 60 65 71 95 96 99 inventory 5 9 10 11 12 16 17 19 31 32 33 36 40 103 105 107 108 113 114 115 118 119 127 5053545556576264667579818283505354 5556576264 667579818283 129 130 131 135 139 140 141 144 150 153 84 85 86 87 88 89 97 98 102 105 106 107 155 157 160 171 174 175 177 181 182 188 108 112 117 127 131 139 148 149 154 163 189 199 208 217 233 235 241 253 258 261 164 166 167 168 176 177 180 185 186 187 276 283 288 289 290 292 294 298 299315299 315 188 189 192 193 195 196 197 198 200201200 201 321 324 325 342 344 347 349 350 354365354 365 205 210 211 213 215 216 226 232 236 244 370 375 378 395 407 415 439 442 449 452 245 251 252 256 257 268 271 277 280 291 456 463 465 466 469 477 481 488 493 498 306 307 309 310 316 323 326 327 328 329 499 504 511 523 524 541 544 555 557 558 330 331 332 338 340 341 352 361 369 371 563 567 570 573 574 590 607 609 617 641 372 378 380 381 384 385 387 388 389 390 642 649 653 655 673 674 675 676 696 713 391 394 395 398 399 403 404 407 408 409 748 756 761 764 773 777 779 780 782 785 410 411 413 417 421 422 423 424 425 426 792 793 801 815 828 835 838 839 849 860 430 446 448 452 453 454 461 462 465 478 862 863 865 868 870 876 884 887 896 897 479 481 482 485 487 496 497 498 499 500 900920923924926927928930933900goo 920 923 924 926996 927 928 930 933 503 504 512 516 525 526 527 529 534 535 desert 30 46 47 48 49 51 58 69 75 90 99 102104102 104 537 539 541 542 547 558 559 565 568 570 109 140 149 153 162 167 199 202 204 206 575 577 583 587 595 601 602 603 604 605 227 263 264 265 266 299 300 302 305 311 606 615 616 622 626 627 628 631 635 636 321 322 349 479 502 512 531 548 549 597 647 650 654 661 665 677 678 689 699 701 622 653 675 676 690 691 695 724 725 726 702 705 707 708 712 716 718 719 721 727 727 743 761 762 789 790 825 831 832 842 731 732 742 744 745 752 755 766 772 773 sti871899907871 899 907 775 784 789 790 796 797 802 804 807 808 disturbance 26 35 37 45 47 50 51 58 93 124 142 810 811 813 814 815 818 820 821 822 823 156 181 221 224 248 287 288 292 294 296 825 826 829 833 840 845 846 847 848 850 344 346 385 386 416 420 464 473 475 476 851 852 853 854 858 866 871 872 874 905 492 515 517 520 528 529 531 540 550 554 909910911926929931909gog 910gio gil911 926 929 931 566 569 582 600 609 633 668 674 676 691 map 2513015020145925 130 150 201 459 460 539 677 895 723 724 725 726 739 750 763 776 778783778 783 model 3838141142147161311141 142 147 161 311 367367428513428 513 693 762 809 813 824 826 834 870 880 881 914 915 relict vegetation 250 483 519 521 534 536 619 804 916919923924916gig 919gig 923 924 843 diversity pattern 291 293 324 360 369 468 610 662 shrubland 6 232324455224 45 52 61 70 77 809080 90 91919410094 loo100 749917 101 103 109 111 116 123 124 126 132 133 early exploration 34 59 152 165 239 254 259 502 135 141 155 156 166 169 170 171 172 173 549 553 589 618 644 645 646 664 697 712 174 175 182 225 243 258 264 279 294 298 735751774805836892735 751774805836 892 302 303 305 312 333 340 353 358 360 379 242 P S bourgeronetalBOURGERON ET AL volume 50

385 386 389 415 416 429 430 431 443 444 550 566 569 571 573 591 594 597 633 637 449 472 477 486 530 540 555 556 557 565 638 639 640 668 669 676 680 682 683 690 579 582 583 588 591 592 594 597 598 599 693 714 715 720 723 724 725 726 739 750 619 637 657 660 669 673 674 679 680 682 753 770 774 776 778 781 783 794 795 799 723 736 737 738 743 757 770 771 791 793 803 809 817 824 831 832 834 842 843 861 794 795 799 801 816 817 838 844 855 856 869 880 881 883 885 889 890 906 915 916 858 860 862 864 865 866 867 868 871 873 919921922923924925927919gig 921 922923 923 924 925923 927 874 876 878 879 882 883 889 916 918 922 wetlandtlandaland 95 96 115 119 120 130 301 311340311 340 351 923925930931923 925 930 931 399 440 442 528 585 586 629 641 659 729 soil vegetation relationship 14 26 42 43 44 70 71 77 769901769 901golgoi 798182838485868788941201291327981 82 83 8485 8687 8894120129132 140 143 157 158 162 204 229 230 234 237 woodlandodland 2 7 8 14 15 23 35 39 52 60 68 73 137 249 254 255 257 260 263 278 286 295 297 142 146 147 175 212 228 235 288 289 290 313 317 322 336 339 345 346 350 370 376 291292291 292 293 294 295 296 318 344 400 401 382 392 429 442 443 445 470 471 472 479 435 464 473 474 482 483 544 545 546 566 491 508 529 534 535 546 552 554 555 556 573 578 583 592 593 599 617 632 638 648 557 563 564 576 588 621 626 630 642 643 665 739 750 776 777 778 779 780 781 782 679 686 687 691 709 710 722 746 749 771 783 787 798 800 822 824 834 857 858 861 827853877879886893894898917827 853 877 879 886 893 894 898 git917 869 871 874 875 884 885 886 887 888 890 898 908 916gig 919gig 920 933 succession 23 35 37 38 47 51 65 78 80 94 110 128 898908916919920933 151 161 172 173 178 179 181 182 188 191 zonationlation 67 72 120 435 551 581 625 653 671 704 209 214 224 233 248 255 258 261 262 269 768788837841888768 788 837 841 888 270 272 273 275 282 287 288 296 300 301 358 359 360 366 367 368 385 401 434 439 440 441 447 457 467 472 473 474 476 480 received I1 july 1990 484 489 490 492 515 517 520 531 532 540 accepted 15 september 1990 great basin naturalist 503 1990 appp 243 248 conservation STATUS OF threatened FISHES IN WARNER BASIN OREGON

jack E williams mark A stern2sterna alan V Munhall3 and gary A anderson 4

abstractabsrract two federally listed fishes the foskett speckled dace and warner sucker are endemic to warner basin in south central oregon the foskett speckled dace is native only to a single spring in coleman valley A nearby spring was stocked with dace in 1979 and 1980 and now provides a second population the present numbers of dace probably are at their highest levels since settlement ofthe region the warner sucker historically occurred throughout much of the warner valley but its distribution and abundance have been reduced by construction of reservoirs and irrigation dams and the introduction of predatory game fishes lentic habitats have become dominated by introduced fishes particularly white crappie black crappie and brown bullheadbullbulibuilheadbead the largest remaining population ofwarner suckers occurs in hart lake where successful reproduction was documented but there is no evidence of recruitment to the adult population two threatened fishes inhabit separate val- deep warner creek near adel the leys in warner basin oregon in coleman warner sucker was listed as threatened pri- valley the only native fish is the foskett marily because of fragmentation of stream speckled dace rhinichthys esculusosculus sspasp habitats by irrigation diversion dams and the which occurs in foskett spring along the west establishment of large populations of intro- margin of the coleman lake bed the lake is duced piscivorous fishes in lentic habitats dry except during years of exceptional rainfall USU S fish and wildlife service 1985b the dace was listed as threatened because of longtimelong time residents recalled that during small population size trampling of its re- the 1930s large numbers of spawning warner strictedstricted habitat by cattle and subsequent suckers referred to as redhorse ascended degradation of the springspringpoolpool area UUSS fish honey creek far into upstream canyon areas and wildlife service 1985a andreasen 1975 by the 1970s the species was fragmented by provide a refuge population free of range numerous irriga- to diversion dams on the lower of the effects of intense livestock grazing 50 tion reaches streams tributary pelican crump and dace from foskett spring were transplanted to lakes andreasen 1975 kobetich 1977 on 14 november 1979 into an unnamed spring hart swensen 1978 coombs et al 1979 hayes now known as spring on bureau of dace 1980 which block spawning runs from the management BLM land land approxi- lakes into streams mately 151.5 kinkm south of foskett spring an- i coombs et al 1979 found that although other 50 dace were transferred into the springsprcprmg habitats had been fragmented resident stream on 26 august 1980 A reproducing population populations still persisted nearly two thirds subsequently established in dace spring and ofall adult suckers 198 of 300 were captured more than 300 dace of three size classes were by coombs et al 1979 in the canal between observed there in 1986 BLM lakeviewLakeview dis- anderson and hart lakes immediately north trict unpublished data of the hart lake spillway adult and larval the presumed historical range of the war- suckers also were captured in snyder creek ner sucker catostomus warnerensiswarnerensis con- in honey creek above the dam at plush at sisted of the main warner lakes pelican the mouth of honey creek in hart lake at crump and hart and other accessible lakes the south end ofwarner valley in twentymileTwentymile and sloughssloughy in warner valley and low to creek between the south end of the valley moderate gradient reaches of tributary floor and the confluence with twelvemileTwelvemile streams the species description by snyder creek and in twelvemileTwelvemile creek immediately 1908 was based on specimens collected from above and below the okeefe diversion dam

division of wildlifeofwildlife and fisheries bureau eflandoflandof land management 18th & C sheetsstreets NWN W washington D C 20240 2theathehe nature conservancy 1205 northwest 25th avenue portland oregon 97210 3bureaubuieaubudeau of land management box 151 lakeviewLakeview oregon 97630 40regonoregon department of fish and wildlife box 1214 lakeviewLakeview oregon 97630

243 244 J E WILLIAMS ETALET AL volume 50

in 1980 coombs and bond 1980 sampled lakes by use of traps gill nets and seines and 22 sites throughout the basin capturing 46 from streams with dip nets trap nets elec warner suckers at 4 localities honey creek shockertroshockertro kick nets and seines most fishes between hart lake and the dam at plush were identified measured and returned to canals ofdeep creek at the east end ofpelican their habitat voucher specimens or those ac- lake the spillway immediately north of hart cicidentally killed during collecting are housed lake and swamp lake in 1983 smith et al at the wildlife and fisheries museum uni- 1984 captured 1 adult warner sucker in versity of california davis Opoperclesopercledercles from crump lake and 2 juveniles approximately five suckers were aged according to the meth- 130 mm total length TL in deep creek ods described by scoppettone 1988 visual between adel and the falls in 1987 an adult observations were made of spawning warner warner sucker was caught by an angler along suckers in honey creek the slough just south of flagstaff lake J E williams personal observation FOSKETT SPECKLED DACE this paper summarizes the current status of these two threatened fishes as determined in 1987 the BLM acquired foskett spring by surveys conducted from 1987 to 1989 and the surrounding 65 ha of which approxi- other native fishes ofwarner valley include a mately 28 ha were fenced to exclude cattle local form of redband trout oncorhynchus the dace population at foskett spring has mykiss sspasp tui chub gila bicolorbwolor and the since expanded to the spring pool its outflow common form of speckled dace R osculus and downstream marsh baseline water qual- the warner valley redband trout largely has ity and vegetation monitoring at foskett and been displaced by introduced trout and is dace springs were initiated by BLM in 1987 listed as of special concern by the american the following data collected on 28 september fisheries society williams et al 1989 1988 from foskett spring and dace spring respectively exemplify the two habitat simi- 19 HABITAT description AND SURVEY METHODS laritieslarities air temperature and 17 C water temperature 17 and 16 C dissolved oxygen the warner basin comprises 6858 sqaq km in 535.3 and 595.9sg mglmgtmg1 conductivity 350 and 250 south central oregon and small portions of mohscmmohscm ph 81si8.1 and 828.2 alkalinity 114 and northeastern california and northwestern ne- 99 mglmgtmg1 caco3cac03cacos hardness 40040.0 and 24724.7 mglmgtmg1 vada fig 1 drainage is internal and is di- and turbidity 141.41 4 and 1181.8isls 8 NTU vided between coleman valley and the much the dace population maintains itself at larger warner valley coleman valley is a dace spring despite a tendency for vegetation separate drainage in the southeastern part of to choke out most open water the introduced the basin and receives sparse runoff observa- population has expanded by movement offishof fishhishbish tions ofthe foskett speckled dace and its habi- through a connecting pipe into a livestock tat in coleman valley were made from 1987 to watering trough just east of the spring no 1989 standardized transects were estab- other fish occur in coleman valley lished along foskett spring and its outflow to monitor vegetation recovery following cessa- WARNER SUCKER tion of grazing and to quantify amounts of open water habitat surveys on twentymileTwentymile creek above and in warner valley all water flows into a se- below the dyke diversion dam located I1 ries of north south oriented shallow lakes adult and 2 larval warner suckers in 1988 sloughssloughy and potpotholespotholedholes during periods with additional 1987 and 1988 surveys failed to above average precipitation as occurred dur- locate warner suckers elsewhere in twelve ing the early 1980s and again in 1989 these mile creek including sections in nevada and lakes fill from the south and eventually over- oregon upstream of the nevada border the flow into the northern part of the valley only canal north of hart lake the slough between the three most southerly lakes pelican flagstaff lake and mugwump lake the crump and hart are permanent fish col- slough between lower campbell and camp- lections in warner valley were made from bell lakes or stone corral lake in april 1987 to 1989 samples were collected from 1989 28 adult suckers were captured at the 1990 threatened WARNERWABNER BASIN FISHES 245

bluejoint lake OREGON sloncorralSlonstonsron corraiCorral warner L basin

bellbeilbeli lake

flagstaff lowercamphellLowerCamphell L lake mugwump L swamp lake anderso t cr cr spillway 1vav af0f

plush hartlakeHarthartlageLake

sixteen kilorneterKilorneter ten miles

crump lallollau

pelican L peep cr

adelladeil

Twentytwentyilelleile

clemaciema oregon lake twjvemjle fattfaitspring california nevada

fig 1 the warner basin of south central oregon 246 J E WILLIAMS ETETALAL volume 50

TABLE 1 frequency of fishes collected in warner basin during 1987891987 89 all collection sites are inm lake county oregon unless otherwise noted collections made at the same habitat are combined large warner tui speckled white black mouth spotted brown location sucker chub dace trout crappie crappie bass bass bullhead twelvemileTwelvemile cr 404 washoe co NV twelvemileTwelve mile cr 591 51 dyke diversion canal I1 irrigation canal along I1 1 1 twentymileTwentymile cr Twentytwentymiletwentymilecrmilemlle cr 6 25 854 5 2 4 greaser reservoir 476 1 2 deep creek 400 40 hart lake 70 12 1620 14 1 449 lower honey cicreekeek 69 upper honey creek 19 canal north of hart lake 7 2 31 30 1 anderson lake 7 10 1 flagstaff lake slough 27 107 59 40 slough between lower 82 39 campbell and campbell campbell lake 371 95 stone corral lake 17 5 total caught 152 649 2269 96 2183 106 2 1 607 relative catch 25 107 374 16 260 17 0 I1 0 1 100 may include native redband trout andor introduced rainbow trout

mouth of honey creek in hart lake and 42 hart lake or the canal immediately north of were captured along the east side of hart the lake introduced fishes dominated the lake fish ranged from 311 to 440 mm TL fauna of hart lake and other lakes and sloughssloughy aagavg 3852385.2 n 70 with most 350 to 410 mm in the valley white crappie pomonispomoxis annu approximately 80 100 other adult warner laris and brown bullhead ictalurus nebulonebule suckers were observed in honey creek be- sus outnumbered native fishes in our collec- tween the most downstream diversion dam tions from hart lake by slightly more than and hart lake these fish were in breeding 251 tui chub which historically was the condition and migrating upstream where most abundant fish in lentic habitats largely they were visible because flow in the creek has been replaced by white crappie was reduced by upstream diversions in mid the warner sucker population appears to may 1989 water began spilling from hart be largest in hart lake but no recent recruit- lake into the canal toward anderson lake ment could be documented except for a suckers dispersed into the canal and 7 small number oflarvae in lower honey creek no suckers smaller than 310 mm TL spannersspawnersspawners were collected there in june stan- were found dard length TL and age of 5 of these were white crappie were abundant at the mouth of honey during june and may 331 357 7 307 361 7333 387 7335 390 creek 331357730736173333877335390 have preyed on sucker larvae as they drifted 9 and 340 397 8 larval suckers also were collected from honey creek just above the into hart lake A single trap net set there in downstream most diversion dam indicating june collected 1530 white crappie and 20 brown bullhead at least limited spawning upstream overall warner suckers constituted only discussion 252.5 of all fishes collected during 1987 89 table 1 nearly all suckers were found in the foskett speckled dace appears to be hart lake honey creek just upstream of near recovery no exotic species are present 1990 threatened WARNER BASIN FISHES 247 in either spring and the primary threats have are needed to determine the extent of any been eliminated some vegetation needs to remaining sucker populations if present be cleared from the pool at dace spring in however recruitment may be prevented by order to provide sufficient open water also populations ofcrappieof crappiecrapple fencing along the boundary of dace spring in conclusion warner suckers once were should be extended to the east to include common throughout the basin but gradually additional habitat continued habitat and declined from about 1900 until the early 1970s population monitoring are necessary at both as a result of agricultural development and springs because the small habitats are vulner- placement ofirrigation structures in spawning able to slight disturbances streams despite habitat fragmentation and the largest remaining population of war- lack ofoffishfishhishbish passage recruitment to lake popu- ner suckers appears to be in hart lake where lations continued until the late 1970s when spawning fish ascend lower honey creek and large populations of piscivorous fishes became the canal north of the hart lake spillway established recruitment of warner suckers populations also may exist in crump and peli- continues in stream habitats but appears from can lakes our observations to be greatly curtailed since successful recruitment of young into the 1979 hart lake population is limited by reduced control of introduced fishes in hart and spawning habitat in honey creek and large crump lakes may be impractical because of populations of crappie white crappie were habitat size 2928 and 3108 ha area respec- introduced into hart lake in 1971 and white tively and large populations recovery of the plus black P nigromaculatus crappie were warner sucker in hart lake therefore at least introduced into crump lake during 1972 and requires increased spawning sites and rearing 1973 oregon department of fish and wild- habitat life unpublished data subsequent collections of the oregon department of fish and acknowledgments wildlife indicated that white crappie black crappie and brown bullhead were common in our fieldwork was aided by W J berg crump lake by 1978 K daily unpublished R G bolton C A macdonald J F data and presumably in hart lake as well morawski G A rosenberg L M swinney adult white crappie commonly feed on small andrand R K white mark warner of the nevada fishes pflieger 1975 thus their abundance at department of wildlife facilitated our collec- the mouth of honey creek during the same tions in nevada earlier drafts of this time that larval suckers were collected from manuscript benefited from reviews by J K the creek increases the likelihood predationofpredationof andreasen W J berg and C D williams on young of year suckers age analysis of the warner suckers was kindly seven irrigation dams on honey creek be- confirmed by G G scoppettone this work tween the lake and plush result in limited was completed while the senior author was on access by adults to upstream spawning areas an intergovernmental personnel act appoint- during 1989 only two fifflesriffles between hart ment with the UUSS fish and wildlife service lake and the first diversion dam contained at the university of california davis our suitable gravel for spawning depending on sincere appreciation to mr joe flynn for ac- stream flows water diversion boards may cess to his property along honey creek and be placed in the irrigation structures before for his interest in the fishes therein during or after the spawning run swenson 1978 reported that during 1978 adult suckers CITED migrated as far as the seventh irrigation dam literature at plush before boards were installed and wa- ANDREASEN J K 1975 systematics and status of the ter diverted for irrigation family catostomidae in southern oregon unpub- A remnant population of warner suckers lished dissertation oregon state university cor- may persist in crump lake as indicated vallis 76 appp COOMBS 1 ANDCAND C BOND by collection in 1989 of young of year in C I E 1980 report ofinvestigaofinvestiga- greaser tions on catostomus warnerensiswarnerensis fall 1979 and twenty mile slough below dam ad- spring 1980 report to U S fish and wildlife ditditionalional surveys of crump and pelican lakes service sacramento california 248 J E WILLIAMS ETALET AL volume 50

COOMBS C I1 C E BOND AND S F DROHAN 1979 SNYDERSNYDEB J 0 1908 relationships of the fish fauna of the spawning and early life history of the warner lakes of southeastern oregon bulletin of the bu- sucker catostomus warnerensiswarnerensis report to USU S reau of fisheries 271907 69 102 fish and wildlife service sacramento califor- SWENSON S C 1978 report of investigations on cato- nia stomus warnerensiswarnerenszsmarnewarnerensis during spring 1978 report to HAYES J P 1980 fish of warner valley pages 131 137 U S fish and wildlife service sacramento cali- in C gilman and J W feminella eds plants fornia 27 appp and animals associated with aquatic habitats of USU S FISH AND WILDLIFE SERVICE 1985a determination warner valley oregon state university corvallis of threatened status for hutton tui chub and fos- KOBETICH G C 1977 report on survey ofwarner valley kett speckled dace federal register 50 12302 lakes for warner suckers catostomus warnerenwarngren 12306 sis report to U S fish and wildlife service 1985b determination that the warner sucker is a Saciasacramentosacramento california 66ppappappp threatened species and designation of its critical PFLIEGER W L 1975 the fishes of missouri missouri habitat federal register 50 39117 39123 department ofconservation 343 appp WILLIAMS J E J E JOHNSON D A hendrickson scoppettone G G 1988 growth and longevity of the S CONTRERASCONTREBAS BALDERAS J D WILLIAMS cui ui and longevity of other catostomids and M NAVARRO MENDOZA D E mcallister AND cyprincyprinidscypnnidsids in western north america transactions J E DEACON 1989 fishes of north america en- of the american fisheries society 117 301 307 dangereddangered threatened or of special concern 1989 SMITH M T STEINBACK ANDANDGG PAMPUSH 1984 distri- fisheries bethesda 146 2 20 bution foraging relationships and colony dynam- icsacs of the american white pelican pelecanus received 30 november 1989 erythrorhynchos in southern oregon and northeastern california oregon department of fish revised 11 august 1990 and wildlife nongame technical report 8300483 0 04 accepted 6 september 1990 great basin naturalist 503 1990 appp 249 256

HOME RANGE AND ACTIVITY PATTERNS OF BLACK TAILED jackrabbitsJACKRABBITS

graham W smithsmithl

ABSTRACT home range use and activity patterns ofblack tailed jackrabbitsjackrabbits lepaslepus californicuscaliforcalifoalifornicus in northern utah were studied using telemetry home range sizes ranged from 1 I1 kmkm2 to 3 kmkm2 and did not differ between sexes or among seasons Jackjackrabbitsrabbits were inactive during daylight became active at dusk and remained active throughout the night animals often traversed their home ranges in a few hours during the breeding season males were more active than females Jackjackrabbitsrabbits were most active during well lit nights and high winds decreased jackrabbitjackrabbit activity the black tailed jackrabbitjackrabbit occupies a wide spring accumulated snowfall in 1983 1984 geographic area and is an important compo- was 69 cm at snowvilleSnowville utah 15 km east of nent of the biota throughout its range in the the study area with snowcoversnowcover persisting great basin the jackjackrabbitrabbit is the most abun- from mid november through mid march dant large herbivore wagner 1981 and national oceanic and atmospheric adminis- serves as an important prey item for many tration 1984 predators considering the central role of the jackrabbitjackrabbit in many ecosystems little research METHODS on the species has been reported detailed black tailed jackrabbitjack quantitative information regarding activity rabbit activity and home range use were monitored via telemetry patterns and home range use is lacking home incidental direct observations ofofjackrabbitsjaekjack range use varies with the patterns of food jackrabbitsrabbits also describing cover and water distribution dunn et al aided in activity patterns black tailed jackrabbitsjackrabbits were captured 1982 I1 examined patterns ofjackrabbitjaekjackofjackrabbitrabbit activ- by night lighting griffiths ity and home range use throughout a calendar and netting and evans 1970 they then year in shrub steppe vegetation in northern were equipped with utah radio transmitters and released periodically during each season additional animals were STUDY AREA caught and instrumented to replace those that had died during winter some jackrabbitsjackrabbits the study was conducted in northern were captured in live traps and handled simi- utah near the wildcat hills in curlew valley larly to those captured by netting sex was about 10 35 km north of the great salt lake determined from external examination ofofgenhofgengen topography and vegetation of the area are italia and age class was estimated from body described in detail by gross et al 1974 size color and relative eye size L C stod- four major vegetation types occur in curlew dart unpublished data valley 1 open stands of juniper juniperus transmitter collars were designed to mini- osteosperma at higher elevations 2 big mize chafing of the jackrabbitjackrabbit s neck wywi- sagebrush artemisia tridentata in the north- alowski and knowlton 1983 I1 assume the ern portions of the study area 3 greasewood transmitters had little discernible effect on sarcobatus wrmiculatusvermiculatus in more saline soils jackrabbitjackrabbit behavior stoddart 1970 donoho closer to the great salt lake and 4 expanses 1972 brand et al 1975 keith et al 1984 of salt desert vegetation primarily shadscaleshadscale telemetry stations on the wildcat hills atriplex confertifoliaconfertifolia and saltbush atriplex overlooked areas with instrumented jack falcata scattered throughout the study area rabbits each station was equipped with two southern portions ofthe valley are typically horizontally stacked 5 element yagi anten- more xeric than northern portions and pre- nas coupled out of phase with a sum and cipitation is most abundant during winter and difference hybrid junction A compass rose

department of fisheries and wildlife utah state university logan utah 84322 present address office of migratory bird management USU S fish and wildlife service laurel maryland 20708

249 250 G W SMITH volume 50

mounted on the antenna mast indicated the comparisons with other published accounts I1 directional orientation of the antennas A also report the 95 contour interval although transmitter placed at a known azimuth from it probably overestimates home range size each station was used as a beacon to orient no statistical rationale exists for the choice of the compass rose to true north the 95 level and its use may result from home range use was assessed by repeat- biologists confusing utilization distributions edly recording azimuths of animals simulta- with alpha levels in statistical tests white and neously from two tracking stations four hour garrott 1990 tracking sessions were distributed throughout the relationships of time of day season the 24 hour day with most occurring between sex amount of moonlight and wind intensity dusk and dawn locations were recorded to jackrabbitjackrabbit activity were assessed using log every 20 minutes for a 4 hourbour period on all linear analyses sokal and rohlf 1981 terms animals whose transmitter signals could be included in the resulting models reflect signif- detected for each reading it was noted icant relationships within the data seasons whether the signal varied in amplitude sug- were defined by curlew valley weather pat- gesting movement of the transmitter antenna terns winter ended with the melting of snow which was interpreted as movement by the in march summer began in late june 1984 animal the amount of night light was classi- with the onset of hot temperatures and ended fied into one of three categories low me- in early september with the arrival offall rains dium or high depending upon the phase of the moon and cloud cover wind intensity RESULTS during the 4 hour period was classified as low or high by noting the wind conditions at the the daily movements of 16 jackrabbitsjackrabbits tracking shelters periods of no wind were were monitored from february through april included in the low category 1984 and those of an additional 44 jackjackrabbitsrabbits home range use was assessed using pro- were monitored from june through novem- gram HOME RANGE samuel et al 1985b ber 1984 I1 determined the sizes of areas used this program offers a series of statistical tests by 30 jackrabbitsjackrabbits with 5 animals having 2 to derive the appropriate home range estima- areas each for a total of 35 areas of use table tor samuel and garton 1985 within each 1 the time periods for which areas were data set locations outside the home range measured ranged from eight days to five were identified statistically samuel and gar- months home range has been defined as the ton 1985 and discarded given a weight ofofo0 if area used by an animal on a day to day basis they appeared to be errors or excursions out- burt 1943 how an animal uses its home side the normal home range area burt 1943 range affects how long the animal must be core areas were identified samuel et al monitored before it traverses its entire home 1985a As recommended by samuel and gar- range typical home range use by black tailed ton 1985 only data sets with 550750i 50 locations jackjackrabbitsrabbits involved extended use of an area were analyzed comparisons of the sizes of measuring 1 kmkm2 animals often traversed areas used by subsets ofthejackrabbitof the jackrabbitjackjaekrabbit popula- the whole area of activity in less than four tion were made using a two way analysis of hours by dawn a jackjackrabbitrabbit was usually back variance SAS institute inc 1985 near the previous day s resting location some because of the model selection criteria of animals maintained this pattern of space use the program I1 used the harmonic mean esti- for up to three months mator dixon and chapman 1980 for all area periodically jackjackrabbitsrabbits changed their of use analyses dixon et al 1981 recom- areas of use these changes involved exten- mended this technique to analyze lagomorph sions of the areas of use into previously un- spatial use because it eliminates many prob- used areas and abandonment ofportions ofthe lems associated with other analyses previously used areas new areas were then choice of a contour isopleth to represent generally used for extended periods these the home range is somewhat arbitrary ander- shifts in areas of use enlarged overall home 2 son 1982 I1 chose the 80 contour for jack ranges to 151.5lsis 3 kmkm2 no differences in the rabbits because it appears to reflect observed patterns of shifts in areas of use among differ- patterns ofland use by these animals to allow ent sex and age segments of the population 199011990 BLACK TAILED jackrabbitJACK RABBIT HOME RANGE 251

TABLE 1 black tailed jackrabbitjackrabbit home range sizes kmkm2 and standard errors in curlew valley utah 1983 1984

harmonharmonicharmonie ie contour areaare a number season ofanimalsofanimals 95 SE 80 SE core SE adults winter male 2 251 077 1661 66 064 096 033 female 6 125 045 083 028 047 015 spring male 5 108 025 073 0180.180 18 043 0130 13 female 4 085 0160.16olg0 16 055 0130.130 13 032 006 summer male 5 288 072 183 045 107 031 female 4 163 049 105 029 062 0160.160olg16 fall male 1 221 1301 30 072 female 2 087 0140.140 14 052 0100.10olo0 10 026 006 juveniles summer fall male 1 1131.131 13 073 043 female 5 1701.701 70 035 092 021 057 015 two way analysis ofofvariancevariance using the 80 contourcontolur df SS F P season 3 3103 10 235 10 sex 1 2142.142 14 485 04 season x sex 3 056 042 74 error 21 154115 41 were apparent core areas as identified by sizes of areas appeared to change season- program HOME RANGE encompassed ally with areas of use being smaller in spring areas slightly larger than half the size of and fall than in winter and summer patterns home ranges the 80 contour table 1 of home range use also appeared to be similar and tended to be near the center of the home throughout the year males tended to have range areas slightly larger home ranges than females Jackjackrabbitsrabbits changed home range areas sea- table 1 too few juvenile jackrabbitsjackrabbits were sosonally where more than one home range instrumented to adequately determine their was recorded for an animal each was analyzed pattern of home range use however sizes of 1 areas independently table traversed dur- areas used by juveniles and adults did not ing seasonal migrations were not considered differ table 1 only juveniles large enough part of either home range burt 1943 in to wear a radio transmitter 3 months ofage march just after the winter s snowcoversnowcover melted Jackjackrabbitsrabbits left wintering areas and were instrumented moved to new home ranges in the fall many Jackjackrabbitrabbit activity changed daily and sea- animals abandoned their summer home range sonally fig 1 significant differences oc- areas and moved to wintering areas smith curred during the day and among the seasons 1987 aot active P3 time F season in fijkdijkf JLL wintering areas were located in stands of aoti fk ab xfikofik pfjkpajk otpfijk tall vegetation primarily greasewood and atpij0 G 1795179.5 18 df P ooiool001.001 cifikcibik G sagebrush although a fewjuniper stands were o5943594.3 20 df P ooi001001.001 pfjkpajk G 8563856.3 30 df also used wintering areas encompassed only P ooi001001.001 api ijkdijk G 1288128.8 15 df P ooiool001.001 a small portion of the available habitat as Jackjackrabbitsrabbits were least active from 0800 1700 Jackjackrabbitsrabbits concentrated in groups areas of hours during the spring summer and fall low vegetation were not used during winter when they often rested in shallow depressions with spring shifts in home range areas under shrubs big sagebrush and grease- Jackjackrabbitsrabbits reoccupied much of the valley wood being the most common in addition smith 1987 some jackrabbitsjackrabbits used badger taxidea taxus 252 G W SMITH volume 50

080.8 during summer and much more so during fall proportion of observations classified as 07-07 the 0 winterwi nteante active was lower during fall table 2 consis- 06og 0 sp 06- springring results of the analysis A susummerm mer tent with the previous 0.505 05 A I1 1 1 fafallfalifailI fig 0 040.4 Jackjackrabbitrabbit activity was apparently influenced by the of light during night t0 0310.31 amount CL hours 3 proportion of observa- 020.2 table the tions in which jackrabbitsjackrabbits were active at night 01oi0.1 changed seasonally with jackrabbitsjackrabbits most ac- and least fall 1800 06000000oooo 0600 1120000 1800 tive in summer active in the time relationship of activity and night light also changed seasonally table 3 during fall winter and spring jackjackrabbitsrabbits were most ac- fig 1 frequency of black tailed jackrabbitjackrabbit activity and relationships with time of day and season of year in tive when night light was greatest a full moon curlew valley utah 1983 1984 and little cloud cover during summer the amount of night light did not appear to influ- TABLE 2 frequency ofblackosblackof blaekblack tailedtalled jackrabbitjackrabbit activity ence Jackjackrabbitrabbit activity and relationships with sex and season of year in curlew Jackjackrabbitsrabbits were most active when there valley utah 1983 1984 the data are the number of was little wind high winds were associated telemetry readings under the categorical conditions the with decreased Jackjackrabbitrabbit activity table 4 proportion active is in parentheses activity in relation to wind intensity changed seapsexpsex W seasonally table 4 with jackrabbitsjackrabbits less likely during when season 17 active a femalefetfeinaienale male to be active the winter winds were high sampling ofJackjackrabbitjaekofjackrabbitrabbit activ- 129 100 winter yes high winds fall insuffi- 24 24.24 29 29.29 ity during in the was no 409 240 cient for basing conclusions spring yes 108 88 lgig 16.16 20 20.20 discussion no 572 336 home range shape and size summer yes 147 156 24 24.24 22 22.22 the shape of most Jackjackrabbitrabbit home ranges no 456 555 tended to be elliptical the shape is not a fall yes 21 4 result of the locations of the tracking shelters ltit 17.17 04 04.04 in relation to radio collared jackrabbitsjackrabbits as no 101 87 very acute or obtuse telemetry bearings were

dijk from analysis A similar elliptical lnin fijkk itA a pjP firk plipijapapiipii arkofikorik arikprikpr arrupriik excluded the aotaiot0i G 17617.6 4 df P 005005.005 shape was noted by rusch 1965 who ob- ark G 94369436df943 6 df P ooi001001.001 tained many locations from snow tracking pr G 6 Ppooiooi001 PFA 52352365236dfdf sizes of black tailed Jackjackrabbitrabbit home G 17.0170 3 df P 00 1 the arrij 1701703df1703 pooip001 afiaff ranges determined in this study table 1 are larger than those reported for the species in burrows during winter especially when deep other studies rusch 1965 and nelson and snows reduced mobility wagner 1973 also working in curlew val- Jackjackrabbitrabbit activity changed seasonally with ley reported that jackrabbitsjackrabbits used areas 020020.22 jackjackrabbitsrabbits least active during fall and most kinkm22 minimum convex polygon for periods of active in winter and summer fig 1 daily one to two months in the fall and winter it is activity patterns also changed seasonally with not clear why the home ranges I1 report are jackrabbitsjackrabbits less active during morning hours greater in size than those described in these after midnight in the fall earlier studies two factors however may the sexes showed different patterns of have had some influence my relocation effort activity table 2 with males slightly more was more intensive than those conducted pre- active than females during winter and spring viouviouslysly resulting in a greater number of re- females were slightly more active than males locations per animal moreover advances in 1990 BLACK TAILED jackrabbitjackbabbitJACKRABBITBABBIT HOME RANGE 253

TABLE 3 frequency ofblack tailed jackrabbitjackrabbit activity and relationships with amount of light at night and season of year in curlew valley utah 1983 1984 the data are the number of telemetry readings observed under the categorical conditions the proportion active is in parentheses

amount oflight P3 season rF active ax low medium high winter yes 133 37 23 21 21.21 60go 60.60 52 52.52 no 477 25 21 spring yes 33 81 48 ig19 19.19 27 27.27 42 42.42 no 138 223 66 summer yes 234 77 143 34 34.34 31 31.31 34 34.34 no 449 175 273 fall yes 12 21 6 09og 09.09 17 17.17 27 27.27 no 126 104 16 in fkfy fl aoti pjp rkI1 otpijap aiaala H frk arriapri xpij G 76076.0 8 df P ooi001 al ik G 1032103.2 9 df P ooi001 nikvik G 7215721.5 12 df P ooi001 aprijk G 56156.1 6 df P ooi001

tabieTABLE 4 frequency of black tailed jackjaekjackrabbitjadkrabbitrabbit activity kmkm2 areas of use were determined in these and relationships with wind intensity and season ofyearofyear in three studies from visual observations of curlew valley utah 1983 1984 the data are the num- marked animals of ber of telemetry readings under the categorical condi- differences in sizes home tions the proportion active is in parentheses ranges may also reflect differences in habitats among study areas habitat in lechleitner s wind intensity P3 and tiemeier s studies consisted of pastures season rF active a low high and cultivated land plant cover on french winter yes 214 14 et al s 1965 study area was similar to the 32 32.32 09og 09.09 native great basin shrub steppe vegetation no 459 134 of curlew valley spring yes 112 52 two limitations of the home range analysis 18 18.18 28 28.28 may have influenced the results these are no 512 211 the precision of the tracking system and serial summer yes 392 107 correlation of the location data I1 believe that 26 26.26 21 21.21 the precision of the tracking system averaged no 1125 396 200 m smith 1987 mills and knowlton 1989 fall yes 33 6 sequential locations had to be at least 200 m 09og 09.09 27 27.27 apart before I1 could be certain that movement no 318 16 had occurred locations closer to the tracking lnfkin fijkdijk 1aaprkaparkpriaprkoti pj rk alijapij arikurik brik otpriik shelters had greater precision probably to a apjjctpcap G 25231523252.3 4 df P ooi001001.001 minimum of 100 m while locations dis- arfearke G 7041704.1 6 df P oolooi001.001 more pfjkpajk G 7597759.7 6 df P ooi001.001 tant from the shelters had less precision apfgk417ijk G 1988198.819883dfp0013 df P ooi001 probably less than 300 m the most precise locations were located along the arearcarmamm created using the baseline between the two towers as radiotelemetricradiotelemetric and analytical procedures the diameter of a circle the reduced preci- since the early 1970s may have played an im- sion in locating distant jackrabbitsjackrabbits may have portant role produced overestimates of home range sizes lechleitner 1958a in california french home range estimation methods assume that et al 1965 in idaho and tiemeier 1965 in locations are serially independent swihart and kansas also reported seasonal ranges of 02020.2 slade 1984 when serially correlated data are 254 G W SMITH volume 50 analyzed these methods underestimate home 1965 and has also been observed in southern range size all the location data sets showed idaho personal observation some degree of serial correlation I1 used all Jackjackrabbitsrabbits appeared to traverse the entire locations obtained for each animal and did not home range in short periods of time similar subsample to decrease serial correlation be- home range use was reported by lechleitner cause for many of the animals too few loca- 19581958aa who observed both female and male tions were obtained to allow me to discard Jackjackrabbitsrabbits covering their ranges in about an data points and still have a minimum sample hour french et al 1965 also reported that size of 50 locations per analysis table 1 be- Jackjackrabbitsrabbits traversed home ranges in a matter cause home range size is a statistic the great- of hours est value of which lies in comparisons among subsets of a population white and garrott factors governing home range 1990 I1 opted for the procedure that gave me shape and size larger samples SEASONAL VARIATION AND WEATHER EFFECTS home range use the greater activity ofmales compared with resting in forms during daylight hours is a females table 2 during winter and spring is behavior observed in virtually all black tailed probably related to reproductive activity the jackrabbitjackrabbit populations vorhies and taylor reproductive season for Jackjackrabbitsrabbits in curlew 1933 lechleitner 1958b rusch 1965 haug valley usually begins in january and lasts 1969 costa et al 1976 and flinders and el- through may or june gross et al 1974 liot 1979 forms are shallow depressions in black tailed Jackjackrabbitsrabbits have a complex mat- or under bushes vorhies and taylor 1933 ing behavior in which males seek out females form use by curlew valley jackjackrabbitsrabbits dunn et al 1982 males would thus be ex- appears typical for the species the use of pected to be more active than females during burrows during the winter in curlew val- the breeding season lechleitner 1958a and ley however appears unusual lechleitner haug 1969 also report greater activity by 1958b Jackjackrabbitsrabbits use burrows to evade males during the breeding season predators vorhies and taylor 1933 personal other researchers have reported seasonal observation and construct shallow burrows to changes in the daily patterns of jackjackrabbitrabbit escape summer heat in the mohave desert activity donoho 1972 and costa et al 1976 costa et al 1976 but daily use of deep bur- reported that Jackjackrabbitsrabbits were generally less rows has not been reported previously I1 be- active during winter the timing of daily ac- lieve jackrabbitsjackrabbits use burrows during winter in tivity observed in this study changed season- curlew valley to reduce the risk of predation ally and was probably related to changes in smith 1987 day length and times of sunset and sunrise Jackjackrabbitsrabbits have been reported to use sys- similar results were reported by donoho tems oftrails to travel about their home ranges 1972 the evening activity peak I1 observed is vorhies and taylor 1933 french et al 1965 similar to that reported by lechleitner rusch 1965 although trails used by jackrab 1958b and haug 1969 fig 1 however bits in curlew valley were obvious in the haug 1969 also described a period ofheight- snow and vegetation I1 was unable to study ened activity just before sunrise trail use because the tracking system could blackburn 1968 and knowlton et al 1968 not locate jackrabbitsjackrabbits with sufficient accuracy reported that jackjackrabbitrabbit activity was influ- home ranges of individual instrumented enced by ambient air temperature I1 was un- jackrabbitsjackrabbits overlapped extensively I1 have no able to record air temperatures at jackrabbitjackrabbit data that suggest individual jackrabbitsjackrabbits in- locations but my finding that Jackjackrabbitsrabbits fluencedfluenced the home range use by other jack were less active during high winds tends to rabbits from spring through fall although support the idea that temperature influences such intraspecific interactions may have oc- jackrabbitjackrabbit activity lechleitner 1958a and curred my study suggests that during winter tiemeier 1965 also reported decreased ac- jackjackrabbitsrabbits were social and gathered in groups tivity during high winds and inclement smith 1987 similar winter behavior was weather lechleitner 1958a also noted that reported earlier from curlew valley rusch Jackjackrabbitsrabbits were more active during bright 199011990 BLACK TAILED jackrabbitJACKRABBIT HOME RANGE 255 moonlit nights a finding consistent with my and feeding areas were not used this reflects results the availability of resources within home the sizes of home ranges measured by this ranges of curlew valley jackrabbitsjackrabbits where study appeared to change with the seasons feeding and resting resources are available in with spring and fall ranges being slightly the same area jackjackrabbitsrabbits do not need to smaller males used larger areas than fe- travel far from daytime forms to nocturnal males tiemeier 1965 and donoho 1972 feeding sites vorhies and taylor 1933 nel- found no significant differences in home range son and wagner 1973 in areas where feeding size between sexes other researchers how- resources are separated from cover jackrab ever reported that female jackrabbitsjackrabbits used bits have been reported to travel distances 1I1 larger areas than males in summer fall lech- kinkm nightly vorhies and taylor 1933 haug leitner 1958a and winter nelson and wag- 1969 Jackjackrabbitsrabbits have also been reported to ner 1973 differences in procedures espe- shift feeding sites to feed in agricultural fields cially analytical make comparisons among bronson and tiemeier 1959 studies difficult many jackrabbitsjackrabbits changed home range areas on a seasonal basis with acknowledgments animals moving to wintering areas in the fall and early winter and leaving in the spring this study was a part of the predator rusch 1965 working in curlew valley and ecology and behavior project of the den- tiemeier 1965 in kansas also reported sea- ver wildlife research center of the UUSS fish sonal shifts in home range areas with animals and wildlife service the center transferred moving to areas with larger shrubs to the animal and plant health inspection AGE As juvenile jackrabbitsjackrabbits mature one service on 3 march 1986 I1 thank F F would expect their home ranges to increase in knowlton and L C stoddart for their support size it appears that juvenile jackjackrabbitsrabbits in- and guidance K corts L S mills and crease the size of their areas of use to roughly K paulin helped with field research I1 thank that of adults within the first six months post- the many persons especially E hanson and partum young jackrabbitsjackrabbits are precocial are W johnson who helped catch jackjackrabbitsrabbits usually weaned by six weeks of age before the F F knowlton J P gionfriddo and arrival of the next litter and are independent A P wywialowski reviewed the manuscript of their dams at a very young age drake F A johnson prepared the figure 1969 population DENSITY Jackjackrabbitrabbit popula- CITED tions in curlew valley undergo changes in literature density on a 10 year cycle L C stoddart ANDERSON D J 1982 the home range a new nonparamet and F F knowlton mathematical model of ricnc estimation technique ecology 63 103 112 coyote jackrabbit demographic interactions BLACKBURN D F 1968 behavior of white tailed and northern utah poster presented at ath4th in- black tailed jackrabbitsjackrabbits of mideasternmideastern oregon unpublished thesis university of idaho mos- terternationalnational theriologicalTheriothermologicallogical congress edmon- cow 47 appp ton alberta canada 1985 my study was BRAND C J R H VOWLES ANDANDLL B KEITH 1975 snow- conducted during a population low iiee 30 shoe hare mortality monitored by telemetry jour- jackrabbitskmjackrabbitskm2 smith 1987 I1 do not know nal ofwildlife management 3974139 741 747 BRONSON F H AND 0 T TIEMEIER 1959 the relation- whether patterns of Jackjackrabbitrabbit home range ship of precipitation and black tailed jackjackrabbitrabbit use change with population density how- populations in kansas ecology 40 194198194 198 ever I1 noted changes in the pattern of use of BURT W H 1943 territoriality and home range concepts wintering areas with fewer wintering areas as applied to mammals journal of mammalogy 24 used during low densities and habitat types 346 352 COSTA R K A NAGY ANDANDVV H SHOEMAKER 1976 1987 ob- smith suggesting the possibility of servationsservations of behavior on black tailed jackjackrabbitsrabbits other changes in home range use with chang- in the mohave desert journal of mammalogy 57 ing density differences between my findings 399 402 and others reported in the literature may be a DIXON K R AND J A CHAPMAN 1980 harmonic mean measure of animal activity areas ecology 61 function of differing Jackjackrabbitrabbit densities 1040 1044 HABITAT Jackjackrabbitrabbit home ranges in this DIXON K R 0 J RONGSTAD ANDANDKK M ORHELEIN 1981 study were contiguous and separate resting A comparison of home range size in sylvilagusSylvilagus 256 G W SMITH volume 50

floridanusfloridanus and S bachbachmaninachmanimani pages 541 548 in K NELSON L JRJK AND F H WAGNER 1973 effects of sub- myers and C D macinnesmachines eds proceedings of lethal cerebral x irradiation on movement activ- the world lagomorph conference 12 16 august ity and home range patterns of black tailed jack 1979 university of guelph press guelph on- rabbits health physics 2550725 507 514 tario canada RUSCH D 1965 some movements of black tailed DONOHO H S 1972 dispersion and dispersal of white jackrabbitsjackrabbits in northern utah unpublished thesis tailed and black tailed jackrabbitsjackrabbits pawnee na- utah state university logan 43 appp tional grasslands unpublished thesis colorado SAMUEL M D AND E 0 CARTONGARTON 1985 home range a state university fortfortcollinscollins 83pp83 appp weighted normal estimate and tests of underlying DRAKE E 1969 maintenance and social behavior of cap- assumptions journal of wildlife management 49 tured jackrabbitsjackrabbits lepus californicuscalifornicus gray un- 513 519 published thesis utah state university logan SAMUEL M D D J PIERCE AND E 0 GARTONGAR TON 1985a 45pp45 appp identifying areas of concentrated use within the DUNN J PR J A CHAPMAN AND R E MARSH 1982 home range journal of animal ecology 54 Jackjackrabbitsrabbits lepus californicuscalifornicus and allies pages 711 719 124 145 in J A chapman and G A feldham- SAMUEL M D D J PIERCE E 0 GARTON L J NELSON AND K R DIXON 1985b s mer eds I1 wild mammals of north america biol- user manual for pro- ogy management and economics johns hopkins gram HOME RANGE university of idaho university press baltimore maryland forest wildlife and range experiment station FLINDERS J TANDCT AND C L ELLIOTT 1979 abiotic charac- technical report 15 69 appp teristicste of black tailed jackjackrabbitrabbit forms and a hy- SAS INSTITUTE INC 1985 SASSTAT guide for personal popothesisthesis concerning form function encycliaencyclicEncyclia 56 computers ath6th ed carygary north carolina 378 appp 34 38 SMITH G W 1987 mortality and movement within a FRENCH N RRR R mcbbldeandjmcbride AND J DETMER 1965 fertil- black tailed jackrabbitjackrabbit population unpublished ity and population density of the black tailed dissertation utah state university logan 101 appp jackjackrabbitrabbit journal of wildlife management 29 SOKAL R R AND F J ROHLF 1981 biometry ad2d ed 14 26 W H freeman and co san francisco califor- GRIFFITHS R E AND J EVANS 1970 capturing jackrab nia 859 appp bits by night lighting journal ofwildlifeofwildlife manage- STODDARTSTODDABT L C 1970 A teleteietelemetricmetriemetric method for detecting ment 3463734.63734 637 639 jackrabbitjackrabbit mortality journal of wildlife manage- 34 501 GROSS J E L C STODDART AND F W WAGNER 1974 ment 3450134.501soi 507 demographic analysis of a northern utah jack SWIHART R K AND N A SLADE 1985 testing for inde- rabbit population wildlife monographs 40 68 appp pendencependence of observations in animal movements 66 HAUGHAUGJJ C 1969 activity and reproduction ofthe black ecology 6611761176 1184 tailed jackrabbitjackrabbit in the coastal cordcordgrassgrass prairie of TIEMEIER 0 W 1965 bionomics pages 5375 37 in the texas unpublished thesis texas aam univer- black tailed jackrabbitjackrabbit in kansas kansas state sity college station 115pp115 appp university agricultural experiment station technical bulletin 140 75 appp KEITH L B J R cabyCARYGARY 0 J RONGSTAD AND M C britt- VORHIES C T TAYLOR ingham 1984 demography and ecology ofaof a de- AND W P 1933 the life histories clining snowshoe hare population wildlife mono- and ecology ofofjackonjackjack rabbits lepus allenialbeni and lepus graphs 90 43 appp californicuscalifornicus sppapp in relation to grazing in arizona university of KNOWLTON F F P E MARTIN ANDANDJJ C HAUG 1968 A arizona agricultural experiment telemetric monitor for determining animal activ- station technical bulletin 49 467 587 WAGNER ity journal ofwildlifeofwildlife management 32 943 948 F H 1981 role lagomorphslagomorphaoflagomorphsof in ecosystems pages myers lechleitner R R 1958a movements density and mor- 668 694 in K and C D macinnes eds 1 proceedings tality in a black tailed jackrabbitjackrabbit population jour- I of the world lagomorph con- nal management 371 ference 12 16 august 1979 university ofguelphofguelph of wildlife 2237122.37122 384 press 1958b certain aspects of behavior of the black guelph ontario canada WHITE G GARROTT tailedjackrabbitjackrabbit american midland naturalist 60 C AND R A 1990 analysis ofofwildwild- life press 145 155 radio tracking data academic inc san california 383 MILLS L S ANDFAND F F KNOWLTON 1989 observer perfor- diego appp wywialowski KNOWLTON mance in known and blind radio telemetry accu- A P AND F F 1983 effects racy tests journal of wildlife management 53 of simulated radio transmitters on captive black 340 342 tailed jackrabbitsjackrabbits proceedings of the international conference on Bio NATIONAL OCEANIC AND atmospheric administration wildlife telemetry 4 1984 climatological data utah national oceanic 1 11 and atmospheric administration asheville north received 1 april 1990 carolina accepted 11 september 1990 great basin naturalist 503 1990 appp 257 264

HUMPBACK CHUB cilaGILAGJLA CYPHA IN THE YAMPA AND GREEN RIVERS DINOSAUR NATIONAL MONUMENT WITH observations ON ROUNDTAIL CHUB G ROBUSTA AND OTHER SYMPATRIC FISHES

catherine A karpi and harold M tyusltyus1tyuse ABSTRACT we evaluated distribution habitat use spawning and species associations ofthe endangered humpback chub atlagilagtla cypha inm the yampa and green rivers dinosaur national monument from 1986 to 1989 adult andjuvenile humpback chub were captured in high gradient reaches of yampa and whirlpool canyons where they were rare n 133 1 of all fish captured the fish was primarily captured in eddy habitats in association with 7 native and 12 nonnativenormative fish species roundtail chub G bustarolobustarobusta were widely distributed in eddies pools runs and fifflesriffles humpback chub n 39 and roundtail chub n 242 in reproductive condition were sympatric in eddy habitats during the 5 6 week period following highest spring runoff river temperatures at this time averaged about 20 C nonnativenormative channel catfish ictalurus punctatuspunctatus were abundant in eddies yielding humpback and roundtail chubs suggesting a potential for negative interactions between the native and introduced fishes the humpback chub gila cypha a large roundtail chub gila robusta are sym- river cyprinid endemic to the colorado river patric with humpback chub in DNM but are basin of western united states is federally more widely distributed and more abundant protected by the endangered species act of banks 1964 vanicek et al 1970 holden and 1973 the fish persists only in isolated loca- stalnaker 1975 miller et al 1982 the fish is tions including canyon reaches in the little not considered threatened or endangered un- colorado and mainstream colorado rivers der the endangered species act of 1973 re- arizona kaeding and zimmerman 1983 up- mains of gila species in indian sites in DNM per colorado river colorado valdez and dating more than 1000 years old leach 1970 clemmer 1982 kaeding et al 1990 green suggest that chub were presumably eaten by and yampa rivers colorado and utah hol- native americans and thus have been present den and stalstalnakerStaInaker 1975a 1975b tyus et al in the area for a long time 1982 and mainstream colorado river utah this study was initiated as part of a larger valdez 1990 all stocks are presumed native program to assess status and habitat needs except in cataract canyon of the colorado of endangered fishes in the yampa river river utah where some fish may be derived tyus and karp 1989 our objectives were from a 1981 stocking of juvenile fish of up- to locate humpback chub in DNM and if per colorado river black rocks parentage successful evaluate habitat use including J valentine personal communication flow and temperature requirements identify distribution and status of humpback chub spawning areas and determine species asso- in the upper green and lower yampa rivers ciationsciations in dinosaur national monument DNM remain poorly documented partly because METHODS canyon bound whitewater habitats are difficult to access and sample the fish is rare and the lower 73673.6 km of the yampa river iei e diagnostic features are not well established yampa canyon deerlodgeDeerlodge park to echo park humpback chub were first reported in DNM fig 1 was sampled weekly from mid may in the 1960s and most captures occurred in through early july 1987 1989 by electrofish the confluence area of the yampa and green ing and angling with native foods eg mor- rivers holden and stalnaker 1970 1975a mon crickets anabrusAnabrus simplex and mega 1975b vanicekvanleek et al 1970 studies in the lopteranlopteran larvae and night crawlers at vari- mid 1970s and early 1980s also noted the ous locations in the water column echo and paucity of the fish in DNM seethaler et al island parks and whirlpool and split moun- 1979 miller et al 1982 tain canyons of the green river werewere sampled

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1- 1 METERS

fig 1 yampa and green rivers colorado and utah showing the boundaries of the study area and dinosaur national monument at least twice each spring in 1987 and 1988 ity use and substrate composition during and lodore canyon green river was sam- low flows pled once each spring survey sampling in- all chubs greater than 85 mm total length cluding trammel netting was conducted TL were identified to species using estab- throughout the monument in july 1986 to lished morphological characters smith et al locate humpback chubs use of trammel nets 1979 douglas et al 1989 we did not evalu- was discontinued after this effort because of ate habitat use of young humpback chub be- trauma to the fish cause we could not reliably distinguish young sampling trips in yampa canyon occurred of the various cilagila species humpback chub at weekly intervals preceding and following greater than 250 mm TL were tagged with first and last capture of ripe fish to insure an uniquely numbered carlin dangler tags for accurate assessment of the humpback chub recapture information ege g growth and move- spawning period our efforts were less inten- ment data sex determination was based only sive in the green river portion of the monu- on expression of eggs or milt from ripe fish ment because earlier sampling had yielded either spontaneously or following manual few adult chubs in these reaches holden and pressure on the abdomen fish with breeding crist 1981 miller et al 1982 sampling pre- tuberclestubtubercledercles but without expressible sex prod- ceded peak flows and was suspended during ucts were considered in reproductive condi- highest runoffrunoffsrunoff22 4 week period depending on tion water year because of sampling problems in Riffifflesrifflesfles small rapids runs eddies pools high water sampling ended each summer and backwaters were sampled because wa- with attainment of base flows approximately ter turbidity precluded visual contact with late june to early july our efforts were re- humpback chub it was necessary to esti- strictstricteded to the spring and early summer be- mate the point of capture physical habitat cause of boat accessibility however two parameters recorded at each humpback chub areas in yampa canyon that yielded hump- capture included water depth temperature back chub in the spring big joe rapid and and substrate type depth was measured vicinity warm springs rapid and vicinity with a calibrated rod gross substrate type was were sampled in september 1989 via heli- described from visual and manual examina- copter and by foot to assess habitat availabil tion and temperatures were obtained with 199019901 HUMPBACK CHUB IN DINOSAUR NATIONAL MONUMENT 259 hand held thermometers methods after n 130 and whirlpool n 3 canyons fig nielsen and johnson 1983 we did not at- 1 the whirlpool canyon fish were captured tempt quantification of water velocities be- in the same location about 6 km downstream cause most humpback chub were captured in of the confluence with the yampa river no habitats where water currents swirled in both other humpback chub were captured in the upstream and downstream directions and ini- green river humpback chub constituted tial efforts with a flow meter yielded a wide 737.3 n 51 of the standardized angling and range of positive upstream and negative 1 n 58 of the standardized electrofish downstream velocities habitat use data was ing catch they were most abundant 85 of not recorded for species other than humpback all humpback chub captures n 113 in the chub river flows were obtained as daily aver- upper 44844.8 km of yampa canyon a moder- ages from the USU S geological survey gauging ately steep gradient section 32323.2 mkmmam domi- station at Deerdeerlodgelodge park yampa river fig nated by rocky runs riffles and rapids lower yampa canyon 288 1 stream gradient was obtained from US km 0 28828.8 a lower gradient geological survey stream profile maps system io10101.0 mkmmam consisting mostly of long deep runs and incised meanders yielded rela- data analyses tively few humpback chub n 17 capture data were analyzed by total catch during spring runoff humpback chub were all years all gear types and sampling and most often captured in larger shoreline eddies 20 100 m2ma that standardized catch 1987 1989 catch of all were either downstream of boulders of rapids smaller species and effort recorded for each sample or upstream or in eddies 20 m2ma within shoreline total catch data were used to describe general runs adult fish 230 mm TL based capture of the fish distribution and standardized data were on smallest ripe fish a 232 mm TL male were used to evaluate relative abundance stan- consistently captured and apparently se- dardizeddardized catch data summed by in were gear lected seasonally flooded shoreline eddies ie angling or electroelectrofishingfishing and for each formed and by yampa ie maintained spring river reach ie lodore whirlpool runoffrunofi these habitats were dominated by and split mountain canyons echo and island low or negative water velocities and influ- parks and catch per effort was cal- unit cfco enced by river surges iei e water velocities at culated by dividing numbers of fish captured any particular point varied in magnitude of by effort angling and electroelectrofishingfishing data from up and downstream currents substrate con- 1986 were not included in cf estimates be- sisted mostly of sand and boulders and water cause numbers of fishes other than chubs depth averaged 1131.33 m at the estimated point of were not recorded and because of significant capture humpback chub were not collected differences in angler ability trammel netting in fifflesriffles and rapids cf was not reported because of limited use eleven of 76 carlin tagged humpback chub electrofishingelectrofisbingElectro fishing was biased toward catch of xY 312 mm TL SD 19 were recaptured larger individuals and small fishes eg non- one week to two years after initial capture 5 native reside shiner richardsoniusRichardsonius balteabaldea within a year 6 from one to two years ten tus and native mottled sculpin cottus sppapp fish were recaptured in the immediate vicin- and juveniles of larger species were not ity of their original capture and one was col- recorded because they often slipped through lected about 8 km downstream from its initial our lin1 ininain2 mesh dip nets angling efforts in capture site eight fish 73 n 11 were september 1989 were excluded from the cf recaptured in breeding condition on at least estimates because this effort represented a one occasion we detected no growth in re- unique fall sample sampling was initiated captured fish late in 1986 and those data were excluded about 22 n 29 of humpback chub were from our evaluation of spawning period juveniles 88 228 mm TQTL these were most often captured by electrofishingelectrofishing in rocky RESULTS shoreline runs and small shoreline eddies one juvenile 122 mm TQTL was taken from and use distribution habitat the stomach of a 61 cm TL garter snake HUMPBACK CHUB humpback chub were thamnophis species caught at the conflu- collected only in whitewater reaches ofyampa ence of the yampa and green rivers 260 CcakarpandhmtyusA KARP AND H M tyus volume 50

ROUNDTAIL CHUB A total of 1482 round- catedbated both were most abundant in split tail chub were captured in all reaches ofdnmof DNM mountain canyon tables 1 2 except split mountain canyon and the upper during september 1989 flows in yampa 29 km of lodore canyon the fish constituted canyon were reduced to less than 2832.83 msm3s 37 n 256 of the standardized angling and and fish habitat was limited to shallow fifflesriffles 15 n 1016 of the standardized electro about 15 cm depth and deeper pools and fishing catch roundtail chub were at least runs about imI1 m depth on september 7 we three times more abundant in yampa canyon collected five chubs four roundtail and one than in the DNM portion of the green river suspected roundtail x humpback chub hy- tables 1 2 and were most prevalent in the brid and seven channel catfish in pools and upper 44844.8 km of yampa canyon 73 of eddies about I1 in deep in big joe rapid kmkin all roundtail chub captures n 1085 the 38438.4 other chubs including a suspected fish was incidental in lodore canyon 1 humpback chub were observed about 0800.88 kinkm n 3 adults and juveniles were most often upstream in a 111 I m deep pool created by captured in eddies pools and shoreline runs shoreline boulders no fish were observed or but they were also taken in fifflesriffles and lower collected in the vicinity of warm springs portions of rapids rapid km 646.4 on september 14 species associations of humpback chub spawning of humpback chub humpback chub were captured in associa- and roundtail chub tion with 7 native and 12 nonnativenormative fish spe- thirty nine humpback chub 16 ripe males cies numbers of native stulpinssculpinssculpins and non- 5 ripe females and 18 tuberculate but nonripe native redsidebedside shiners not recorded species fish were captured in shoreline eddy habitats that dominated the standardized catch inin a 48 km reach km 20820.8 68868.8 in yampa cluded flannelmouth sucker catostomus canyon n 37 and in a 2 km reach kinkm latilatipinnispinnis blueheadbluebeadbluehead sucker C discobolus 5456545.6 5472547.2 in whirlpool canyon n 2 roundtail chub common carp cyprinus car- turbidity precluded direct observation of the pio and channel catfish ictalurus buncpunc fish thus spawning behavior and microhabi tatus tables 1 2 tat use were not documented A total of 350 fish were captured by angling all ripe fish were silvery colored with gold in eddies occupied by humpback chub flecks on the dorsum ripe males always had roundtail chub composed about 45 chan- some orange coloration on the lower side of nel catfish 35 and humpback chub 15 the head operclesopopercledercles abdomen and paired and of this catch more channel catfish were cap- anal fin bases ripe males and females usually tured by angling than was any other species bore light tuberculation on portions of the n 328 47 ofangling catch and it was the head nuchal hump opoperclesopercledercles and paired fins most abundant nonnativenormative fish in eddies that this tuberculation was more robust in males also yielded humpback chub other species ripe males averaged 311 mm TL n 16 SD including colorado squawfish ptychocheilus 35 range 232 370 mm and 229 g n 14 lucius flannelmouth sucker common carp SD 67 range 130 348 g ripe females aver- black bullhead ameiurus melas and rain- aged 300 mm TL n 5 SD 20 range bow trout oncorhynchus mykiss composed 280 333 mm and 230 g n 4 SD 75 less than 5 of the angling catch electrofish range 160 336 g and nonripe tuberculate ing catch was dominated by flannelmouth fish averaged 303 mm TL n 18 SD 35 n 2049 29 and blueheadbluebeadbluehead n 1801 range 232 382 mm and 203 g n 17 SD 26 suckers and these fishes were common 62 range 92 356 g in canyon habitats table 1 and open parks ripe humpback chub were collected table 2 following highest spring discharges from the most abundant introduced fishes in mid may to late june 1987 to 1989 table 3 DNM were common carp n 1321 and fig 2 captures of nonripe but tuberculate channel catfish n 1153 these species fish also occurred within this 5 6 week period were relatively common in canyon bound table 3 although sampling in 1986 did not whitewater reaches and lower gradient slow include preprerunoffrunoff conditions and thus was ex- water sections standardized cf data indi cluded from figure 2 four humpback chub in 1990 HUMPBACK CHUB IN DINOSAUR NATIONAL MONUMENT 261

TABLE 1 total catch N and catch per unit of effort offishes collected by standardized angling AN and electrofish ing EL 1987 1989 yampa lodore whirlpool and split mountain canyons dinosaur national monument total effort in hours spent angling angler hours and electroelectroflshingelectrofishingfishing

split yampaYampa lodore whirwhirlpoolipoolippol mountain canyonC nyon canyon cancanyonyon canyon species N AN EL EL AN EL EL native species flannelmouth sucker 2159 030 2894 2472 000 2782 209220 92 blueheadbluebeadBluehead sucker 1812 001 2235 1443 000 3196 8367 roundtail chub 1238 325 IT170909 028 102 266 000 humpback chub 109 065 103 000 023 000 000 colorado squawfish 27 001 030 028 000 053 102 razorback sucker 4 000 007 000 000 000 000 mountain whitefish 2 000 002 000 000 013 000

introduced species common carp 1100 024 1506 1094 023 919gig9.199 19 2551 channel catfish 1091 401 964 1791.791 79 261 826 7194 trout 277 001 016 2226 000 386 102 black bullhead 31 011 021 009 068 04 051 northern pike 15 000 0180.180 18 000 000 0130.130 13 204 white sucker 13 000 0140 14 018 000 173 000 smallmouth bass 6 000 011 000 000 000 000 green sunfish 1 000 002 000 000 000 000 total fish 7885 653 5349 795 42 641 405 total efforteffortseffort1effortb1 76 56 11 9 8 2 includes rainbow cutthroat brown and lake trouts broun&drounded

TABLE 2 total catch N and catch per unit of effort of TABLE 3 capture dates of humpback and roundtail fishes collected by standardized electrofishingelectrofishing ELEQ chubs in reproductive condition yampa and green riv- 1987 1989 island and echo parks dinosaur national ers dinosaur national monument 1986 1989 monument total effort in hours spent electroelectrofishingfishing tuberculate males females fishhish b island echo ripe ripe park park humpback chub species N EL EL 1986 jul 5 15 juljuijulajul55 1987 may20may 20 jun29jun 29 may 18 jun 16 may 18 jun 22 native species 1988 jun 7 28 jun 15 jun 6 15 flannelmouth sucker 185 2637 2634 1989 jun 7 may 27 jun 6 bluebeadBluehead sucker 145 2154 1995 roundtail chub roundtail chub 42 322 8188.188 18 1986a1986s juljuijulg6 29 mountain whitefish 1 032 000 1987 may 18 jun 20 maylamayl7may 17 jun23jun 23 may 17 jun 29 introduced species 1988 jun 7 jul 5 jun 16 jun 7 29 1989 may 27 jun 7 jun 20 may 22 jun 20 common carp 125 1929 1662 no channel catfish 55 sampling prior to july 5 in 1986 119 46 fuberculatetuberculate fish were not ripe but exhibited secondary sex characters trout 16 064 358 black bullhead 2 000 051 northern pike 1 000 026 breeding condition two of each sex were col- white sucker 1 032 000 lected in july of that year ripe fish were green sunfish 2 032 026 captured at water temperatures of about 19519.5 C 145 C total fish 575 261 314 range 14514.5 23 Q total efforteffortseffortbeffort1111 3 4 roundtail chub in reproductive condition includes rainbow cutthroat brown and lake trouts n 242117242 117 males 6 females and 119tuber119 tuber broundedbRoungroundedroundedded eulate but nonripe bshfish were darker than 262 C A KARPKABP AND H M TYUS volume 50

most prevalent in and presumably selected 300 eddy habitats in moderate to steep gradient reaches whereas roundtail chub were ubiqui- tous in pparksarks and most canyons in eddies 200 riffles and runs both fishes were most abundant inm yampa canyon neither was captured in split mountain canyon and the humpback chub was absent and roundtail a too10000 the chub rare in lodore canyon the paucity of colorado river chubs in split mountain and lodore canyon reaches

1 I I1 indicates a general decline ofofgilagila species rel- MAY J my ative to earlier decades e g banks 1964 MONTH vanicek et al 1970 holden and stalStaIstalnakernaker 1975a this may be related to the loss of fig 2 relationship between averageaveiaver age distribution hhy-Y drodrographgraph and spawning period for humpback and round- historic temperature and flow regimes due to tail chubs yampa river 1987 1989 dashed vertical regulated flow releases from flaming gorge lines delineate first and last capture of ripe humpback dam and to the proliferation of nonnative chub solid vertical lines delineate first and last capture of fishes particularly channel catfish and com- ripe roundtail chub 1986 not included because sampling mon carp current rarity of colorado was initiated late in spring runoff the river chubs in split mountain canyon was also noted by the authors min 10 hours ofopporofoppor- humpback chub and exhibited more robust tunistictunistic sampling and by the state of utah tuberculation and more brilliant orange col- during their 1988 89 studies T chart utah oration patterns of tubtuberclestubercledercles and breeding division of wildlife resources personal com- coloration were similar between the two munimunicationcation chubs ripe male roundtail chub averaged capture of 133 humpback chub including 344 mm TL n 117 SD 24 range 292 419 39 breeding adults and 29 juveniles indicates mm TL and 329 g n 100 SD 84 range that a reproducing population exists in yampa 190 652 g and ripe females averaged 363 canyon however only one ripeapenpe fish a male mm TL n 6 SD 15 range 343 380 mm was collected inm the green river iei e whirl- TQTL and 363 g n 3 SD 104 range pool canyon and it is unknown whether it 276 478 g nonncenonripenonnpe tuberculate fish aver- spawned there or was a stray from the yampa aged 351 mm TL n 119 SD 29 range river collection of ripe roundtail chub inm 264 447 mm TQTL and weighed about 364 g n canyon reaches yielding ripe humpback chub 77 SD 123 range 140 844 g indicates some temporal and spatial overlap in ripe roundtail chub were captured in pools habitat use during the spawning period as and shoreline runs and eddies during the observed by others in the upper colorado period of declining spring runoff fig 2 river kaeding et al 1990 humpback and roundtail chubs in breeding ripe humpback and roundtail chubs were condition were collected topicallysynsyntopicallysyntonically on 13 collected during declining spring flows and occasions although this indicated overlap increasing river temperatures after highest in use of shoreline eddies during spring spring runoff this occurred in may and june runoff ripe females of both species were in low- ege g 1987 1989 and average ege g not syntopicsyncopicsyntopic 1988 flow years but extended into july in the 1986 high flow year no humpback chub in discussion breeding condition were captured during pre runoff and late postpostrunoffpostronoffposrunofftronoff periods and we humpback chub and roundtail chub were presume the fish spawned only during the sympatric in DNM in the reach from upper 5 6 week period following highest spring yampa canyon to upper whirlpool canyon flows capture of only a few ripe female although humpback chub were rare 1 I1 chubs five humpback and six roundtail chubs of total catch and only 8 of the two gila 4 of all breeding captures suggested that species combined humpback chub were females may be ripe for a limited time ripe 1990 HUMPBACK CHUB IN DINOSAUR NATIONAL MONUMENT 263 humpback chub were captured at tempera- holden and stalstalnakerStaInaker 1975a tyus and tures Yx 19519.5 range 14514.5 23 QC that minckley 1988 tyus and nikirk 1990 indi- approximate optimum egg incubation condi- cate a potential for negative interactions be- tions ie 20 C marsh 1985 these tempera- tween these fishes although the incidence of tures are similar to the 14 24 C range noted predation by channel catfish on native fishes is by kaeding et al 1990 but slightly higher unknown observations of bitbitelikeelike abrasions than the lisils11511.5 16 C temperatures noted by on some chubs collected in DNM suggest valdez and clemmer 1982 both in the up- channel catfish predation because no other per colorado river piscivorous fish in that system could have all humpback chub and most roundtail caused such damage humpback chub re- chub in breeding condition were captured in mains were found in channel catfish stomachs shoreline eddies our recapture data indicate from the little colorado river W L minck- that adult humpback chub remain in or near ley personal communication and channel specific eddies for extended periods and that catfish are known to consume fish fish parts they return to the same eddy during the and eggs in DNM tyus and nikirk 1990 spawning season in different years they ie only a few common carp were captured syn exhibit a fidelity to a specific site of the ten topically with humpback chub however we I111 I1 recaptures were captured in the same eddy speculate that their abundance may also have as the initial capture 50 in two different some negative impact on the native fishes spawning seasons and 73 were captured in due perhaps to predation on breeding condition at least once we do not eggs know whether these fishes deposited eggs in the humpback chub persists in only a few these eddies or used such habitats only for canyons in the colorado river basin and staging resting or feeding however we planned water development projects may fur- consider the use of such habitats as part of the ther jeopardize its survival the yampa river breeding requirements of humpback chub in in DNM supports all native fishes known to the yampa river shoreline eddy habitats in have occurred there including the endan- yampa canyon were ephemeral ie disap- gered humpback chub colorado squawfish peared with declining summer flows and it and razorback sucker xyrauchen tetanustexanustexanus was obvious that the fish moved elsewhere existing flows of the yampa river may be after the spawning period our observations singularly responsible for enabling the persis- of gila species in pools near big joe rapid in tence ofchubs in the yampa and green rivers september 1989 suggest that some fish re- alteration of yampa river flows could reduce main in nearby deep habitats during low flow the availability or character of chub spawning periods habitat and presumably adversely affect their feeding habits of humpback chub are not reproduction aid in further proliferation of well known and were unknown in DNM cap- introduced competitors and predators and ture of some fish in the interfaces between reduce the quality and quantity of usable shoreline eddies and adjacent runs suggests habitats dinosaur national monument should that chubs use these areas for feeding on drift be considered a refugiumrefugium for native fishes stomachs of two humpback chub that died in and efforts should be made to protect flows of trammel nets contained hymenopterhymenopteransans and the yampa river plant debris and gross examination of fecal material taken from live fish indicates exten- acknowledgments sive use of hymenopterhymenopteransans and other terrestrial insects eg mormon crickets as food this study was funded in part by U S fish we observed humpback chub and other fishes and wildlife service bureau of reclamation eg roundtail chub common carp feeding national park service and the northern col- on mormon crickets at the water surface in orado water conservancy district J beard eddies P clevenger and L trinca were among the high numbers of channel catfish in several who assisted with data collection habitats used by humpback chub and round- P B marsh C 0 minckley and W L tail chub and the gross overlap in foods minckley improved an earlier draft of the consumed and in feeding habits banks 1964 manuscript 264 C A KARP AND H M TYUS volume 50

literature CITED SEETHALERSEETHALEB K H C W MCADA AND R S WYDOWSKI 1979 endangered and threatened fish in the BANKS J L 1964 fish species distribution in dinosaur yampa and green rivers of dinosaur national national monument during 1961 1962 unpub- monument pages 605 612 in proceedings of the lished thesis colorado state university fort first conference on scientific research in the collins 96 appp national parks vol 1 national park service DOUGLAS A E W L MINCKLEY AND H M TYUS 1989 transactions and proceedings series 5 qualitative characters identification of colorado SMITH G R R R MILLER AND W D SABLE spe- gila 1979 river chubs cyprinidae genus and the cies relationships among fishes oftheodtheofthe genus gila in of well 19896531989.6531989 653 662 art seeing copela the upper colorado river drainage USU S national HOLDENHOLDENPP BBANDLAND L W CRISTCBIST 1981 documentation of park service transactions proceedings series 5 changes in the macromacroinvertebrateinvertebrate and fish popula- 613 623 tions in the green river due to inlet modification of flaming gorge dam BIOWEST PR 16 5 TYUS H M AND C A KARP 1989 habitat use and logan utah streamstreamflowflow needs of rare and endangered fishes yampa HOLDEN P B AND C B STALNAKERSTALNAKEB 1970 systematic river colorado US fish and wildlife studies of the cyprinid genus gila inm the upper service biological report 8914 27 appp colorado river basin copela 197019701091970.109109log 120 TYUS H M AND W L MINCKLEY 1988 migrating mor- 1975a distribution and abundance ofmainstrearnofmainstream mon crickets anabrus simplex orthoptera tetti- fishes of the middle and upper colorado river gonigoniidaeidae as food for stream fishes great basin basin 1967 1973 transactions of the american naturalist 48 25 30 104 231 fisheries society 217 TYUS H M AND N J NIKIRKNIKIBK 1990 abundance growth 1975b distribution of fishes in the dolores and and diet ofchannelofchannel catfish ictalurus punctatuspunctatus in yampa river systems of the upper colorado the green and yampa rivers colorado and utah basin southwestern naturalist 1940319.40319 403 412 southwestern naturalist 35 188 198 KAEDING L R AND M A ZIMMERMAN 1983 life history of the humpback chub in the little colorado and TYUS H M B D BURDICK R A VALDEZ C A HAYNES colorado rivers of the grand canyon transac- T A LYTLELYTLEANDCAND C R BERRY 1982 fishes of the tions of the american fisheries society 112 upper colorado river basin distribution abun- 577 594 dance and status pages 127012 70 in W H miller H M C A eds KAEDING L R & D BURDICKBURDICKPP A SCHRADER AND C W tyus and carlson fishes oftheodtheofthe RB schraderandc upper mcada 1990 temporal and spatial relations be- colorado river system present and future tween the spawning ofhumpbackofhumpback chub and round- american fisheries society bethesda maryland tail chub in the upper colorado river trans- VALDEZ R A 1990 the endangered fish of cataract actions of the american fisheries society 119 canyon final report biowestBio West inc logan utah 135 144 VALDEZ R AANDGA AND G H CLEMMERCLEMMFR 1982 life history and LEACH L L 1970 archaeological investigations at de- prospects for recovery of the humpback chub and luge shelter in dinosaur national monument bonetailbonytailbonytail chub pages 109 119 in W H miller unpublished dissertation university ofcoloradoofcolorado H M tyus and C A carlson eds fishes oftheodtheofthe boulder 336 appp upper colorado river system present and future MARSH P C 1985 effect of incubation temperature american fisheries society bethesda maryland on survival of embryos of native colorado river fishes southwestern naturalist 303012930.129129 140 VANICEK C D R H KRAMER AND D R FRANKLIN 1970 green fishes MILLER W H D L ARCHER H tyus AND R distribution of river in utah and A A gorge MCNATT 1982 yampa river fishes study final colorado following closure of flaming report US fish and wildlife service and na- dam southwestern naturalist 14297 315 tional park service cooperative agreement 14 16000681931600068193116 0006 81 939311 salt lake city utah 107pp107 appp received 3 april 1990 NIELSEN L A AND D L JOHNSON 1983 fisheries tech- 17 niques american fisheries society bethesda revised august 1990 maryland 468 appp accepted 6 september 1990 great basin naturalist 503 1990 appp 265 272

management OF endangered SONORAN TOPMINNOW AT BYLAS SPRINGS ARIZONA description CRITIQUE AND recommendations

paulcpaul C marshmarshi and W L Minckley 2

ABSTRACT efforts between 1982 and 1990 have failed to recover and secure three natural populations of endangered sonoran topminnowtopminnow poeciliopsispoecihopsts 0 occidentalisoccidentoccidentalistalis at bylas springs arizona flooding in the gila river in 1977 78 allowed ingress by predatory mosquitmosquitomosquitoishmosquitobshmosquitofishBshofish Gamgambusiabusta afalfinisaffinisfinis and topminnowstopmmnowstopminnows began to decline since that time 1 one stock has been replaced twice and is again nearly gone because of depredations by mosquitmosquitofishofish that resisted two eradication attempts 2 toptopminnowstopmmnowsminnows at a second spring were extirpated through vegetation encroachment after fencing to protect the habitat from livestock and 3 a third population was lost to mosquitmosquitofishofish restocked after the nonnativenormative was removed and the restocked population is again in jeopardy or extirpated since mosquitmosquitofishofish re invaded recommendations for a more intensive program of recovery are based on reassessments of past efforts and new suggestions for eradication and exclusion of mosquitmosquitofishofish the sonoran gila topminnowtopminnow poecilioppoecifiop with west and eastcast sources through siliSIIIS iliIII111 sis 0 occidentoccidentalisoccidentalistalis is a poeciliid fish endemic west to east and provide a synonymy below to and once widespread in the gila river basin descriptions andor mention of the springs of arizona and new mexico USA and the have appeared in papers noted above and rios gila Concepcconcepcionfon and sonora basins of others by meffe 1983a 1983b meffe and sonora mexico hubbs and miller 1941 marsh 1983 hendrickson and minckley minckley 1973 vrijenhoek et al 1985 it was 1985 williams et al 1985 taylor 1987 listed in 1967 as an endangered species and hershler and landye 1988 US fish and wildlife service USFWS all the springs are small with base flows of 1989 because of predation by introduced a few liters slisllsiiSIIS II11 middle spring which mosquitmosquitofishofish gambusia afalfinisaffinisfinis and habitat dried temporarily in 1990 to a few tens of degradation in the gila river basin miller liters per minute S 1 with west and east 1961 meffe et al 1983 meffe 1985 most sources bos and medicine springs respec- efforts to recover the sonoran topminnowtopminnow tively siliSIIIS iliillliiIII111 salt creek siliSIIIS lililiIII111 has the great- have emphasized reintroductions within its est base flow S 1 and sllslisilSIIS II11 rise through alluvial native range USFWS 1984 this paper deals fill along a stony escarpment they have essen- only with attempts to maintain its natural pop- tially no surface watersheds other than their ulationsulations which in the USA are now restricted immediate surroundings SIIIS III rises in the to fewer than 10 sites all in arizona channel ofan otherwise ephemeral watershed ofof5050 kmburkhamkmkm2 burkham 1976a all are intermit- STUDY AREA tent in their lower reaches originally isolated from the gila river either by desiccating on or three natural topminnowtopminnow populations oc- percolating into the terrace that parallels the cupied a series of small springs adjacent to the river s broad floodplainfloodplain SIS I1 and silsllSIIS II11 or by gila river on san carlos apache indian tribal falling over an alluvial escarpment 25252.5 m lands near bylas graham county arizona high siliSIIIS lililiIII111 two were discovered in 1968 johnson and unusually high flooding in the gila river kobetich 1970 and the third in 1981 meffe et in winter 1977 78 afforded an opportunity al 1983 no other fishes were present al- for mosquitmosquitofishofish to invade S 1 and that spe- though collectively known as bylas springs cies plus red shiner cyprinellaCyprinella lutrensislutrensis the habitats gained various coined names in recorded only once also colonized siilSIIIS illiliIII111 sub- the literature for brevity we term them SIS I1 sequent declines in topminnowtopminnow populations

center for environmental studies arizona state university tempe arizona 85287120185287 1201 department of zoology arizona state university tempe arizona 85287150185287 1501

265 266 P C MARSH AND W L MINCKLEY volume 50 were immediate and dramatic in both sys- each new barrier were fenced to exclude tems and various management strategies domestic livestock were planned and implemented in attempts S 1 was again poisoned with antimycin A in to eradicate the introduced fish and restore april and june 1984 and restocked in july with topminnowstopminnows this paper recollects those ef- 200 topminnowstopminnows and uncounted inverte- forts provides a status update on topminnowstopminnows brates removed prior to poisoning all native at bylas springs and offers recommendations animals soon reestablished in large numbers to perpetuate the stocks however mosquitmosquitofisbmosquitofishofish again survived downstream to reinvade above the barrier when it THE SEQUENCE OF EVENTS was bypassed by spate augmented flows in late summer the introduced species consti- quantitative data are lacking on the abun- tuted 24 of total fishes caught in december dance of topminnowstopminnows at the times of their 1985 69 in september 1986 and 98 discovery in bylas springs but these fish by july 1987 simons 1987 in january were present in substantial numbers espe- 1989 topminnowstopminnows n 9 in the fenced cially in summer throughout the systems and west source of SIS I1 were accompanied by in winter in and near the springspringheadsheads after mosquitmosquitofishofish n 6 and only mosquitmosquitofishofish invasion of mosquitmosquitofishofish topminnowtopminnow in SIS I1 were taken downstream the unfenced east between summer 1980 and spring 1982 re source was not sampled three topminnowstopminnows bainedmained in the upper reaches of runs 62 72 and one mosquitmosquitofishofish were collected from the of all fishes present but declined to rare or fenced source in april 1990 while tolmintopmin absent downstream where mosquitmosquitofishofish were nows composed 44 of 189 fish taken from the numerous A parallel situation existed in unfenced source no topminnowstopminnows were found silisillSIIIS illiliIII111 slisllSIIS II11 was never invaded by mo in the pool above the barrier or in the stream squitsquitofishofish and topminnowstopminnows remained com- below where mosquitmosquitofishofish were abundant mon until 1988 in summer 1989 in anticipation of another SLS L1 in march 1982 SIS I1 was poisoned with poisoning which has not yet occurred the antimycin A meffe 1983b no live fish were channel of SIS I1 was realigned to again flow over found three weeks later and 150 tolmintopmin its barrier and low 05050.5 m high babionsgabions of nows along with large numbers of indigenous wifewire constrainedconeonstrained and loose rock were ex- invertebrates salvaged prior to poisoning tended for 19 24 m on either side to replace were reintroduced in the springspringheadsheads by the earthen bermsbarms in summer 1990 notches july 1982 substantial populations of both top in all the barriers were reshaped from the minnows and invertebrates were reestab- original YV to approximately rectangular lished at the same time however mosquito doubling their areas to 1500 cm2 to accommo- fish were discovered downstream since the date larger flows the upstream base of the gila river had not again flooded poisoning barrier at SIS I1 was also sealed with bentonite to must have failed to remove them from the prevent seepage under the structure which lower part of the system meffe 1983b rec- had become substantial and a large saltcedarsalteedarsaltsaitsaltesaiteedarcedar ommendedommended massive repeated poisoning com- tamarix chinenchinensissis that threatened damage bined with construction of barriers as a possi- was removed ble method of maintaining the topminnowtopminnow slisiiSHSJJS HJJ this spring experienced no dis- both strategies were embraced in the species charge that passed over the barrier and re- recovery plan USUSFWSFWS 1984 mained secure from mosquitmosquitofishofish neverthe- barriers intended to prevent upstream less topminnowstopminnows disappeared in 1990 due to movements of mosquitmosquitofishofish were constructed surface water depletion invasion by cattail on the runs of all three springs during winter typha angustifolia after livestock exclusion 1983 84 each consisted of low 07070.7 080.8 m discussed below resulted in increased evajoevapo concrete V notch weirs with earthen wings transpiration and accumulation of vegetative at SIS I1 and sillsiliSIIIS liilililiIII111 extending laterally and up- debris and entrained sediment that dried the stream for 353.5 666.6gg6 6 m no bermsbarms were con- system strucstructedted lateral to the barrier at slisllSIIS II11 in addi- siiliSIIIS III topminnowsTopminnows were extirpated from tion the western source of SIS I1 and the single SIIIS liililIII111 by 1984 A flood in the salt creek water- source of slisllSIIS II11 plus the areas surrounding shed in 1983 incised the lower part of the run 1990 TOPMINNOW management 267

to destroy the former waterfall the intermit- major habitat changes occurred as a result tent lower part of the spring run now led into of barrier construction and fencing in the by- the river over a gentle cataract that should las springs system all the runs begin at a have posed no reasonable deterrent to fish relatively low gradient which is reduced even passage despite this the reach above the further as they pass onto the gently sloping artificial barrier remained fishless from the surface of the gila river floodfloodplainplain this pre- time mosquitmosquitofishofish were poisoned in april cluded construction of high barriers without 1984 until 300 topminnowstopminnows from silslisllSIIS II11 were extensive concrete work or excavation for stocked in 1986 brooks 1986 topminnowsTopminnows long lateral bermsbarms the structures decided became abundant and no exotic fish was cap- upon were placed as far downflow as practica- tured in 1986 1987 or winter 1988 the run ble ca 400 and 200 m downstream from the flooded around the barrier sometime in 1988 respective west and east sources in S 1 60 m and mosquitmosquitofishofish again appeared in spring in sllsliSIIS II11 and 575 m in siliSIIIS iliIII111 to protect maxi- summer 1989 by april 1990 mosquitmosquitofishofish mum lengths of spring runs all barriers are constituted 76 of all fishes collected above 700 m or more from the gila floodplainfloodplain none the barrier and were abundant downstream spanned the entire flood channel however where only a single topminnowtopminnow was captured and those at SIS I1 and sillsiilSIIIS illliililIII111 failed to direct high discharges over the concrete no topminnowstopminnows were observed in siliSIIIS lilliiIII111 in weirs since sur- july 1990 face runoff from one or more precipitation around in 1989 in anticipation of future renova- events cut them berm replacement by longer rock babionsgabions in 1989 again failed tion and as at S 1 lateral babionsgabions of wire at S 1 constrained and loose rock were extended where surface flow bypassed the barrier in july 1990 10 11 m on either side of the structure and the barriers also created small ponds on the the channel of S III was realigned to again SIII low gradient runs concern existed that such pass over its artificial barrier ponds might enhance mosquitmosquitofishofish and the exotic species did in fact quickly expand its discussion population as soon as the lentic habitat was achieved however because they had already the inability to secure populations of the invaded prior to the presence of ponds the endangered sonoran topminnowtopminnow at bylas question was moot the pond habitats were springs is disturbing given that nearly a transitory anyway sedimentation was exten- decade has passed since management efforts sive and all three were quickly invaded by moreover began of the 10 known natural catcattailscoattailstails by 1990 emergent vegetation was so stocks of sonoran topminnowstopminnows remaining in dense above the barriers on SIS I1 and siilSIIIS liililillIII111 that the gila river basin including SIS I1 and silsliSIIS II11 open water scarcely existed cattail stands the latter now extirpated 6 were sympatric had trapped even more sediment so that by- with mosquitmosquitofishofish in 1987 simons 1987 and passing by floodwatersfloodwaters may have been forced expected to disappear in the near future in part by boundingmounmoundingding of vegetative debris and there is little doubt that two of the three silt upstream from the weirs given sufficient populations at bylas springs would already be time accumulations of cattailscatcoattailstails became so ex- gone had mosquitmosquitofishofish not been partially con- tensive that we are convinced the barriers trolledtrolled but the facts remain that 1 a tolmintopmin would have been clogged and breached with- now stock at S 1 although removed and re- out high water placed twice is again nearing extinction in january 1989 the barrier pool at slisllSIIS II11 through depredations by mosquitmosquitofishofish that re- held only small pockets of water 20 30 cm in sisted two attempts to remove them 2 a diameter and scarcely a centimeter deep native stock at sllslisilSIIS II11 has been extirpated several dozen topminnowstopminnows had died in these through encroachment of vegetation after pockets likely due to combined low tempera- fencing to protect it from livestock and 3 one ture and oxygen depletion in the thin layer of population sillsiliSIIIS iliillliiIII111 was lost to mosquitmosquitofishofish water overlying organic sediments the popu- necessitating restocking from silsliSIIS II11 after the lation had dwindled to a few individuals and a nonnative was removed and the restocked single adult male was caught in april and july population is again in jeopardy or extirpated 1990 the habitat was only moist no fish were since mosquitmosquitofishofish reinvaded found despite exhaustive examination 268 P C MARSH AND W L MINCKLEY volume 50

A similar sequence occurred after fencing atelyabely funded commitment to maintain the to exclude cattle from the source of slisllSIIS II11 the bylas springs topminnowtopminnow stocks piecemeal headspring and its outflow were rapidly inin- efforts to date have largely failed because hy- vaded by catcattailscoattailstails and by january 1989 the drodrologiclogic and vegetational dynamics and com- plants had formed a virtual mound of living plexity were either not understood or taken and dead vegetative materials with surface into account habitat responses to manage- water only along the margins we speculated ment prescriptions were thus unpredicted that the site would be uninhabitable by fishes A formal plan for recovery must be imple- the next growing season and by april 1990 no mented followed by programmed event surface water was present and the topminnowtopmmnowtopminnOw responsive monitoring for the foreseeable population was gone succession had pro- future ceeded to include invasion of a large bulrush next it is appropriate to define the degree scirpus sp in july 1990 water was present of isolation of the three bylas springs the but fishless and the cattail stands here and at presence of endemic hydrobiid snails mem- the downstream barrier pool were dead for bers of a specialized group restricted to unknown reasons springs in the american west taylor 1987 vegetation in the western source of S 1 re- hershler and landye 1988 indicates consid- sponded differently to livestock exclusion erable antiquity of the habitat these animals marsh and springhead although 5 0 the in are essentially unknown in streams and their in diameter were avoided by livestock pre- in presence in large erosive rivers like the gila sumably due to its dangerously spongy is even less probable presence in S 111 quaking deposits overgrown by their sillsiliSIIIlilliiIII organic which rises within a channel that floods on of small sedges eleo- cienega vegetation occasion is unusual topminnowsTopminnows could have and grasses was charis sp only its periphery colonized the springs at any time heavily grazed over the years minckley un- Mosquitmosquitofishofish were not locally available to published data found three dead cattle mired invade bylas springs until perhaps the 1930s in the center of this tiny marsh in it remains chamberlain 1904 collected none in the saf- similar today inside fence al- its protective ford area and the species was found nowhere though there is a slightly more luxuriant is else in arizona until 1926 when it appeared in growth of the same low vegetation but no the colorado river at yuma and salt river in of catcattailscoattailstails invasion by other than invasion tempe miller 1961 miller and lowe 1967 low especially adapted sedges and grasses invasion progressed rapidly and mosquito be precluded by saturated reduc- may water fish were abundant statewide at low eleva- hydrohydrosoilsbydrosoilssoilssolls as suggested by hendrick- ing tions by the 1940s minckley 1973 topmin son and minckley 1985 interestingly the tolmin nows were thus protected for about 40 years spongy marsh has now solidified and no longer until flooding in 1977 78 was sufficient to per- appears dangerous to livestock currently mit ingress by the nonnativenormative species recon- fish only a hmnocrenelimnocrene the habitat comprises structionst of the history of these habitats helps 50 long wide and 25 deep ca cm 25 cmem cm understand how and why mosquitmosquitofishofish were outflow and its originally excluded source of S I1 was always larger the eastern SI gila channel 90 in width in than any but salt and the river in other spring creek the period 1875 1903 was eroded to an aver- so was not fenced and an remains it remains age of 600 in in width in 1905 17 burkham flowing limnocrenehmnocrene 1.0io10 across 2.020 open 10lomin 2020min 1972 turner 1974 terrace on which long and 25 cmem deep combination of the the bylas springs now occur was not present in wateimggrazmgtramplingwateringgrazingtrampling by live- intense the latter period and the springs were much stock plus human although un- activities nearer or could have flowed directly into the sightly and outwardly to damage appearing river the system precludes overgrowth by river semisemi- during 1905 1906 a cone of coarse allu- aquatic vegetation vium was washed into the gila river channel by flash flooding in salt creek the channel in recommendations which siliSIIIS iliIII111 rises by 1914 the river was being our recommendation is that the USU S fish deflected southward and threatening the and wildlife service make a firm appropriappropnappropre town of bylas US army corps of engineers 1990 TOPMINNOW management 269

1914 olmstead 1919 this was accompanied barriers so that upstream ponds and their by deposition on the north side of the channel associated problems will not exist downstream of the salt creek alluvial cone once mosquitmosquitofishofish are eradicated a barrier as documented by photographic evidence should not be needed on slisiiS 11 forty years of burkham 1972 and indirectly by the sizes of protection by natural isolation would seem an mesquite trees prosopis sp see gavin 1973 acceptable period of time for management of that could only have colonized after the ter- an endangered short lived species such as race was formed see also minckley and clark the sonoran topminnowtopminnow we cannot predict 1984 in part because of saltcedarsaltcedar invasion the recurrence interval of major floods but burkham 1976b the channel again nar- such events some of which have exceeded rowed to average 120 m by 1964 68 the 4000 m3ma secsee 1 in a river that averages 14 m3ma river remained on the south side of its flood seesec 1 at safford burkham 1970 US geologi- plain thus bylas springs were isolated cal survey 1989 cannot be engineered flooding in 1977 78 and again in 1983 contin- against such a flood directed against the ued to erode southward stimulating major north side of the gila river floodplainfloodplain would engineering attempts to stabilize and control destroy the system as it now exists the channel personal observation despite these data and pronouncements since it was unknown whether invasion by mapping of the entire spring complex is mosquitmosquitofishofish in 1977 78 was an isolated event clearly in order for future management refer- or whether the system was changed enough to ence the effort should include aerial pho- assure continued access by exotic fishes we totography at the time of minimum vegetative reexamined it in july 1990 SIS I1 ended as it did development in winter accompanied by ex- before 1977 78 in a variably wetted sump tensive ground truth to confirm intricacies of formed within salteedarsaltcedarsaltsaitsaltesaiteedarcedar thickets on the ter- the aquatic system the extent of aquatic race this sump may exceed 0050.5os 5 ha in area and habitat should be determined under both was heavily utilized by livestock also as be- drought and wet conditions to assure an un- fore there was no apparent outlet to the gila derderstandingstanding of actual and potential intercon- river which lay at least a kilometer farther nections south and considerably lower in elevation ca next mosquitmosquitofishofish must be eradicated an 505.0 m slisllSIIS II11 similarly remained equally as iso- operation which must be preceded by related in 1990 as it was in the recorded past As moval of substantial numbers of native fish noted before the channel into which silisillSIIIS lilliiIII111 and other animals from each spring to secure rises passed unimpeded into the gila river refugia fortunately fish from slisllsilSIIS II11 have al- and was thus accessible to mosquitmosquitofishofish dur- ready been transplanted to a spring in roper ing flood lake state park near safford arizona where As long as mosquitmosquitofishofish exist in the sump they established a large reproducing popula- of S 1 an artificial barrier will be required tion arizona game and fish department but it must be designed to function under all AZGFDAZGFDI unpublished data A stock from but the most severe conditions silsllsliSIIS II11 seems SIS I1 is being similarly maintained at arizona sufficiently isolated without a barrier assum- state university the stock originally inhabiting fish habitat can again be established SIIIS lilliiillIII111 ing siliSIIIS ililiiIII111 is extinct will require a barrier for the foreseeable fu- invertebrates may present a problem es- ture since mosquitmosquitofishofish will continue to oc- pecipeciallyally the two endemic hydrobiid snails cupy the gila river existing weirs especially tryoniabryonia filaegilae and the monotypic apachecoc those for sillSIIIS lililiillIII111 will require further modifica- cus arizonaearizonanarizonae taylor 1987 see also hershler tions to accommodate high discharges or must and landye 1988 which have previously be replaced with structures that will do so been held on site for no longer than a few days the last is difficult because of the gently slop- and in aquaria for about three months ifthese ing surrounding terrain which may necessi- animals are to be held captive for a longer tate bermsbarms extending tens of meters on each period special treatment or facilities may be side each should be equipped with wide required typically invertebrates may be spillspillwaysways of nonerodable material we also reintroduced soon after the poison dissipates recommend installation of soil pipe or some generally a few days after the final application other means of passing base flows through all numbers of all animals retained for stockingrerestocking 270 P C MARSH AND W L MINCKLEY volume 50

should be large enough to assure maintenance could be gated and opened periodically to of genetiegenetic variability and a reasonable proba- allow removal of vegetation by livestock if bility for representation of rare alleles meffe not we recommend the fences be removed 1986 1987 meffe and vrijenhoek 1988 from headheadspringssprings and barrier pools alike considering the negative results of previ- cattail and other emergent plants could ous attempts piscipiscicidecide should be applied re- also be controlled by cutting burning or peatpeatedlyedly perhaps three or more times at chemical herbicidesherbicides but these methods are weekly or longer intervals application should labor intensive and may be more habitat dis- be accompanied by physical dedewateringwatering of ruptive than livestock a situation that should runs lateral pools and the downstream sump be avoided ifpossible experimental manipu- of S 1 if possible footprints of livestock in lations to determine reasonable vegetation muds of the sump provide tiny but effective control measures might be attempted at life support sites in which mosquitmosquitofishofish may sllsilSIIS II11 which is now fishless once tested ac- survive immediately adjacent to toxic water cepceptabletable control techniques could be applied temporary exclusion of livestock might well to other sites alleviate this problem alternatively water could be retained upstream by temporarily conclusions damming the spring run so the sump could be repeatedly watereddewatererdedewatered at least to a degree the current and past recovery efforts in and then flooded to inundate such refugia behalf of sonoran topminnowstopminnows in the gila with poisonous water river basin brooks 1985 1986 simons A fishless period of at least a year should be 1987 simons et al 1989 have emphasized required for the entire system before tolmintopmin introductions and introductionsreintroductionsre within his- nows are restocked presence of mosquitmosquitofishofish toric range far more than dealing with natural in any ofthe springs allows children or adults populations resulting in continuing jeopardy to inadvertently or intentionally move them to natural populations in fact according to from place to place humans are attracted to the current recovery plan USFWS 1984 all springs in otherwise and lands and the bylas natural populations could disappear without springs although reasonably isolated are pe- influencing the down listing or dedelistinglisting crite- ririodically used by local residents for recre- ria and since the criteria have been satisfied ation simons et al 1989 the species could con- in order to assure longtermlong term success the ceivaceivablybly be down listed to threatened status area must be inspected frequently and man- in 1990 the USFWS desert fishes recov- aged to assure maintenance of habitat in- ery team recommended against such action tegrity and continued exclusion of mosquito minckley unpublished data fish and to detect and interdict local land uses we do not underestimate the difficulties that may prove detrimental we recommend associated with habitat renovation for main- a cooperative agreement with the san carlos tenance of native topminnowtopminnow stocks but apache tribe to perform a quarterly or more we suggest a concerted effort be directed frequent schedule of surveillance and moni- toward accomplishing that end at bylas toring biannual intensive surveys should en- springs if topminnowstopminnows cannot be secured list the assistance of a professional biologist here which undoubtedly represents some the entire system is small and may be thor- of the least complex habitats occupied by nat- oughly examined in one day ural populations of topminnowstopminnows in the USA the activities of domestic livestock which it seems highly unlikely that any natural popu- must have precluded overgrowth by cattail in lation threatened by mosquitmosquitofishofish has much the past may be a necessary part of the ecol- hope ofa persisting yet repeated attempts to ogy of these springs encroachment by semi- eradicate mosquitmosquitofishofish have failed and other aquatic vegetation destroys small isolated efforts to manage the habitat have had unde- habitats and this must be avoided the habi- sirable results it is more efficient to devote tat disruption and apparent degradation by necessary planning manpower and material livestock is preferable to loss of the habitats to initial operations even if the assurance of and topminnowstopminnows if reliable close order success is costly than to expend lesser surveillance is developed the existing fences amounts on repeated unsuccessful operations 1990 TOPMINNOW management 271 over time whereby cumulative costs become hendricksonHENDricksonBICKSON D A AND W L MINCKLEY 1985 clenecigne exorbitant and the populations are still lost A gas vanishing aquatic climax communities of the american southwest plants 6 new round of effort is clearly needed and desert 131175 HERSHLER R AND J J LANDYE 1988 arizona hydro quickly biidaeboidae prosobranchia rissoaceaRissoacea smithsonian contributions in zoology 4591459.1459 1 63 HUBBSHUBBSCC LANDRL AND R R MILLER 1941 studies ofthe fishes acknowledgments of the order cyprinodontes XVIIXVIL genera and many species of the colorado river system occasional individuals have been involved in papers of the university of michigan museum of research and management efforts at bylas zoology 4424421944211 9 springs USFWS personnel have included JOHNSON J E AND G KOBETICH 1970 A new locality for gila among others J brooks formerly AZGFD the topminnowtop minnow poeciliopsis occidentoccidentalisoccidentalistalis poeciliidae southwestern naturalist 1436814 368 S jacks D parker B J hansen robertson MEFFE G K 1983a ecology of species replacement in deceased and S stefferud B bagley the sonoran topminnowtopminnow poeciliopsis occidentoccidentalisoccidentalistalis D hendrickson and L simons AZGFD and the mosquitmosquitofishofish gambusia alfinisaffinisaffinis in ari- provided data and assistance as did a number zona unpublished dissertation arizona state of students and others from arizona state university tempe 143 appp 1983b attempted chemical renovation of an university D langhorst G meffe R tim- arizona springspringbrookbrook for management of the en- mons and S vives deserve special mention dangereddangered sonoran topminnowtopminnow north american the san carlos apache indian tribe permit- journal offisheries management 3 315 321 ted access through their wildlife depart- 1985 predation and species replacement in american southwestern fishes a case study south- ment and tribal wardens physically assisted western naturalist 30 173 187 in many operations all deserve special 1986 conservation genetics and the management thanks we of course take full responsibility ofendangered fishes fisheries 11 14 23 for any errors in fact or interpretations 1987 conserving fish denomesgenomesgenomes philosophies and practices environmental biology of fishes 18 3 9 literature CITED MEFFE G K D A hendrickson W L mlnckleyMINCKLEY AND J N RINNE 1983 factors resulting in decline of BROOKS J E& 1985 factors affecting the success of cliacilaclidgilagild the endangered sonoran topminnowtopminnow poeciliop topminnowtopminnow introductions on four arizona national sis occidentoccidentalisoccidentalistalis atheriniformes poeciliidae in forests report to the USU S fish and wildlife ser- the united states biological conservation 25 vice albuquerque new mexico arizona camegame 135 159 and fish department phoenix 35 appp MEFFE G K AND P C MARSH 1983 distribution of BROOKS J E 1986 status ofnatural and introduced sono- aquatic macroinvertebratesmacroinvertebrates in three sonoran ran topminnowtopminnow poeciliopsispoecthopsis 0 occidentoccidentalisoccidentabsoccidentalistabsalisallsails popu- desert springspringbrooksbrooks journal of arid environ- lations in arizona through 1985 report to the ments 6 363 371 USU S fish and wildlife service albuquerque MEFFE G KKANDBAND R C vrijenhoek 1988 conservation new mexico arizona game and fish depart- genetics in the management ofdesert fishes con- ment phoenix 43 appp servation biology 21572 157 169 MILLER 1961 man BURKHAM D E 1970 precipitation stream flow and R R and the changing fish fauna of the southwest papers major floods at selected sites in the gila river american of the michigan academy of drainage above coolidge dam arizona U S ge- science arts and letters 46 365 404 ological survey professional paper 655 B 1 33 MILLER R R AND C H LOWE 1987 part 2 fishes of 1972 channel changes ofthe gila river in safford arizona pages 133 151 in C H ed ver- valley arizona 1848 1970 USU S geological sur- lowe tebrateste ofarizona ad2d printing university of vey professional paper 6556556 G 1 23 arizona press tucson 1976a from small watersheds flow adjacent to the MINCKLEY W L 1973 fishes ofarizona arizona game study reach of the gila phreatophyte pro- river and fish department phoenix 293 appp ject U S geological survey professional paper MINCKLEY W L AND T 0 CLARKCLABK 1984 formation and 6551655655111 1 19 11 destruction of a gila river mesquite bosque com- 1976b hydraulic effects of changes in bottom munity desert plants 6 23 30 land floods gila vegetation on three major river OLMSTEADOLMSTEADFF 11 1919 gila river flood control a report in southeastern arizona USU S geological survey on flood control of the gila graham professional paper river in 655 J 1 14 county arizona US 65th congress ard3rd ses- chamberlain F W 1904 unpublished arizona field sion senate document 4364361436.11 94 notes filed at the smithsonian institution SIMONS L H 1987 status of the gila topminnowtopminnow poe washington DC 52 appp handwritten ciliopsisciliopsis occidentalisoccidentoccidentalistalis occidentoccidentalisoccidentalistalis in the united GAVIN T A 1973 an ecological survey of a mesquite states report to the US fish and wildlife ser- bosque unpublished thesis university of ari- vice albuquerque new mexico arizona game zona tucson and fish department phoenix 36 appp 272 P C MARSH AND W L MINCKLEY volume 50

SIMONS L H D A hendricksonhindricksonhl NDRICKSON AND D papoullasPAPOUIPAPOULIASlasIAS threatened wildlife and plants subpart B lists 1989 recoveiyofthegilatopmmnowrecovery of the gila topminnowtopminnow a success endangered and threatened wildlife CFR 171117.11 story conservation biology 3 1 5 and 171217.12 US government printing office TAYLOR D W 1987 freshwaterfresh water mollusksmolluscsmolluscs from new washington DCD C 25 appp mexico and vicinity bulletin of the new mexico USU S geological SURVEYS URVEY 1989 water resources data for bureau of mines and mineral resources 116 arizona water year october 1987 to september 1 50 1988 US government printing office washing- TURNERTURNLR R A 1974 quantitative and historical evidence ton D C 358 appp for vegetation changes along the upper gila river vrijenhoek R C M E DOUGLAS AND G K MEFFE arizona USU S geological survey professional 1985 conservation genetics of endangered fish paper 655 H 1 20 populations in arizona science 229 400 402 USU S ARMY CORPS OF ENGINEERS 1914 san carlos irriga- WILLIAMS J EK D B BOWMAN J E BROOKSBKOOKS A A tion project arizona USU S 63rd congress 2ndand ECHELLE R J EDWARDS D A hendrickson session house document 791 1 168 AND J J LANDYELAN DYE 1985 endangered aquatic USU S FISH AND WILDLIIWILDLIIEWILDLIFEE SERVICE 1984 recovery plan ecosystems in north american deserts with a list for gila and yaqui topminnowtoptopmmnowminnow poeciliopsispoectltopsis occi of vanishing fishes of the region journal of the dendentalsdentalisdentahstalistails baird and girard U S fish and wildlife arizona nevada academy of science 20 1 62 selvleeserviceSeivice region 2 albuquerque new mexico 67 appp US FISH AND WILDLIFE SERVICE 1989 title 50 received 26 july 1990 wildlife and fisheries part 17 endangered and accepted 6 september 1990 great basin naturalist 503 1990 appp 273 281

DAMSITEDAM SITE SELECTION BY BEAVERS IN AN EASTERN OREGON BASIN

william C mccombmccomb1mccombb james R sedellsedellasedell2 and todd D buchholzbuchholz1buchholzz1

ABSTRACT we compared physical and vegetative habitat characteristics at 14 dam sites occupied by beaver castor canadensis with those at 41 random unoccupied reaches to identify features important to dam site selection in the long creek basin grant county oregon stream reaches with dams were shallower and had a lower gradient than unoccupied reaches beaver did not build dams at sites with a rock substrate bank slopes at occupied reaches were not as steep as those at unoccupied reaches and occupied stream reaches had greater tree canopy cover especially of thinleafthintwinleafleaf alder alnus tenuifolia than did unused reaches A discriminant model using transformations of bank slope stream gradient and hardwood cover classified all beaver dam sites correctly and 35 of 41 random sites as unoccupied sites the 6 misclassified sites had rock substrates we also tested four habitat suitability models for beaver in this basin three models produced significantly different P 05 scores between occupied and random unoccupied reaches suggesting that they might have some utility for this region beaver castor canadensis have long been forniaformamorma used multivariate techniques to iden- recognized as having a significant effect on tify habitat features associated with beaver riparian ecosystems through alteration of occupied reaches slough and sadlier 1977 stream flow they impact soil moisture developed a land capability classification sys- biomass distribution soil redox potential tem for beaver in british columbia based on ph and plant available nitrogen in riparian regression relationships however no mod- areas naiman et al 1988 creation of pool els have been developed for beaver in andaridarld habitat is important to some salmonsalmonidsids cardgardcardgard habitats and none ofthe existing models have 1961 and other pool inhabiting animals par- been tested on independent data from andaridarld ticularlyticularly in areas lacking pools formed by nat- habitats urally occurring coarse woody debris pool our objectives were 1 to locate all beaver habitats can be particularly important for dams in a third order basin representative some species in andaridarld regions where water lev- of andaridarld habitat in eastern oregon 2 to iden- els decrease substantially during the summer tify habitat features potentially important to As central place foragersforagers beaver also create beaver 3 to develop a habitat classification early seralserai stage patches that add to habitat model for beaver in the basin and 4 to test complexity and may influence the diversity of four existing habitat classification models terrestrial organisms naiman et al 1988 al- beaver management represents a low cost STUDY AREA ternativeternative to intensive riparian rehabilitation activities such as cabling coarse woody debris the long creek basin drains approxi- in streams but its success depends on the mately 490 kmkm22 of grant county oregon ability of land managers to predict where fig 1 elevations range from 760 to 1900 m beaver are likely to build dams and thus average annual precipitation is 30 35 cm create pools with most of that occurring in the winter not all portions of all streams are suit- temperatures range from about 10 to 30 C able beaver habitat alienallenailen 1983 developed franklin and dyrness 1973 a habitat suitability index HSI model for the area is dominated by shrub steppe evaluating lacustrine riverine and palu- vegetation typical of andaridanidannd eastern oregon in strine habitats for beaver A similar model was the blue mountains physiographic region developed by urich et al 1984 in missouri franklin and dyrness 1973 sagebrush howard and larson 1985 in massachusetts artemisia sppapp dominates with junipersjunijuniperuspers and beier and barrett 1987 in northern cali Yunijuniperusperus sppapp and ponderosa pine pinus

department of forest science oregon state university corvallis oregon 97331 USA 2usdaSDA forest service pacific northwest research station forestry sciences laboratory 3200 SWS W jefferson corvallis oregon 97331 USA

273 274 W C mccomb ETALET AL volume 50

U U U

U U U U U U U b U b U

U U b

U U U U b dhadaa101101 U U U

U U U UU U b U

OREGON

b BEAVER DAM SITE u unoccupied SITE

fig 1 location of long creek basin grant county oregon and distribution of beaver dams b and random unoccupied reaches u in the basin ponderosa occurring in the higher eleva- METHODS tions riparian vegetation is primarily thinleafthintwinleafleaf on 2 september 1988 we examined 98 km alder alnus willow salix sppapp tenuifolia of perennial streams in the long creek basin hawthorn crataegus sppapp and cottonwood from the air at an altitude of 200 300 m this populus trichotrichocarpacarpa the dominant land included 48 km of long creek 21 km of pass use is grazing and the land is privately owned creek 11 km of pine creek 15 km of basin except for the portion ofthe upper basin in the creek and 3 km of unnamed streams thirty ochoco national forest sites showing signs of possible beaver activity 1990 DAMSITEDAM SITE SELECTION BY BEAVERS 275

ponds pools or felled trees were marked on TABLE 1 variables measured at 14 beaver dam sites a topographic map and then visited on the and 41 unoccupied random stream reaches in the long creek basin giantglantgrant county oregon 1988198819891989 ground fourteen of the possible beaver sites were actually occupied by beaver the others variable method either natural pools were or human induced stream gradient average ofgradientofgradient upstream disturbances or structures in september and downstream from dam or at 1988 we recorded habitat characteristics at a random point on unoccupied the occupied sites and at 16 randomly selected reaches measured with a clinometer unoccupied reaches random reaches were stream width m high width immediately selected by drawing random to iden- water numbers below dam or random point tify points that corresponded to distances in stream depth cm high water depth immediately meters from the mouths of the streams these below dam or random point reaches happened to be skewed toward the Floodfloodplamfloodplainplamplainpiam width m width of area dominated by lower basin so an additional 25 randomly se- alluvial soils at the dam or lected unoccupied reaches were visited in random site march 1989 to obtain a better representation bank slope average ofbankorbankofbank entrance angle of riparian habitat available throughout the on both sides ofthe stream mea- a clinometer basin resulting in a total of 41 unoccupied sured with reaches bank type classified as predominantly dirt or small cobble 20 cm diame- twenty two habitat characteristics includ- ter cobble 21 cmem diameter ing those used in previous studies existing or solid rock models and some that were potentially im- distances m distance to nearest road build- portant in this basin were measured at each ing or bridge dam site n 14 and each unoccupied reach drainage area kkm2m2ma area drained above a dam or anznzn 41 table 1 stream variables were mea- random point sured immediately below the dam at occupied plant cover ocular estimates averaged over or at the randomly selected two 40 m diameter plots see sites point on text for grasses and sedges unoccupied reaches terrestrial habitat was forbs thinleafalderthinleafthintwinleafleaf alderaider willow measured at two 40 m diameter plots per site hawthorn cottonwood juniper plots were established on both sides of the and other conifers mostly pon- stream and were immediately adjacent to the derosa pine dam at occupied sites or to the streamstreambankbank at hardwood the sum ofalderoffalderof alderaider willow hawthorn and cottonwood unoccupied reaches values for the two plots covers were averaged to characterize each site hall shrub the cover of all stems 1 I1 cm 1970 found that 90 of woody food was cut diameter within 30 m of the stream edge and johnston total canopy the sum ofhardwoodofhardwood and conifer and naiman in press reported that most for- covers aging occurred within approximately 35 m of grazing pressure classed as low 25 stems the stream therefore we assumed that 40 eaten medium 25 50 stems m diameter plots adequately sampled eaten high 50 75 stems terres- eaten or very high 75 stems trial habitat for beaver additional variables eaten were measured to characterize dam sites dam height cm pond surface area mam2 average basal diameters of woody stems by species retained for subsequent analysis continuous cut by beaver and percentage of available variables were examined for normality using woody stems by species that had been cut by the W statistic SAS institute inc 1982 580 beaver nonformalNonnonnormalnormormalNorm ormalormainormal data were subjected to square univariate comparisons were made be- root or logarithmic transformations to address tween occupied and unoccupied reaches assumptions behind parametric analysis any with a t test linear correlation between all variables either raw or transformed with combinations of pairs of variables was con- W 070.70 7 max 101.0io1 0 were excluded from ducted for pairs with r 8080.80 only the vari- multivariate analyses based on these criteria able that seemed most biologically meaning- 10 oftleoftbeof the original 20 continuous variables were ful to beaver dam building in this basin was retained for analysis the subset of these 276 W C MCCOMBmccombetalETAL volume 50

TABLE 2 average SE habitat characteristics measured at beaver dam sites and unoccupied reaches long creek basin grant county oregon 1988 1989

habitat occupied unoccupied characteristic transformation W n 14 n 41 Pptt stream gradient log 840 2302230.22 30 2 64056.4056 40 5 0001 stream width m log 936 3908390.83 90 8 3303330.33 30 3 5386 stream depth cm none 899 5406540.65 40 6 134111341.113 41 1 0001 floodplainFlood plain width m log 947 135241352.413 52 4 1201412.01412 01 4 5894 bank slope sqrt 939 111261112.611 12 6 241252419524 12 5 0009 road distance m none 753 539113539 113 65468654 68 3880 house distance m none 490 84384843 84 93624936 24 2995 bridge distance m none 473 86476864 76 90340903 40 6332 drainage area km2 sqrt 921 19223192 23 17022170 22 5864 plant cover grass none 952 546505465.054 65 0 424344243.442 43 4 0669 forb sqrt 894 168231682.316 82 3 83j68 31 6 0067 thinleafthmleaftwinleafThinleaf alder none 631 112371123.711 23 7 3008300.83 00 8 0452 willow none 280 4336433.64 33 6 0903090.30 90 3 3579 hawthorn none 423 5522552.25 52 2 11071 10 7 0836 cottonwood none 281 0000000.00oooo00 0 00000.000oooo0 00 0 1000 hardwood log 904 210712107.121 07 1 5013501.35 01 3 0001 shrub none 650 231672316.723 16 7 63l5633563156 31 5 0313 juniper none 198 3636363.63 63 6 0502050.20 50 2 4040 other conifer none 385 2121212.12 12 1 14041 40 4 7298 total canopy none 630 267802678.026 78 0 6 91 5 0001

10 variables best able to separate occupied from per 7 km of stream but the distribution of the unoccupied reaches was selected by stepwise dams was highly clumped in the middle of the discriminant analysis A classificatory model for basin fig 1 in a study covering a compara- the original data set was developed from this ble area 600 kmkm2 and stream length 153 km subset with canonical discriminant analysis beier and barrett 1987 recorded 43 active discrete data are reported as frequencies beaver colonies in a sierra nevada basin with chi square goodness of fit analyses con- other investigators have reported beaver ducted for among class comparisons values dam densities of one per 010.10oi 1 363.6 km beier from random unoccupied reaches were used and barrett 1987 naiman et al 1988 to establish expected frequencies from our data we evaluated the effective- physical habitat relationships ness offour existing habitat suitability models dam heights xE 55 cm and pond surface 1 the US fish and wildlife service US- areas x 167 mm2ma were highly variable CV FWS HSI model allenailenalienalienallen 1983 2 the mis- 62 and 88 respectively beaver dams souri HSI model urich et al 1984 3 occurred exclusively at sites with dirt rather the massachusetts model howard and lar- than bedrock or cobble dominated banks son 1985 and 4 the truckee river model whereas only 37 of the unoccupied reaches beier and barrett 1987 A new version of the had dirt banks because beaver in this basin missouri HSI model includes beaver in the dennedbenned in banks or lodges adjacent to the list of species evaluated scores were re- banks dirt substrates were probably a requi- corded for each occupied and unoccupied site for adequate dens reach and then compared with a t test for each immediately below the dam streams were model we assumed that a significant differ- shallower had a gentler gradient and had a ence P 05os05.05 in scores between occupied and gentler bank slope than at unoccupied reaches unoccupied reaches indicated potential utility table 2 the features of dirt banks with gen- for a model in long creek basin tle slope low stream gradient and relatively shallow water were best met in the middle of RESULTS AND discussion the basin further upstream the gradient was beaver dams were not abundant in the steep downstream the water was deep and basin we found an average of one beaver dam would probably result in volumes that could 1990 DAMSITEDAM SITE SELECTION BY BEAVERS 277

wash out dams during high flows all unoccu- occupancy and previous grazing practices pied reaches downstream from the occupied kindschy 1985 johnston and naiman in sites were dominated by bedrock drainage press many of the preferred food species basin area stream width and floodplainfloodplain may have been eliminated from the area prior width did not differ between occupied and to this study unoccupied reaches table 2 distances to features that might have af- habitat classification fected the likelihood of dam placement such bank slope stream gradient and hardwood as bridges roads or buildings did not differ canopy cover best separated FP 0001.0001 pil- between occupied and unoccupied reaches lai s trace 0620.62 occupied from unoccupied beaver will apparently live in close proximity reaches the model was to humans and human made structures if all response variable habitat requirements are met 37533.753 vbankbank slope 02720.272 logio10910 stream gradient 52395.239 vegetative habitat relationships 1090logic hardwood cover 12731.273 beaver cut exclusively hardwoods at the with zero as a decision level negative val- dam sites use of thinleafthintwinleafleafalder 6 willow ues of the response variable were classified as 9 and hawthorn 3 was nearly in beaver dam sites and positive values were proportion to availability 0 use in propor- classified as random unoccupied reaches low tion to availability however percent cover values for bank slope and stream gradient and of a site by thinleafthintwinleafleaf alder was higher on occu- high values for hardwood cover produced pied than on unoccupied stream reaches negative values the model correctly classi- table 2 because thinleaftwinleafthinleaf alder was a domi- fied all dam sites and 35 of the 41 85 unoc- nant plant along the riparian area hardwood cupied reaches misclassified unoccupied cover and total canopy cover were higher on reaches were dominated by either bedrock or occupied than on unoccupied reaches alder cobble therefore when all sites except those also dominated the shrub category stems 1 with dirt banks were deleted from the data set cm dahdbh thus shrub cover differed between prior to running the model classification was occupied and unoccupied reaches cover by 100 the accuracy of this model in other other potential food and dam construction drainage basins of this size in eastern oregon plants willow and hawthorn was highly vari- is unknown but it seems likely that these able among sites and so did not differ between habitat characteristics would influence beaver occupied and unoccupied reaches nor did dam building elsewhere in the region cover by conifers table 2 cover by forbs was higher on occupied than on unoccupied assessment of existing models reaches probably resulting from the higher the only model that produced scores that water table around dam sites grass cover at did not differ significantly between occupied dam sites did not differ from that at unoccu- and unoccupied reaches was the massachu- pied reaches setts model table 3 this model was de- assuming that this basin is typical of many signed for use in small watersheds 750 ha in eastern oregon then beaver were most in the northeastern united states and in- abundant prior to intensive beaver trapping in cluded variables that did not pertain to condi- the late 1800s and early 1900s followed by tions in eastern oregon soil drainage class grazing of the area finley 1937 kindschy and abandoned field proximity the other 1985 reported that grazing can adversely af- three models produced scores that differed fect beaver habitat in the region by reducing between occupied and unoccupied reaches willow abundance grazing pressure was FP 006006.006 suggesting that they can provide rated as low to none at 64 9 of 14 of beaver an index to beaver habitat quality in this dam sites and 49 20 of 41 of unoccupied basin stream reaches xaXx2 0730.730 73 P 2.2 although beier and barrett 1987 used stream depth we did not detect any association between a classificatory variable in their study and grazing and dam site selection vegetation re- stream gradient to identify beaver occupied sponses may have been obscured by historic and unoccupied reaches in the truckee river cutting patterns of beaver length of pond basin california when we assessed these 278 W C mccombetalMCCOMB ETAL volume 50

tailleTABLE 3 average SE scores for four models tested with data from beaver occupied and random unoccupied reaches long creek basin grant county oregon 1988 1989 occupied unoccupied model n 14 n 41 ptP t massachusetts 0560140560.140 560 14 0520060520.060 520 06 0813 truckee riverb 1440051440.051 440 05 0390140390.140 390 14 0001 missouri HSI originaloriginal0originals0 0670030670.030 670 03 0550020550.020 550 02 0006 modified 0690030690.030 690 03 0540020.540020 540 02 0006 USFWS HSI originabriginaoriginaloriginaid1 0390060390.060 390 06 0200030200.030 200 03 0005 USFWS HSI food 1460231460.231 460 23 0490090490.090 490 09 0001 USFWS HSI water 0500000500.000 500 00 0430020430.020 430 02 002 USFWS HSI modified 0790110790.110 790 11 0290050290.050039005290 05 0001 USFWS HSI food 1460231460.231 460 23 0490090490.090 490 09 0001 USFWS HSI water 1000001000.001 000ooo 00 0780050780.050 780 05 0003 this study 1510121510.121 510 12 1340231340.231 340 23 0001 allaliailhowaldoward and larson 1985 beier and barrett 1987 trichmrichwrichurlurichetalUrichetalet al 1984 lienilendallendalienllen 1983

variables on the long creek basin data model resulted in average life requisite scores stream gradient drove the model the stream for water of 101.0io1 0 on occupied reaches and 0780.780 78 depth variable was not sensitive to conditions on unoccupied reaches table 3 average at long creek beaver habitat suitability in- HSI scores for occupied reaches increased creased with stream depth in the truckee from 0390.39 to 0790.79 as a result of this change river basin but we found an opposite rela- while scores at unoccupied reaches did not tiontionshipship in the long creek basin change appreciably 02000200.2020 to 0290.290 29 we do not the missouri HSI model produced accept- suggest changes in the calculation of the life able results in eastern oregon but weak- requisite value for food and dam construction nesses were apparent for example this material because that score differed signifi- model places high habitat suitability value on cantly between occupied and unoccupied stream sections with steep banks whereas reaches table 3 beaver in the long creek basin selected gentle bank slopes for dam placement reversing assessing site suitability the suitability index scores for this variable williams 1965 indicated that in addition to made the model more sensitive to conditions sufficient food suitable habitat for beavers in the long creek basin A variable describ- requires a channel gradient 15 and stable ing the proximity to cropcroplandslands did not pertain water levels in riverine habitats stream gra- to long creek basin and was eliminated dient is the most significant factor determin- making these alterations however changed ing the suitability ofhabitat for beaver slough the scores of the original model by only and sadlier 1977 gradient was considered 0 02 units an important habitat feature by retzer et al the USFWS HSI model produces life 1956 slough and sadlier 1977 alienallenailen requisite values between 0 and I1 for both food 1983 urich et al 1984 howard and lar- and water the HSI is the minimum of these son 1985 beier and barrett 1987 and two values the water life requisite value is naiman et al 1988 gradients on beaver oc based on suitability indices SI for water level cupiedaupied reaches in the long creek basin fluctuation and stream gradient all sites mea- ranged from 151.5lsis1 5 to 404.04 0 while those on unoc- sured in the long creek basin were classified cupied reaches were as high as 12 exclud- as having moderate fluctuations in water level ing all stream segments with gradients that could have influenced lodge entrances 12 could facilitate identification of suit- SI 050.50 5 so this variable was not sensitive able dam building segments along long to conditions at occupied and unoccupied creek and its tributatributariesries at most sites gradi- reaches stream gradient was a better predic- ents 7 are probably only of marginal value tor of occupied reaches eliminating the vari- retzer et al 1956 however gradient alone able for stream level fluctuation from the is probably not the best indicator of dam site 1990 DAMSITEDAM SITE SELECTION BY BEAVERS 279

U U b BEAVER DAM SITE U u unoccupied SITE U U B MEAN VALUE OF USED sltesSITES U belerBEIER AND BARRETT 1987 UU U

U 0 0rt001 U U U z ujLU a 7

rrCE U U 0 U U U U U U u U U U U U U

b U U U b U b b B U b b U b b b b b 0 o808oa 292.9 505.0 717.1

2 STREAM AREA m

fig 2 relative stream gradient diagram stream gradient relative to stream cross sectional area five random unoccupied reaches fell below 0088 m2crossm2mccrossma cross sectional area values from random unoccupied reaches below the diagonal line were classified as unusable beaver habitat because of stream substrate or food availability see text

suitability the relationship between gradient in the truckee river basin beier and barrett and dam building is influenced by the cross 1987 although stream depth width and sectional area of the stream because small drainage area above the dam were important high gradient streams can be dammed up to features in other studies howard and larson a point but large high gradient streams can- 1985 beier and barrett 1987 the degree not similarly large streams of low gradient to which these variables indicate habitat qual- can be dammed but again only up to a point ity for beaver is largely dependent on the 5 mm2ma cross sectional area on long creek length of stream sampled and the location our data support this concept as does the of sampling in the watershed in first and mean value from active colony sites B fig 2 second order streams these variables must 280 W C mccomb ET AL volume 50 be sufficiently large to provide adequate wa- shrubs or both once a dam is built forb ter for beaver howard and larson 1985 in abundance will probably increase table 2 large streams depth and width have a nega- resulting in improved food quantity and qual- tive association with dam building because ity in the summer jenkins 1981 the force of the water can prevent dam persis- tence during high flows sampling a wide conclusions range of stream sizes resulted in a gaussian distribution of these factors with similar for streams similar to those in the long means for occupied and unoccupied reaches creek basin we suggest that land managers due to the location of beaver dams in the may evaluate the potential for beaver dam central basin but the range of values for establishment using either the alienallenailen 1983 width and depth is narrower for occupied than HSI model modified for eastern oregon con- for unoccupied reaches using relative stream ditions or the beier and barrett 1987 model gradient cross sectional stream area at a the discriminant model that we developed given gradient overcomes this problem provided excellent classification of the origi- substrate type can also be used to further nal data and used habitat features identified refine selection of potential dam sites ap- by other investigators as important to bea- proximately 63 of long creek and its tribu vers but it has at least two weaknesses first tarnestannes passes through substrates of rock or variable transformations obscure direct rela- large cobble that seem to restrict dam con- tiontionshipsships between beaver and the habitat structionst slough and sadlier 1977 reported characteristic the square root or logarithm of that beaver in their study area did not use a variable may not be as meaningful as the lakes with rocky margins original value second the model has not bank slope is another physical feature that been tested on an independent data set seems important to dam site selection urich an alternative to using the alienallenailen 1983 or et al 1984 considered steep banks important beier and barrett 1987 models is to use the to beaver in missouri probably because they following logic based decision tree A stream offer suitable locations for dens along large segment may support beaver 1 if the rela- streams in our study and that of beier and tive stream gradient falls in the domain below barrett 1987 beaver were associated with the diagonal line in figure 2 2 if the stream gentle bank slopes the influence of bank substrate is not rock or cobble and 3 if the slope on habitat suitability may be a locally hardwood cover is 7 if hardwood cover is important variable and should not be univer- 7 then the land manager has the option of sally included in habitat models improving the section of stream habitat by an adequate and accessible supply of food encouraging woody plant growth to increase and dam construction materials must be the volume of pool habitat in a stream by present for establishment of a beaver colony encouraging beaver the land manager should slough and sadlier 1977 on our study area identify reaches with adequate geomorphic sites with 7 hardwood tree cover were characteristics reestablish hardwoods if nec- unlikely to be dam sites based on a 95 confi- essary and minimize trapping ofbeaver until dence interval denney 1952 summarized the population is well established for suit- the food preferences ofbeaver in north amer- able stream sections this approach would be ica and reported that aspen populus tremuteemu more economical than adding logs or similar loides willow cottonwood and alder were instreaminstream structures that could be better used most often selected the food species present elsewhere may be less important in determining habitat quality than are physiographic and hydrologic acknowledgments factors jenkins 1981 alienallenailen 1983 if food is not adequate but the geomorphic features B H smith salmon national forest already described for dam placement are met salmon idaho C dahm department of then the land manager can encourage the biology university of new mexico albu- growth of food and dam construction materi- querquequerque and R J naiman center for als by restricting grazing of the riparian area streamside studies university of washing- by artificial regeneration of the trees and ton seattle reviewed and improved the 1990 DAMSITEDAM SITE SELECTION BY BEAVERS 281 manuscript this research was supported by HOWARD R J AND J S LABSONLARSON 1985 A stream habitat funds provided through the cooperative ex- classification system for beaver journal of wild- oregon life management 49 19 25 tension service college of forestry JENKINS S H 1981 problems progress and prospects in state university this is paper no 2622 ofthe studies of food selection by beavers pages forest research laboratory oregon state 559 579 in J A chapman and D pursley eds university worldwide furbearerforbearerFurbearer conference proceedings vol 1 university of maryland frostburgFrostburg JOHNSTON C A AND R J NAIMAN in press browse se- literature CITED lection by beaver effects on riparian forest composition canadian journal of forest research KINDSCHY R R 1985 response ALLEN A W 1983 habitat suitability index models of red willow to beaver beaver western energy and land use team use in southeastern oregon journal of wildlife management 49 26 28 division of biological services USU S fish and NAIMAN R A wildlife service fort collins colorado 20 appp J C JOHNSTON AND J C KELLEY 1988 alteration of north american streams by beaver belerBEIERBEIEB P AND R H BARRETT 1987 beaver habitat use bioscience 38 753 762 and impact in the truckee river basin california RETZER J L H M SWOPE D REMINGTON AND W H journal of wildlifeofwildlife management 51 794 799 J rutherford 1956 suitability of physical factors DENNEY R N 1952 A summary of north american for beaver management in the rocky mountains beaver colorado game management fish and ofcoloradoofcolorado colorado department ofgameofgame fish department report 28 colorado of division and parks technical bulletin no 2 denver wildlife 14 denver appp 33pp33 appp FINLEY W L 1937 the beaver conserver of soil and SAS INSTITUTE INC 1982 SAS user s guide basics SAS water transactions of the north american institute inc cary north carolina 921 appp wildlife and natural resources conference 2 SLOUGH B G AND R M F S SADLIER 1977 A land 295 297 capability classification system for beaver castor FRANKLIN J F AND C T DYRNESS 1973 natural vegeta- canadensis kuhl canadian journal of zoology tion ofoforegonoregon and washington U S department 55132455.1324 1335 ofagricultureofagriculture general technical report PNW 8 URICH D L J P GRAHAM AND E A GASKINS 1984 US forest service portland oregon 417417ppappp habitat appraisal of private lands in missouri GARD R 1961 effects of beaver on trout in sagehen wildlife society bulletin 12 350 356 creek california journal of wildlife manage- WILLIAMS R M 1965 beaver habitat and management ment 2525221221 242 idaho wildlife review 17 3 7 HALL J G 1970 willow and aspen in the ecology of beaver in sagehen creek california ecology 41 received 12 april 1990 484 494 accepted 23 june 1990 I1 great basin naturalist 503 1990 appp 283 285

SMALL MAMMAL RECORDS FROM DOLPHIN ISLAND THE GREAT SALT LAKE AND OTHER localities IN THE bonneville BASIN UTAH kenneth L cramericramer A lee foote and joseph A chapman 1 collections made during 1985 and 1986 re- of the bonneville basin in addition we feel it sulted in the following notes on reproduction noteworthy to mention a siting of a sagebrush extensions of geographic ranges and speci- vole on the extreme northern newfoundland mens of rare and uncommon small mammals mountains tant6n r13w s17 because of from the bonneville basin in northwestern the isolated nature of this range which is utah collapsible sherman live traps and vic- surrounded by barren salt flats the vole tor snap traps baited with a mixture of rolled observed one afternoon in june 1985 was oats peanut butter chopped raisins and ba- clearly identified by its short tail and very con fat were used for collections exact locali- light pelage ties and dates of capture are reported under little pocket mouse perognathus longi each species description membris gulosusgulosus thirteen specimens were vagrant shrew sorex vagransmagrans vagransvamagransgrans collected in may 1986 from the western edge three individuals were captured in june 1986 of floating island tooelethoele co utah tant2n at twin springs in tooelethoele co a small spring r16w s22 approximately 50 km northeast dominated by saltgrasssaltgrass distichlis spicataspicatespicata ap- of wendover utah near the end of silver proximately 35 km south of wendover utah island mountains the site had fine sandy tst9s r16w one specimen was found in an soil and the dominant shrub was desert milk- insect pitfall trap two additional specimens wort polygala intermontana three speci- were caught 21 march 1986 in the grassy mens were also collected from the north end mountains tant3n r11w s26 in a shallow of the newfoundland mountains also re- narrow dry ravine the female contained six ported there by H egoscue personal com- embryos 8 mm in length these records ex- municationmunication these records confirm durrant s tend the known range of this subspecies 35 km 1952 hypothesized distribution for this sub- to the north previous record durrant 1952 species in the western deserts of utah ibapah utah and substantiate the occurrence of this subspecies in this of the area SMALL MAMMALS OF DOLPHIN ISLAND bonneville basin sagebrush vole lagurus curtatuscurtatus inter- we trapped for two nights in august 1986 medius two females were recorded from when the great salt lake was at a peak level of the grassy mountains near the area in which 4212 feet above sea level the high lake lev- the vagrant shrews were captured one cap- els reduced this island to an area of 25 ha tured 23 february 1986 was lactating and had area calculated based on the 4210 foot con- four placental scars the other was captured tour line A drop in lake level to 4200 feet in september 1985 the latter specimen was expands the island area to 210 ha although prepared and deposited in the department of much of this area is unvegetated mud flats in fisheries and wildlife teaching collection at 750 trap nights on the island tant9n r10w utah state university these records support only dipodomys ordiiordia and P longimembrislongimembris the general distribution of this subspecies in were captured this contrasts markedly with northwestern utah postulated by durrant goldman s 1939 and marshall s 1940 cen- 1952 and establish the occurrence of the suses of the island 50 years ago goldman sagebrush vole in this western central range spent two nights on the island and found a

department of fisheries and wildlife utah state university logan utah 84322521084322 5210

283 284 NOTES volume 50 much more diverse small mammal fauna personal communication don wilson US in only 37 trap nights at that time the great biological survey however our specimens salt lake was at a historic low and the island are not identical to marshallimarmarshallashalli they are slightly was connected to the mainland by a low sand paler and the tails are darker than marmarshallimarshallashalli bar goldman reported capturing deer mice the skulls of all specimens are very similar peromyscus maniculatusmamculatus ground squirrels these comparisons of our specimens with the spermophilus towntownsenditownsendiasendi and both ord s original series collected by goldman were D ordiiordia and chisel toothed D micronsmicropsmicrops confirmed by comparisons to specimens of kangaroo rats also he recorded evidence D 0 marmarshallimarshallashalli at the university of utah of desert woodralwoodrat neotoma lepida coyotes museum of natural history thus we feel caniscams latranslatranolatrans and a carcass of a porcupine that our specimens of D ordiiordia collected on erethizon dorsatumdorsatum goldman 1939 named dolphin island are more closely related to the a new subspecies of chisel toothed kangaroo subspecies D 0 marmarshallimarshallashalli than to the original rat D TO russrusseoluseolus and ord s kangaroo rat subspecies D 0 cineraceuscineraceous described by D 0 cineraceuscineraceous based on specimens he goldman 1939 durrant 1952 earlier ques- captured on the island tioned the validity of subspecific status for we saw sign of runways of S towntownsenditownsendiasendi cineraceuscineraceous noting frequent connection of through dense stands cheatgrassofofcheatgrasscheatgrass although dolphin island with the mainland and a lack of we saw no aboveground activity in august nearby mainland specimens when we visited the island in addition we little pocket mouse perognathus longi saw droppings and weathered nests of neo- membris gulosusgulosus six specimens of the lit- toma but none of them were recent suggest- tle pocket mouse were collected on dolphin ing that there may be no woodratswoodrats left on this island this species has not been recorded island while no live lagomorphslagomorpha were ob- previously from any island in the great salt served on the island two weathered disartic- lake goldman 1939 marshall 1940 bowers ulated skeletons of jackrabbitsjackrabbits lepus sp 1982 few records are available for this spe- weiewere also found but these could have been cies in the bonneville basin durrant 1952 earnedcarried there by raptorsraptores the island almost shippee and egoscue 1968 the nearest from certainly has no peromyscus remaining in kelton utah on the north shore of the lake other west desert areas we would normally trapping on the nearby mainland at higher catch a minimum of 10 15 deer mice for 750 elevations 5500 feet in the hoguphogop moun- trap nights of effort even in very poor habitat tains failed to produce any individuals of this such as the cheatcheatgrassgrass bromus tectoriumtectectorumtorum species this may have been due to the ab- monoculture dominating the island we sence of habitats usually preferred by this caught no specimens of D micronsmicrops and be- species the P longimembrislongimembris specimens col- lieve that the subspecies named for its occur- lected on dolphin island are much darker rence on the island D m russrusseoluseolus is ex- overall than gulosus although still within the tinct range of variation of this subspecies but ap- ord s kangaroo rat dipodomys ordiiordia mar pear identical in skull morphology speci- shalli we captured 11 individuals of this mens examined from dolphin island are de- subspecies five specimens were deposited posited in the national museum of natural in the national museum of natural history history 3 and the university of utah mu- and another five reside in the university of seum of natural history 3 utah museum of natural history the complete isolation of the island from the specimens of ord s kangaroo rat do the mainland for several years probably ex- not appear to fit within the range of variation plains these faunal changes high lake levels for D 0 cineraceuscineraceous the subspecies first de- have inundated formerly choice dune habitats scribed by goldman 1939 as endemic to dol- occupied by the heteromyids that still persist phin island our specimens are much darker on the island it is likely that the island fauna than cineraceuscineraceous paipal titularlyticulticularlyparticularlyarlyariy the tails in addi- has changed repeatedly over the years as a tion all of our specimens have black facial result of lake level fluctuations that alternately markings like the mainland subspecies D 0 isolated it from and connected it with the marmarsmarshalhmarshallimarshallahalhhalbhaibshalli only one specimen from the origi- mainland in the 1900s alone the island has nal series of cineraceuscineraceous has these markings been isolated from and reconnected to the 1990 NOTES 285 mainland on at least three separate occasions ter of utah state university provided vehicles gwynn 1980 this could account for the ap- and other logistical and technical support parent reinvasion of the island by D 0 mar the utah department of wildlife resources shalli and possible swamping of variation granted the necessary collecting permits ad- found in the subspecies cineraceuscineraceous frequent ditditionalional thanks are due to don wilson of the and periodic invasions and subsequent isola- US biological survey for his comparison of tion make dolphin island a very dynamic sys- our specimens of D ordiiordia with the topotypestopotypes tem whose mammalian fauna could change from dolphin island dramatically as often as lake levels fluctuate with varying precipitation patterns major literature CITED changes include the extinction of a unique subspecies D m russrusseoluseolus and the potential BowbowersBOWFRSrRsers M A 1982 insular biogeography of mammals in great great of a new subspecies of little pocket the salt lake basin naturalist 42 creation 589 596 mouse DURRANT S D 1952 mammals of utah taxonomy and whereas marshall 1940 recorded seven distribution university of kansas publications species of mammals on dolphin island when lawrence was connected by a sandbarsandbag to the GOLDMAN E A 1939 nine new mammals from islands inm it narrow great mainland now apparently only two small the salt lake utah journal of mammalogy per- 2035120 351 357 mammal species dipodomys ordiiordia and GWYNN J W 1980 great salt lake a scientific historical ognaognathusthus longimembrislongimembris survive with possi- and economic overview utah geological and bly a third species spermophilus townsendtownsendiitownsendiaii mineral survey utah department of natural res- surviving as well ources bulletin 116 MARSHALL W H 1940 A survey of the mammals of the islands in great salt lake utah journal of acknowledgments mammalogy 2114421 144 159 SHIPPEESHIPPCE E A ANDANDHH J EGOSCUE 1958 additional mam- this study was funded in part by US air mal records from the bonneville basin utah force department of defense contract no journal of mammalogy 39 275 277 f42650 84 c3559 through the utah state received 20 february 1990 university foundation the department of revised 24 may 1990 fisheries and wildlife and the ecology cen accepted 24 may 1990 great basin naturalist 503 1990 p 287

TWO PRONGHORN ANTELOPE FOUND LOCKED TOGETHER DURING THE RUT IN WEST CENTRAL UTAH

david R smithsmithl

american pronghorn antelope antilocapra wood sarcobatus vermiculatus and shad anieanteamericanaricana americana ord 1818 bucks begin scale atriplex confertifoliaconfertifolia one buck had showing signs of entering the rutting period in heart shaped horns 39439.4 cm in length with late august in western utah smith and beale inward curving tips 383.8 cmem apart during the 1980 bucks exhibit territorial dominance fight this buck evidently thrust its horns up- by marking vegetation with subaricularsubaricsubauricularular gland ward on the underside of the second buck s secretions urine and feces and by direct in- neck the horn tips flexed far enough apart to teractionsteractions with other bucks kitchen 1974 allow the second buck s neck to pass through yoakum 1978 this territorial behavior as- the horns were then locked around the sec- sures that the larger healthier males do the ond buck s neck the second buck s neck was majority of the breeding kitchen 1974 of- rubbed raw and heavily scabbed indicating ten fights between bucks result in injury that the two animals may have remained kitchen 1974 on rare occasions these fights locked together for some time before dying have resulted in the deaths of male mule deer geist 1981 and white tailed marchin- deer literature CITED ton and hirth 1984 when their horns have become locked together the occurrence of GEIST V 1981 behavior adaptive strategies in mule prongpronghornhornborn bucks locking together as a result deer pages 157 223 in 0 C wallmo ed mule fighting and black tailed deer of north america univer- of has been documented very few sity of nebraska press lincoln times spencer 1942 reported a case in south KITCHEN D W 1974 social behavior and ecology of the park colorado in which the right horn of prongpronghornhornborn wildlife monograph no 38 96 appp one buck pierced the underside of the marchinton R L AND D H HIRTH 1984 behavior second buck s jaw while its left horn locked pages 129 168 in L K halls ed white tailed deer ecology and behind the second buck s right horn yoakum management stakepoleStakepole books harrisburg pennsylvania personal communication 1988 reported that SMITH A D AND D M BEALE 1980 antelope in utah two bucks with interlocked horns were found some research and observations utah division of dead on the hart mountain national wildlife wildlife resources publication 801380 13 87 appp refuge in oregon during the late 1940s SPENCERSPENCEKCC C 1942 antelope with locked horns journal on 17 september 1986 a rancher from of mammalogy 239223.92 sutherland utah found two dead pronghorn YOAKUM J D 1978 pronghorn pages 103121103 121 in J L bucks locked together pair were found schmidt and D L gilbert eds big game of the north america stackpole books harrisburg approximately 10 km west of sand mountain pennsylvania in juab county utah t14s r6wraw secsee 99. this area is flat saltbrushsaltbrush desert and domi- received 7 march 1990 nant vegetation species include black grease accepted 15june15 iunejune 1990

utahtah division of wildlifeofwildlife resources 1950 W 1120 S delta utah 84624

287 great basin naturalist 503 1990 appp 289 294

microbiology AND WATER CHEMISTRY OF TWO NATURAL SPRINGS IMPACTED BY GRAZING IN SOUTH CENTRAL NEVADA deborah A hallandhailandhallahall1 and penny S amylamy this study was initiated to monitor the springs with duplicate samples taken from water chemistry and microbial populations at one site each month on a rotational basis two sites in southern nevada ash springs and weather conditions water levels and flow condor canyon cattle impact was suspected were monitored by ocular estimation water to be a causative factor in increased mortality and air temperatures were taken using a cali- of two endangered fish species white river brated thermometer conductivity corning springfish Creniccrenicthysthys baileyibaileyy bailebaileyibaileyyyi in ash model PS 17 ph hanna instruments springs and big spring spinspindacedace lepidomedaLepidomeda model 0624000624 00 and dissolved oxygen hach mollispinismollispinis prapratensistensis in condor canyon were tested in the field using calibrated condor canyon located at the northern equipment water was collected in sterile end of the meadow valley wash in south cen- nalgene bottles by hand dipping the bottle tral nevada is approximately four miles long rinsing and refilling without sampler related and contains a stream system that runs alter- contamination nately through bureau of land management total bacterial counts were evaluated by and private land site I1 is furthest down- dilution and spread plating on r2araa agar stream near the bottom of the canyon site 2 is diacodifco as well as by membrane filtration gel- within the canyon and site 3 is near the man gn6gna during the winter months when mouth of the canyon adjacent to the spring counts were low plates were incubated at source site 4 is closest to another spring room temperature for five to seven days source on delmuesdeamues s ranch a private ranch total coliformscoliforms were measured using the where cattle currently graze cattle are also most probable number method MPN occasionally present near site 3 american public health association 1985 ash springs in pahranagat valley is a and membrane filtration followed by growth warm water spring with temperatures vary- on mendo agar difeo at 37 C fecal coli ing from 33 C to 35 C between summer of forms were also cultured after membrane fil- 1986 and spring of 1987 cattle were present tration and support on mfcmac agar diacodifco at at the headpoolheadpool and there was a marked de- 44 C each coliform colony and positive MPN cline in springfish and other endemic species tube was confirmed by inoculation into bril- removal of the cattle by fencing initiated by liant green bile broth difcodiaco tubes and each the BLM in 1987 allowed fish numbers to was scored positive when gas and acid were increase to the same levels as prior to the produced decline taylor et al 1990 because of the pseudomonas aeruginosa and Aeaeromonasromonas recovery this area served somewhat as a con- hydrophilahydrophilyhydrophila were also evaluated by membrane trol but residual manure continues to influ- filtration bacteria on the filters were grown ence the spring whenever precipitation oc- on mpa agar american public health associ- curs the headheadpoolpool is still utilized by the ation 1985 at 41 C for isolation of P aerugiaerugo public as a hot tub nosa positive colonies were confirmed by collection trips were planned for both streaking on skim milk agar with clearing of spring sites on a monthly basis beginning in the milk by colonies macconkey agar diacodifco september 1988 and continuing until august was originally used for enumeration of A hy 1989 we gathered water samples from four drophila but it did not clearly select for that sites in condor canyon and two sites at ash organism A comparison of several selective

department of biologicalofbiological sciences university of nevadaofnevada las vegas 4505 maryland Parkpafkparkwayvaymay las vegas nevada 89154

289 290 NOTES volume 50

WATER temperature

40U SITE 5535 H D B a 0 0011 gb el 2 5030 A a5aaA 3 A a4aaA 4 25 el 5 0 A- 0 dad5 A A A A 0 0 6 Q- 20 A A A E AAA Q ft u 15 A 8 A A 0 10 0 x A aoeare e A 0 5 0

i i 7 1 0n t- t 9 I1 I1 S 0 N D J F M A M J J A MONTH

fig 1 water temperatures in ash springs and condor canyon sites 1 4 were in condor canyon and sites 5 and 6 at ash springs

AMMONIA LEVELS media for enumeration of A hydrophilahydrophilyhydrophila by arcos al 1988 found ma 55 site et agar rippey and A 0 01 cabelli 1979 to be the most effective there- A 2 45 A A 3 fore we replaced the macconkey agar with A A 4 ma agar beginning with the november sam- 6 5 35 pling incubation was carried out at 37 C and A suspected positive colonies by a yel- 0 denoted E 25 0 6 0 low E color were inoculated into AH semisoft A 0 A A agar A tubes kaper et al 1979 A positive reac- 0 0 tion was scored in tubes exhibiting alkaline 0 C conditions at the top and acid production at 54454 4 1 1 S 0 N D J F M A M J J A the butt of the tube confirmed organisms mont h were motile produced indole and did not 55 site 0 05 produce hydrogen sulfide E 45 06 water temperatures in condor canyon exhibited a gradient with higher temperatures 35 at sites 3 and 4 near the source of the springs and cooler temperatures within the canyon 0 0 E 25 0 temperatures in the canyon dropped during E a 0 the winter months and increased again during 8 a 15 to a the summer while in ash springs they re- 0 0 0 80 mained stable throughout the year fig 1 5 ammonia levels in canyon S 0 N D J F M A M J J A the condor month fluctuated throughout the year but a general increase was observed during december and fig 2 ammonia levels in parts per billion top frame january along with a less significant increase depicts condor canyon bottom frame ash springs in june fig 2 sites 5 and 6 at ash springs exhibited a similar pattern during these months 1990 NOTES 291

when the nitrate plus nitrite levels were NITRATE PLUS NITRITE LEVELS graphed together a similar to the 500 pattern nhanh3 A SITE data was seen but there was a lag of A 0 0 1 time A I21 400 A A 3 nearly a month in the peaks for both condor A A 4 A A A canyon and ash springs fig 3 there was a 300yoo A 0 A A nitrate plus nitrite gradient in the condor 0 A j canyon sites with site 4 at the top of the 200 A A A canyon being highest and site 1 at the bottom A loo100 0 0 lowest 0 0 0 A large peak in organic phosphorus OP 0 Q Qr 0 0 s 0 N D J F M A M J J A levels during march correlated with nhanh3 peaks observed during that month ash canyon 375 springs and condor both exhibited SITE this 4 ada a 5 phenomenon fig 500300 0dM N 6 total viable counts in condorincondor canyon were dramatically similar to OP levels during 225 4 2 march fig 5 the peak at site 2 in january A ajavja 150 bla was notable at ash springs total counts lit varied little A slight peak was observed in 75 march but this may not be statistically valid pseudomonas an 0 aeruginosa opportunistic S 60 N D J F M A M J J A fish pathogen was found on a regular basis month only at site 5 in ash springs with rare colonies fig 3 nitrate plus nitrite levels in parts per billion 6 A appearing at site pattern similar to that top frame depicts condor canyon bottom frame ash exhibited in the total viable counts can be springs seen with a dramatic peak in march and also with higher numbers during the summer and early fall fig 6

ORGANIC phosphorus LEVELS 70 A SITE 60 0 0 1 0 2 50 zA Aaa3 A Aa54 0no 1 5 nQ 404-40 el dad5 0 0 N E 6 V Q 30 0 zlal 0 0 QU 20

10 E el

0 S 0 N D J FE M A M J J A MONTH

fig4figfiga 4 organic phosphorus levels in parts per billion sites 1 4 were in condor canyon sites 5 and 6 at ash springs 292 NOTES volume 50

TOTAL VIABLE COUNT 1500015 000ooo

1250012.50012 500 0 SITE 0 0 0 1 E 1000010 0004- A 2 0 A A a3aaA 3 0 A A a4aaA 4 775007.500500 A 0 dad50 5 0 0 0 6 5000-5 000ooo 16 0 9 A 0 A A A 2 X 2500500 A A A D 0 0 8 0000oooo0 000ooo 00 11

25002 500 S 0 N D J F M A M J J A MONTH

fig 5 total viable counts in colony forming units per 100 ml sites 1 4 were in condor canyon sites 5 and 6 at ash springs total Pseudopseudomonasmonos oeruginoso counts viable counts fig 5 the peak at site 2 35- in january appeared also 50500 the increase of ammonia levels in condor canyon 0 during december january and june S 25 correlated with precipitation and cattle pres- I1 20 N ence in the canyon rain and snow were abundant in and 15 december january with the additional summer showers typical of the mojave 10 desert occurring in june cattle were present

5 1 at site 3 in december and the streamstreambanksbanks 1 Y I were trampled in january when cattle were 0 S 0 N D J t M A M J J A present they deposited nhanh3NH in the form of month urine and feces and runoffduring winter rain storms washed additional nitrogen into the fig 6 pseudomonas aeruginosaaerugmosa counts in colony system from residual feces because nitrogen forming units per 100 ml at site 5 in ash springs is often a limiting nutrient in ecosystems monitoring nitrogen containing chemical spe- Aeaeromonasromonas hydrophilahydrophilyhydrophila another oppor- cies is important the peaks in nitrogen spe- tunistictunistic fish pathogen was found at all sites cies at sites 5 and 6 ash springs in december colony forming units CFU in ash springs and june were probably due to precipitation followed the pattern seen with pseudomonas rather than cattle presence fig 2 A peak aeruginosaaerugmosa levels showing peaks in march was also noted there during march it was and july fig 7 in condor canyon a slight raining during this sampling trip peak was also observed in march with in- the lag time observed between the higher creased numbers during the summer months ammonia and nitrite plus nitrate levels was when total microbial counts were also higher mostmost likely due to the oxidation of ammonia the shape of the graph of total coliform through nitrite to nitrate fig 3 this most levels in condor canyon fig 8 is similar to often occurs as a metabolic process of nitrify the graph of total viable bacterial CFU fig ing bacteria and this conversion can become 5 fecal coliformscoliforms were elevated in march slower in cool temperatures and when low and figure 9 markedly resembles the graph of numbers of nitrifyingnitrifying bacteria are present 1990 NOTES 293

ALFOAEROMONASollttrnhydrophilyhydrophilaHYDRO tilamilaPHILA COUNTS TOTAL COLIFORMS 25 6000 r SITE SITE

1 0 0 0 010 1 10 2 5000 20 0 E 0 022 A 3 0 A 6 3 E 4 0 a3aa A A A a4aa 0 4000 15 A 3000 A 10 2000

5 1000- A 0 A A A

I1 te 0 1 0 t N D J F M A M J J A S 0 N D J F M A M J J A MONTH 25 500 D ED 5 SITE 0 066 0 05D 5 20 E 400 E 0 6 0 0 tet 15 300- 0 to 200- 0 5 100 0 0 0 A 0 0 n a a B ii 0 0 n N D F M A M A j D F M M J J J S N BJJ A J J A 0 MONTH MONTH

fig 7 aeromonasAeromonas hydrophilahydrophilyhydrophila counts in colony form fig 8 total coliform counts in colony forming units ing units per 100 ml top frame depicts condor canyon per 100 ml top frame depicts condor canyon and bot- and bottom frame depicts ash springs tom frame depicts ash springs

the elevated level of total viable bacteria at ash springs the numbers of coliformscoliforms re- during march was probably due to the flected precipitation rather than cattle pres- availability of OP and other nutrients washed ence as exhibited by the peaks in march and in by the rain from both feces and any fertiliz- july when precipitation occurred there had ers used by the nearby rancher figs 5 6 at been no cattle access to ash springs for six other times bacterial growth was probably months prior to the beginning of this study limited by lack of phosphate the higher but the effect was still seen during times of counts in summer and early fall are likely due sufficient precipitation increased fecal colicolt to the warmer water temperatures in condor form numbers were similarly observed during canyon at ash springs where water tem- the rainy season and when water temperature peraperatureture remained stable total counts varied was warmest in condor canyon fig 9 little pseudomonas aeruginosa levels fol- As expected prior to this study bacterial lowed a similar pattern although they were levels were influenced by water tempera- found only in ash springs tures with higher counts correlating with figure 7 depicts aeromonasAeromonas hydrophilahydrophilyhydrophila warmer water in condor canyon increased data beginning with november and extending numbers reflected these changes during the through august only because of the change in warmer months while in ash springs they the isolation medium explained above remained fairly stable throughout the year on the total coliform graph fig 8 the bacterial levels also reflected increased pre- peaks from february through march in con- cipitation and cattle presence because of dor canyon again probably reflect precipita- the influx of nutrients necessary for growth tion the significantly higher numbers at site of microorganisms influence of cattle could 4 during the warmer months correlate with be seen months after their physical presence the time periods when cattle were most when precipitation allowed an influx of nitro- prevalent and water was slowest and warmest gen and phosphorus 294 NOTES volume 50 acknowledgments FECAL COLIFORMS 400 SITE we greatly appreciate the help ofhermiof hermi 0 0 1 2 D hiatt in sampling and microbiological test- 300 A 3 A A ing thanks also go to john unrue for the E 4 dr purchase of a piece of equipment critical to 200 L this study this research was funded by the U S of 0 US bureau land management under D contract uvo53uv053 rog 11 to the university of loo100 A 0 nevada las vegas A A A axaAYA x0 1 2 6 I1 01- t v esa I1 S 0 N D i F M A M J J A literature CITED MONTH 400 arcosABCOS M L A DE VlVICENTECENTE M A MORINIGOMOKINIGO SITE P ROMERO 0 050 5 AND J J BORREGOBORBEGO 1988 evaluation of several 10 01 6 E selective media for recovery of Aeofaeromonasaeromonasromonas hy a 300 ophila from polluted 0 dr waters applied and envi- ronronmentalmental microbiology 5454278654.278627869786 2792 AMERICAN PUBLIC HEALTH association 1985 standard 200 methods for the examination of water and waste water 16th ed american public health association washington 100 0 DC KAPER J R J SEIDLER H LOCKMAN AND R R COLWELL E 0 1979 medium for the presumptive identification 0 B 8 B of aeromonasAeromonas hydrophilahydrophilyhydrophila and enterobacteri S 0 BIBBIBBN 0 J F M A M J J A acealaceae applied and environmental microbiology MONTH 38102338.1023 1026 RIPPEY S R AND V J CABELLICABELLIL 1979 membrane filter fig 9 fecal coliform counts in colony forming units procedure for enumeration of Aeaeromonasromonas hy per 100 mlmi top frame depicts condor canyon and bot- ophiladrdrophila in fresh waters applied and environ- tom frame depicts ash springs mental microbiology 3810838.10838 108 113 TAYLORTAYLOB F R L P GILLMAN AND J W PEDRETTIPEDKETTI 1989 impact ofeattleofeattlecattleeattie on two isolated fish populations in pahranagat valley nevada great basin naturalist 4949149 491 495 received 5 february 1990 revised 22 august 1990 accepted 9 september 1990 great basin naturalist 503 1990 appp 295298295 298 BIRDS OF A SHADSCALE ARTRIPLEX confertifoliaCONFERTI FOLIA HABITAT IN EAST CENTRAL NEVADA dean E medinmedini despite widespread distribution of shad hopsagehousage grabiagrayia spinosa fourwingFourfousrwingfourlingwing saltbush scale atriplex conferticonfertifoliafolia habitat in the atriplex canescentcanescenscanescens black greasewood sar- great basin desert fowler and koch 1982 cobatus vermiculatus and rubber rabbit it has been largely ignored by avian ecologists brush chrysothamnus nauseosus occasion- there are few quantitative assessments of ally occur along shallow washes three breeding bird populations in these vast areas perennial grasses indian ricegrassricegrass oryzopsis used primarily for livestock grazing but see hymenoideshymenoides galleta hilaria jamesiijamejamesiesii and fautin 1946 for western utah smith et al squirreltail sitanionSitanion hystrix occur through- 1984 for southwestern idaho this informa- out the site cheatgrassCheatgrass bromus tectoriumtectorumtectorum tion is basic to understanding the ecology of an annual is a frequent associate plant names desert birds and the stewardship of their habi- follow holmgrenHohngren and reveal 1966 tats in this paper I1 describe breeding bird a densities of shadscaleshadscale community in the METHODS snake valley of east central nevada and com- pare them with other quantitative studies A 20 ha plot was censcensusedcensusesused for breeding from shadscaleshadscale habitats birds using the spot map method international bird census committee 1970 A cen- STUDY AREA sus plot chosen as the best representative of the shadscaleshadscale community was selected by ex- the study area is located 4 km north of amining the vegetation and topography of the baker in southeastern white pine county general area A square plot was surveyed and nevada at a median elevation of approxi- gridded with points numbered and marked mately 1600 inm the study area is a flat valley with stakes at 75 m intervals ten census vis- bottom bounded by foothills and mountains its to the plot were made annually from 29 there are no seeps springs or live streams on march to I1 june from 1981 to 1983 most spot the site although dry washes cross the valley mapping was done from sunrise to early after- floor climatically the area is a cold desert noon when birds were most active different with cold winters and hot dry summers max- census routes through the plot were used imum temperatures in summer frequently ex- with different starting and ending points dis- ceed 35 C and minimum temperatures in tritributed as evenly as practicable among the winter often drop to 2929cC houghton et al visits to ensure complete coverage the plot 1975 annual precipitation ranges from 10 to was censusedcenscensusesused by walking within 50 in of all 20 cm houghton et al 1975 the area is points on the grid recorded bird observa- grazed lightly by cattle trailing to and from tions extended a minimum of 75 m beyond spring fall ranges R jenson personal com- plot boundaries municationmunication at the end of the sampling period clusters vegetation in the study area comprises of observations and coded activity patterns on a mixture of low shrubs with a sparse species maps were circled indicating areas of herbaceous component dominant shrubs are activity or approximate territories fractional shadscaleshadscale green molly kochia americana parts ofboundary territories were determined common winterwinterfatfat eudotiaeurotiaEurotia lanata bud by estimating the portion of each edge cluster sagebrush artemisia spinescentspinescensspinescens and spiny that fell within the study plot oelke 1981

intermountain research station forest service USU S department ofagricultureofagriculture boise idaho83702idaho 83702

295 296 NOTES volume 50

TABLE 1 passerine breeding bird densities individualindividualshasha in shadscaleshadscale vegetation east central nevada 1981 1983

foraging nestingnesting Breedingbreeding bird densitydens ity species categorya subsubstratessubstratebitrate1 1981 1982 1983 horned lark GGO G 128 1521.52 1321.32 efemEremeremophilaerenwphilaophila alpestrisalpestris brewer s sparrow GGI B 0080.08 olo0100.10OJOono 0080.08 spizella breweribrebrewereweri sage thrasher GGI B 0020.02 4 C 005oos0.05 oreoscoptesoreoseoptes montanus total individualindividualshiindividualshashishai 138 1621.62 1451.45 11 biomass ghaahadghadgha11 42 49 44 species richness n 3 2 3 after degraafDeGraadegraafetfetet aat 1985 GGO ground gleaning omnivore GGI ground gleaning insectivore bafterbafferabterabherafter harrisonharrlson 1979 CG ground nester B bush nester c indicates the species was observed infrequently less than three registrations dspeciesspecies weights from dunning 1984

and verner 1985 summarized methodologi- row pooecetes gramineusgraminousgramineus black throated cal and other special problems of the mapping sparrow amphispiza bilineatabilineatebilineata and western method meadowlark sturnella negneglectaneglectslecta total bird biomass was calculated annually horned lark breeding territories were by summing the products of breeding bird contiguous on the study plot from 91 to species densities and average bird species 95 of the total bird density each year was body weights dunning 1984 bird nomen- accounted for by the horned lark table 1 clature is from the 1983 AOU checklistcheck list this species inhabited areas in which the veg- american ornithologists union 1983 etation was open and low growing with considerable bare ground horned larks sang from the ground while perched or from RESULTS AND discussion the air during nuptial flight displays five three passerine bird species bred on the horned lark nests were found during the study site table 1 by far the most common study all were placed on the ground in shal- breeder was the horned lark Eremeremophilaophila low excavations partly beneath or beside a low alpestrisalpestris A permanent resident this broadly shrub or grass tussock incubating females distributed bird occurred throughout the were first observed on 22 april 1983 and study plot less common and in more re- nestnestlingslings were last observed on 19 may 1983 strictstricteded locations were two summer resi- in the great basin horned larks are usu- dents the brewer s sparrow spizella brew ally most abundant in andaridarld valleys but may eri and the sage thrasher oreoscoptes occur in suitable habitat on mountain plateaus montanus or in montane fields ryser 1985 as well as in other species observed as occasional visi- cold northern desert scrub sagebrush and tors on or over the study plot during the subalpine grasslands behle and perry 1975 breeding season included northern harrier brewer s sparrows were a consistent but circus cyaneuscycyaneousaneus red tailed hawk buteoja relatively minor avian component of the shad maicensis ferruginous hawk buteo re- scale community in this study table 1 As a galis golden eagle aquila chrysaetos breeding bird it was largely restricted to scat- american kestrel falco sparveriussparverius prairie tered clumps of black greasewood fourlingfourfourwingwing falcon falco mexicanusmexicanus mourning dove saltbush and rubber rabbitbrushrabbitbrush occurring zenaida macrouramacroura burrowing owl athene near a shallow dry wash that crossed the study cunicularcuniculariaia short eared owl asio flam site I1 found no nests of brewer s sparrow but meus violet green swallow tachycineta observed singing courtship pairing and thalassinathalassinothalassina cliff swallow hirundo pyrrhopyrrhon other breeding activities this sparrow nor- nota barn swallow hirundo rustica com- mally breeds in big sagebrush artemisia tri mon raven corvus boraxcorax loggerhead dentata habitats but will also nest in a variety shrike lanius ludovicianusludovicianus vesper spar of other suitable shrubs short 1984 1990 NOTES 297

TABLE 2 breeding bird densities individualshaindividualsha in shadshadscalescale communities of the great basin desert total location year density species reference southwestern idaho 1979 1541.541 54 3 smith et al 1984 1980 154 3 11 11 11 southwestern utah 1984 1381.38 2 medin 1986 1984 1391.39 2 1984 0980.98ogs 3 1984 1161.16ilg1 16 3 western utah 1940 106log1.06 3 fautin 1946 east central nevada 1981 1381.381 38 3 this study 1982 1621.621 62 2 1983 1451.45 3 identified as the salt desert shrub vegetation type shrub species included shadscaleshad scale bud sagebrush common winterwinterfatfat blackmack greasewood fourfourlingfourwingfourwmgwing saltbushnuttallsaltbush nuttall saltbush atriplexfalcataAtriplex falcata and httleleafhorsebrushlittleleafborsebrush tetradymiatetradymw glabrataglabrateglabrata

I1 recorded relatively low densities of sage to those from shadscalesbadscaleshadscaleseale habitats elsewhere in thrashersThrashers in the shadscaleshadscale community sage the great basin desert overall the number thrashersThrashers bred on the study plot only two of of bird species breeding on a census plot in the three study years table 1 mapped shadscaleshadscale habitats ranged between two and breeding territories included the tallest black three breeding bird densities in shadscaleshadscale greasewood shrubs associated with the dry habitats were relatively uniform between wash that crossed the area sage thrashersThrashers years and locations ranging from 0980.980ogs98 to 11621.6262 were not common on the study plot and no individualindividualshasha but there were pronounced nests were found although considered by compositional differences in the breeding some investigators to be a sagebrush obligate bird communities of several bird species re- ege g braunetalbraun etalet al 1976 sage thrashersThrashers occur ported breeding in shadscaleshadscale habitats only in other plant communities behle and perry the horned lark was common to each census 1975 list the sage thrasher as a regular but plot observed differences in the composition relatively uncommon bird of the great basin of breeding bird communities may have been desert scrub formation that includes shad related to physiognomic and floristic differ- scale black greasewood and rubber rabbit ences in the vegetation at each location brush fautin 1946 classified the sage thrasher as a summer resident in greasewood acknowledgments habitats of western utah few other assessments of breeding bird I1 gratefully acknowledge the field and office densities in shadscaleshadscale habitats are available assistance of J R groves J shochat of the table 2 fautin 19461946287287 reported an aver- humboldt national forest at baker nevada age summer population from actual counts on and W P clary and J W kinney of the 4 ha plots of 106log1.06 birdshabirdbirdshalsha in shadscaleshadscale com- forestry sciences laboratory at boise idaho munimunitiesties of western utah nesting birds in- provided support D A klebenow and cluded horned larks rock wrens sal R E sjogren reviewed an early draft of the pinctespinches obsoletesobsoletus and black throated spar- manuscript rows medin 1986 570 found total densities ranging from 00980.9898 to 1391.391 39 birdshabirdbirdshalsha on several literature CITED sample plots on the desert experimental AMERICAN ornithologists UNION 1983 checklistcheck list of range in southwestern utah breeding birds north american birds ath6th ed american or included horned larks black throated spar- nithologists union washington DC 877 appp BEHLE W AND M PERRY Sh al H L 1975 utah birds check- rows and loggerhead shrikesshaikesrikes smith et list seasonal and ecological occurrence charts and 1984 263 reported a total density of 1 54 guides to bird finding utah museum of natural passerine birdshabirdbirdshalsha in a salt desert shrub com- history university of utah salt lake city 144144ppappp munity in southwestern idaho BRAUN C E M F BAKER R L ENG J S GASHWILER from study breed- AND M H SCHROEDERSCHBOEDER 1976 conservation com- results my three year of mittee report on effects of alteration of sagebrush ing bird populations in a shadscalesbadscaleshadscale habitat in communities on the associated avifauna wilson east central nevada were numerically similar bulletin 8888165165 171 298 NOTES volume 50

DEGRAAF R M N G TILGHMAN AND S H ANDERSON internationalrernatlonal BIRD CENSUS COMMITTEE 1970 an inter- 1985 foraging guilds of north american birds national standard for a mapping method in bird environmental management 9 493 536 census work audubon field notes 24 722 726 MEDIN DUNNING J B JR 1984 body weights of 686 species DIN D E 1986 grazing and passerine breeding birds of north american birds western bird banding in a great basin low shrub desert great basin association monograph 1 3838ppappp naturalist 46 567 572 OELKELKE H 1981 limitations of the mapping method FAUTIN R W 1946 biotic communities of the northern pages 114 118 in C J ralph and J M scott eds desert biome ecological inc shrub in western utah estimating numbers of terrestrial birds cooper monographs 1625116 251 310 ornithological society studies in avian biology FOWLER D AND D KOCH 1982 the great basin pages 6 630 appp 7 63 in G L bender ed reference handbook RYSERSERger F A JR 1985 budsbirds ofoftheodthethe great basin university on the deserts of north america greenwood of nevada press reno 604 appp press westport connecticut 594 appp SHORTORT H L 1984 habitat suitability index models HARRISON H H 1979 A field guide to western birds brewer s sparrow USU S department of the inte- nests houghton mifflin company boston mas- rior fish and wildlife service FWSOBS 82 sachusetts 279 appp 108310 83 16 appp SMITHITH G W N C NYDEGGERNYDEGGEB ANDANDDD L YENSEN 1984 HOLMGREN A H AND L REVEAL 1966 checklist of J passerine the vascular plants of the intermountain region bird densities in shrubsteppeshrubsteppe vegeta- US department of agriculture forest service tion journal offield ornithology 55 261 264 VERNERRNER J 1985 assessment of techniques intermountain forest and range Experexperimenexperimentimenj counting pages 247 302 in R F johnston ed station research paper INT 32 ogden utah m current ornithology vol 2 plenum press new york 160 appp 378 appp HOUGHTON J G C M SAKAMOTO AND R 0 ciffordGIFFORD 1975 nevada s weather and climate nevada bu- received 29 january29january 1990 reau of mines and geology special publication 2 revised 26 july 1990 7878ppappp accepted 8 september 1990 great basin naturalist 503 1990 appp 299 302

california GULL populations NESTING AT GREAT SALT LAKE UTAH don S paul joseph R jehl jr 2 and pamela K yochem2 although the california gull larus califor location of the colonies was determined by nicus is the state bird of utah the history and making an aerial survey of the entire lake status of colonies nesting at great salt lake shore and the islands in 1982 and 1983 a have not been well documented stansbury walk through strip count of active nests and 1852 reported gulls nesting in 1850 and the young was the primary method used to deter- population has been studied sporadically mine colony size in several colonies size was since then behle 1958 22 32 provided a determined by using a spotting scope to count comprehensive review of the history of the nests or adults in several others photo tran- colonies and commented on the reliability of sects were made from the airplane in 1986 early estimates many of which were greatly and 1987 estimates of the adult population exaggerated he made the first complete sur- were made using direct nest counts at small vey in 1931 behle 1958 23 reporting ap- colonies at larger colonies estimates were proximately 80000 adults breeding on the made by comparing aerial photographs to great salt lake islands behle 1958 32 con- those made in earlier years the largest colo- tinued to study the population through the nies were estimated by paul using knowledge 1950s and concluded that from past experience of the colony and its size 1989 numbers the only generalization one can make is that there are in were estimated from aerial population shifts constantly going on and there seems surveys on most islands and by strip transects to be a movement from the remote colonies ofthe lake of mainland colonies ground counts were eastward closer to the foot of the wasatch front and made by paul and jehl at the large colony at closer the food supply is to it is not certainly known the morton salt company and also on gun- whether there has been an actual increase of the total population of gulls for the entire region during late nison island DWR also censcensusedcensusesused colonies at years as some claim it is my feeling that such is not the utah lake and Nepneponsetonset reservoir in 1982 case rather by moving their nesting colonies to new and 1983 for details see paul 1983 1986 locations to the east or at the several refuges the 19871989 seagulls are more conspicuous the results table 1 show that in the 1980s behle 1958 32 provided several examples of gulls nested at 16 20 sites around the lake relocations most notably fig 1 and that at some sites numbers varied hat island which once supported 20000 gulls has been from year to year most of the changes could completely abandoned and the gunnison island popu- be associated with a 10 foot fluctuation in lake lation has been reduced from 60000 to 10000 oior levels which caused the desertion of some 15000 in contrast the rock island utah lake colony colonies and the formation of others the lake increased from a few hundred to 27850 level rose from 4202 in 1982 to 421184211.8 in sporadic records and unequal effort make 1987 and then receded to 420654206.5 in 1989 data from 1932 through 1981 difficult to inter- for example up to 18000 gulls nested at pret to gain a better understanding of the antelope island in the mid 1960s even current situation the utah department of though terrestrial predators badger fox coy- natural resources division of wildlife res- ote had access to the colony paul 1983 and ources DWR repeated behle s work by personal observation no gulls were present conducting aerial and ground censuses of the in 1982 the first year of intensive surveys colonies in 1982 and 1983 paul 1983 addi- perhaps because disturbance at a nearby exca- tional surveys were made in 1986 1987 and vation site discouraged nesting large num- 1989 paul 1986 1987 1989 in all years the bers nearly 33000 returned in 1983 and

utah department ofnaturalofnatural resources 515 east 5300 south ogden utah 84405459984405 4599 2seaaseaea world researchKe search institute 1700 south shores road san diego california 92109

299 300 NOTES volume 50

TABLE 1 numbers of breeding adult california gulls at great salt lake utah 1982 1989 data from utah department of wildlife resources paul 1983 1986 1987 1989 colony 1982 1983 1986 1987 1989 remarks great salt lake 1 salt creek WMA 200 50 50 100 adbndbnd11 controlled population 2 promontory point 1172 0 0 0 0 flooded 1983 3 bear river refuge 2492 4270 0 0 0 flooded after 1983 controlled population 4 perry sewer lagoon 0 0 1200 3165 35 isolated after 1983 connected to mainmalnmain- land 1989 5 GSL mineral 5356 0 0 NA 4500 flooded 1983 6 rocky island 2037 976 20 20 300 reduced in size by flooding inm 1983 again 1986 7 ogden bay a pintail flats 8706 2400 0 0 0 flooded after 1983 b umtlumalunit 1 0 2000 0 0 3592 flooded after 1983 c unit 2 112 0 0 0 0 flooded after 1982 d pasture 0 0 1000010.000 12000 0 occupied as lake rose 8 egg island 3502 1170 0 0 0 reduced in size in 1983 flooded 1984 9 white rock 0 492 200 200 200 area reduced 25 after 1983 10 antelope island 0 32940 34600 34000 0 see text 11 farmington bay 20 250 0 0 0 flooded after 1983 a turpin dike 20 0 0 0 0 flooded in 1983 b interior 0 250 0 0 0 controlled population 12 morton salt co 9476 9660 9500 9500 43025 incorporated antelope island colony in 1989 13 lake point salt co 2740 7072 0 0 0 flooded after 1983 14 hat island 10997 9507 9800 9800 12000 minor reduction in area 15 gunnison island 3032 9450 10000 10000 12700 no change in habitat 16 locomotive springs 0 0 1100 225 200 subtotal GSL 49862 80487 76470 79010790104 76552 other colonies geneva steel utah lake 6591 5982 ND ND ND white lake 8981 7855 ND ND ND Nepneponsetonset reservoir 3680 4856 ND ND ND subtotal others 19252 18693 18000180001 18000 18000180001 grand total 69114 99180 94470 9701097.010 94552 11515 no datidata the colony remained fairly stable until 1989 farther inland and occupied a pasture 12000 when it was abandoned for unknown reasons birds in 1987 which was accessible to mam- concurrently the morton salt colony which malian predators the pasture colony bred had been stable at about 9500 birds from successfully through 1988 but was deserted in 1982 1988 increased to over 43000 pre- 1989 when the dike used in 1985 resurfaced sumably by incorporating the antelope island and was reoccupied birds rising water in the early 1980s also isolated at ogden bay several shoreline nesting the dikes at perry sewage lagoons allowing a areas were inundated by rising water in 1982 colony to form there in 1984 the colony grew 83 paul personal observation causing the to over 3000 in 1987 and then was virtually gulls to move inland in 1984 to a dike separat- abandoned 35 nests in 1989 after its isola- ing waterfowl management units when the tion was destroyed by the falling lake levels dike was inundated in 1985 the gulls moved paul and jehl personal observation 199011990 NOTES 301

there are several other nesting locations in the vicinity of great salt lake of which three are island colony at utah lake the rock 16 estimated at 2000 adults in 1932 by behle grew to 27850 adults in 1942 beckbeckbeek 1942 A subsequent rise in lake level in 1944 made the north am island largely unavailable at that time a new 3 site developed on a dike at the newly estab- 2 lished geneva steel plant beekbeck estimated its 4 size at 6800 gulls in 1946 in 1979 the DWR 6 15 estimated 12320 pairs there 1982 and 1983 causeway 5 counts were 6591 and 5924 adults respec- 6 7 tively white lake a southern extension of utah lake has been active since at least the south am 1960s estimated pairs the DWR 12124 in 14 8 1979 in 1982 and 1983 there were 8981 and 7855 breeding adults respectively 9 in rich county east of the wasatch range gulls have nested at Nepneponsetonset since reservoir 12 at least the 1960s counts in 1982 and 1983 were 3680 and 4856 adults the utah lake and Nepneponsetonset reservoir colonies are still active and currently hold 13 approximately the same numbers as in the early 1980s paul personal observation fig 1 outline map of salt lake numbers refer to locations ofcalifornia gull colonies listed in table 1 discussion man J bich M halpin W kirschke the california gull is a highly adaptable D krueger S manes C marti R mckay species despite a 10 foot fluctuation in lake M miller D peterson M schega level which led to major changes in the T schmidt D shirley and P wood partici- availability of breeding sites and in the size pated inm field studies we thank S I1 bond of individual colonies in the 1980s the num- and D mcdonald for help inm compiling the ber of breeding adults at great salt lake manuscript and M tove and D fischer for has remained essentially constant at about helpful comments 75000 8000080 000ooo birds through that decade and apparently since behle s 1931 survey this is literature CITED surprising in view of the major population BECK D E 1942 life history notes on the california increase this species has undergone in the gull great basin naturalist 3&4 91 108 twentieth century conover 1983 and an ap- behlBEHIBEHLE E W H 1958 the bird life of great salt lake parent increase in winter population at great university ofutah press salt lake city 203 appp salt in recent years tove and fischer CONOVER M R 1983 recent changes in ring billed and lake gull 1988 only apparent anomaly the num- california populations inm the western united the in states wilson bulletin 96 362383362 383 ber of breeding birds is the drop in 1982 PAUL D S 1983 current and historical breeding status which evidently resulted from the temporary of the california gull in the creatgreat salt lake re- abandonment of antelope island estimating gion unpublished report utah department of 18000 birds at nearby colonies gives a total natural resources 71 appp 1986 notes on the 1986 great salt lake california ofofcaca 93000 98000 breeding adults for the gull breedingbleeding population unpublished report great salt lake region utah department of fish and game 1987 1987 great salt lake california gull colony locations and adult breeding population estimates acknowledgments unpublished report utah department of fish and game this study was supported by the los ange- 1989 1989 great salt lake california gull colony les department ofwater and power V bach locations and breeding adult population estimates 302 NOTES volume 50

unpublished report utah department of fish TOVE M H AND D L fischerFISCHEB 1988 recent changes in and game natural resources the status of wintering gull populations in utah STANSBURY H 1852 exploration and survey ofthe valley american birds 42 82 190 of the great salt lake of utah lippincott received 5 february 1990 grambo and co philadelphia 487 appp revised 1 august 1990 accepted 11 august 1990 information FOR AUTHORS the great basin naturalist welcomes previously references IN THE TEXT are cited by author and unpublished manuscripts pertaining to the biologi- date ege g martin 1989 or martin 1989 multiple cal natural history ofwestern north america pref- citations should be separated by commas and listed ce will be to manuscripts ofup to in chronological order use et al after name of erenerencei given concise 1212000000ooo words first author for citations having more than two au- manuscripts SHOULD BE SUBMITTED to james R thors barnes editor great basin naturalist 290 acknowledgments under a centered main MLBM brigham young university provo utah heading include special publication numbers 84602 A cover letter accompanying the manu- when appropriate script must include phone numbers of the author literature CITED also under a centered main submitting the manuscript it must also provide heading lists references alphabetically in the fol- information describing the extent to which data lowing formats text or illustrations have been used in other papers mack G D and L D flake 1980 habitat rela- or books that are published in press submitted or tiontionshipsships of waterfowl broods on south da- soon to be submitted elsewhere authors should kota stock ponds journal of wildlife man- adhere to the following guidelines manuscripts not agement 44 695 700 so prepared may be returned for revision 44695 sousa W P 1985 disturbance and patch dynam- CONSULT THE MOST RECENT ISSUE of this journal ics on rocky intertidal shores pages 101 124 for general style and format also refer to the CBE loi101124 S in T A pickett and P S white eds 1 style manual ath5th edition council of biology edi- I the torslors 9650 rockville pike bethesda MD 20814 ecology of natural disturbance and patch dy- press USA 24 namics academic new york coulson R N and A 1984 TYPE AND DOUBLE SPACE all materials including R J witter forest literature cited table headings and figure legends entomology ecology and management john avoid hyphenated words at righthandright hand margins wiley and sons inc new york 669 appp underline words to be printed in italics use stan- TABLES are double spaced on separate sheets and dard bond 22 X 28 cm leaving 25 cm margins on designed to fit the width of either a single column all sides or a abagepageapage use lowercaselower case letters to indicate foot- SUBMIT 3 COPIES of the manuscript number all notes pages and assemble each copy separately title photocopies OF FIGURES are submitted ini- page abstract and key words text acknowledg- tially with the manuscript editors may suggest ments literature cited appendices tables figure changes lettering on figures should be large legends figures enough to withstand reduction to one or two TITLE PAGE includes an informative title no lon- column width originals must be no larger than ger than 15 words names and addresses ofauthors 22 x 28 cm a running head offewer than 40 letters and spaces NOTES if your manuscript would be more ap- footnotes to indicate change of address and author propriatepropriate as a short communication or note follow to whom correspondence should be addressed if the above but do not include an ab- other than the first author instructions stract ABSTRACT states the purpose methods results A CHARGE of 45 per page is made for artiartlartieresarticleschescreseres and conclusions of the research it is followed by published the rate for will 40 6 12 key words listed in order of decreasing im- subscribers be per portance to be used for indexing page however manuscripts with complex tables andor numerous half tones will be assessed an TEXT has centered main headings printed in all additional charge purchased capital letters second level headings are centered reprints may be at the time of publication an order form is sent with in upper and lowercaselower case letters third level headings begin paragraphs the proofs FINAL CHECK VOUCHER SPECIMENS authors are encouraged to designate properly prepare label and deposit cover letter explains any duplication of infor- high quality voucher specimens and cultures docu- mation and provides phone numbers menting their research in an established perma- 3 copies of the manuscript nent collection and to cite the repository in publi- conformity with instructions cation photocopies of illustrations ISSN 0173614017 3614 GREAT BASIN naturalist volvoi5050.50 nonoano33 october 990wo1 CONTENTS articles sprouting and seedling establishment in plains silver sagebrush artemisia cana pursh sspasp cana C L wambolt T P walton and R S white 204201 bibliography of nevada and utah vegetation description P S bourgeronBourgeron L D engelking J S tuhy andandaandjJ D brotherson 209 conservation status ofthreatened fishes in warner basin oregon jack E williams mark A stern alan V munhall and gary A anderson 243 home range and activity patterns of black tailed jackrabbitjackjaekjackrabbitsrabbitrabbitss graham W smith 249 humpback chub gila cypha in the yampa and green rivers dinosaur national monument with observations on roundtail chub CG robusta and other sympatric fishes catherine A karp and harold M tyus 257 management of endangered sonoran topminnowtopminnow at bylas springs arizona description critique and recommendations paul C marsh and W L minckley 265 dam site selection by beavers in an eastern oregon basin v william C mccomb james R sedell and todd D buchholz 273 notes small mammal records from dolphin island the great salt lake and other localities in the bonneville basin utah kenneth L cramer A lee foote and joseph A chapman 283 two pronghorn antelope found locked together during the rut in west central utah

I1 I1 david R smith 287 microbiology and water chemistry of two natural springs impacted by grazing in south central nevada deborah A hall and penny S amy 289 birds ofa shadscaleshadscale atriplex confertifoliaconfertifolia habitat in east central nevada dean Eemedinmedin 295 california gull populations nesting at great salt lake utah don S paul joseph R jehl jr and pamela K yochem 299 GREAT BASIN naturalist MEMOIRS SERIES

the great basin naturalist memoirs series was established in 1976 for scholarly works in biological natural history of greater length than can be accommodated in the great basin naturalist the memoirs series appears irregularly approval for the production is the respon- sibisibilitylity of the great basin naturalist editorial board information concerning the production of the memoirs series can be obtained from james R barnes editor great basin naturalist 290 MLBM brigham young university provo UT 84602

GREAT BASIN naturalist MEMOIRS

all memoirs are available for purchase direct inquiries to editorial office attnanttn carolyn backman 290 MLBM brigham young university provo UT 84602 only checks or money orders drawn on US banks or US currency are accepted please include 2 per copy for mailing and handling charges no 1 the birds of utah 10 no 2 intermountain biogeography a symposium 15 no 3 the endangered species a symposium 6 no 4 soil plant animal relationships bearing on revegetation and land reclamation in nevada deserts 6 no 5 utah lake monograph 8 no 6 the bark and ambrosia beetles of north and central america coleoptera scolytidae a taxonomic monograph 60 no 7 biology of desert rodents 8 no 8 the black footed ferret 10 no 9 A utah flora 40 no 10 A reclassification of the genera of scolytidae coleoptera 10 no 11 A catalog of scolytidae and platypodidaePlatypod idae coleoptera part 1 bibliography 30 no 12 research in the auchenorrhyncha homoptera a tribute to paul W oman 30